Memoirs of the NATIONAL MUSEUM Of VlCtOria Melbourne Australia 7 July 1978 Ml :te f H Number 39 Front Cover— Squirrel Glider. Back Cover — Plains-wanderer. MEMOIRS of the NATIONAL MUSEUM OF VICTORIA MELBOURNE AUSTRALIA No. 39 Director John McNally Deputy Director Thomas A. Darragh Editor Douglas M. Stone PUBLISHED BY ORDER OF THE COUNCIL 7 JULY 1978 © National Museum of Victoria Council 1978 Printed by Brown Prior Anderson Pty Ltd Burwood Victoria NATIONAL MUSEUM OF VICTORIA Members of Council Sir Robert Blackwood, MCE BEE FIE Aust (President) Professor J. W. Warren, MA PhD (Calif) (Deputy President) James C. F. Wharton, BSc (Treasurer) Professor E. S. Hills, CBE PhD (Lond) Hon DSc (Dunelm) DSc (Melb) FIC FAA FRS Professor Sir Sydney Sunderland, CMG MD BS DSc FRACP FRACS FAA Dr H. K. Worner, CBE DSc (Melb) Hon DSc (Newcastle) ABSM FAA FTS FRACI FIM FIMM MAIMM MAIME Dr H. N. B. Wettenhall, MD BS FRCP FRACP Brigadier P. P. Jackson, CMG BE FIE Aust FIMechE FAIM Henry G. A. Osborne, BAgrSc John McNally, ED MSc Chief Executive Officer I. J. Curry, AASA Secretary to Council Staff (as at 1/5/1978) Director: John McNally, ED MSc AMAA Deputy Director: Thomas A. Darragh, MSc DipEd AMAA Administration: A. G. Parsons (in charge) Debbie Valentine E. Rowley R. Domican Patricia Batchelor Judith Freeman Kayelene Traynor Lynette Anderson Ursula Gawronski Scientific Staff Geology and Palaeontology: Curator of Fossils: T. H. Rich, BA PhD Curator of Minerals: W. D. Birch, BSc (Hons) PhD Assistant Curator of Fossils: Juliet O'Conor, BSc Assistants: Kerryn Gent, Susan Gibson, H. Veitch Zoology: Senior Curator (Zoology) : B. J. Smith, BSc PhD AMAA Curator of Vertebrates: Joan M. Dixon, BSc (Hons) Curator of Birds: A. R. McEvey, B.A. OIC Herpetology: A. J. Coventry Curator of Insects: A. Neboiss, MSc PhD FRES Assistant Curator of Insects: A. A. Calder, BSc (Hons) Curator of Invertebrates: Vacant Assistant Curator of Invertebrates: Suzanne Boyd, BSc Assistants: Elizabeth Matheson, Rhyllis Plant, Joan Phillips, Linda Huxley, BSc, LORENE REID Anthropology: Curator of Anthropology: A. L. West, BA DipSocStud Assistant Curator of Anthropology: Alison M. Oates, BA Research Assistant: Judith Wiseman, BA Assistant: Christine Hogarth, BA Survey: Survey Officer: J. D. Blyth, BSc Assistants: H. G. Bawden, Alena Glaister, BSc Conservation: Conservation Officer: D. S. Wood, BA Service Staff Library: Librarian in Charge: Clara Myers, BA DipLib Librarian: Christine McKenzie, BA DipLib Assistant: Susan Alexander Display and Preparation: Display Officer: A. G. Soliman, BA (Cairo) Assistants: Diane Stephens, Dip Ad, John Challis, DipArt, Sandra Brown, DipArt Senior Preparator: L. Chapman Preparators: M. G. Traynor, B. Hall, K. Kelly Photography: Photographer: F. Coffa, DipArt (Photo Illust.) Education Service: R. N. Miller, DipArt (in charge) A. Perry TPTC Julie Carter, BA (Hons) DipEd Rosemary Taylor, BA TPTC Wendy Proudlock, TPTC Annette Seeman, DipTeaching Valerie McCormack (Typist) Honorary Associates with year of appointment Geology: A. A. Baker, 1951 A. W. Beasley, MSc PhD DTC, 1973 A. C. Collins: FRAIA ARIBA AMTPI MACE, 1953 E. D. Gill, ISM BA BD FGS FRGS, 1973 Professor J. F. Lovering, MSc PhD, 1974 D. J. Taylor, MSc, 1966 J. A. Talent, MSc PhD, 1966 H. E. Wilkinson, BSc, 1970 Vertebrate Zoology: C. N. Austin, 1955 C. W. Brazenor, 1962 A. G. Brown, MRCS (Eng) LRCP (London), 1968 N. J. Favaloro, 1945 A. K. Lee, BSc (WA) MA PhD (Calif), 1972 M. J. Littlejohn, PhD (WA) MSc, 1972 P. A. Rawlinson, BSc, 1968 C. Tanner, 1953 R. M. Warneke, MSc BAgrSc, 1966 Invertebrate Zoology: K. N. Bell, BSc DipEd, 1973 J. Hope Black, MSc, 1966 R. F. Burn, 1962 A. N. Burns, MSc FRES, 1966 D. F. Crosby, FRES AASA AFAIM, 1968 R. L. Jensz, BSc DipEd, 1968 C. McCubbin, 1974 A. E. Monger, LS MIS (Aust), 1974 E. T. Smith, 1960 Jeanette E. Watson, ASMB AMTC, 1970 L. Winsor, DipMedTech FAIMT AAIST, 1974 Anthropology: J. H. McNamara, MB BS FRCPA, 1969 N. M. Wallace, 1970 Sir William Dargie, CBE FRSA, 1977 CONTENTS GEOLOGY 1. Geology and Structural Development of the Cerberean Cauldron, Central Victoria. By W. D. Birch, A. J. W. Gleadow, B. W. Nottle, J. A. Ross and Whately. INVERTEBRATE PALAEONTOLOGY 1 2. Gastropods from the Early Devonian Bell Point Limestone, Cape Liptrap Peninsula, Victoria. By C. B. Tassell. (Plate 1) 19 3. The Omphalocirridae: A new family of Palaeozoic Gastropoda which exhibits sexual dimorphism. By R. M. Linsley. (Plates 2-10) 33 VERTEBRATE PALAEONTOLOGY 4. Propleopus chillagoensis, a new North Queensland species of extinct giant rat-kangaroo (Macropodidae: Potorinae). By M. Archer, A. Bartholomei and L. G. Marshall. (Plate 11) 55 5. Detailed comparisons of the dentitions of extant Hexanchid Sharks and Tertiary Hexanchid Teeth from South Australia and Victoria, Australia. (Selachii:Hexanchidae). By N. R. Kemp. (Plates 12-15) 61 6. Raemeotherium yatkolai gen et. sp. nov., a primitive diprotodontid from the medial Miocene of South Australia. By T. H. Rich, M. Archer and R. H. Tedford. (Plate 16) 85 7. Synonymy of the fossil wombat Vombatus pliocenus (McCoy) with the living species Vombatus hirsutus (Perry). By H. E. Wilkinson. (Plates 17-18) 93 INVERTEBRATE ZOOLOGY 8. Comments on the species concept in some Australian Anisops Spinola (Hemiptera:Notonectidae). By I. Lansbury. 101 9. Revision of the Australian Euasthetinae (Coleoptera: Staphylinidae). By V. Puthz. 117 10. Leptanthura and new related genera (Crustacea, Isopoda, Anthuridea) from Eastern Australia. By G. C. B. Poore 135 11. The Spionidae (Polychaeta) from Southeastern Australia with a revision of the genera of the family. By J. A. Blake and J. D. Kudenov. 171 VERTEBRATE ZOOLOGY 12. Survey of the Vertebrate Fauna in the Grampians-Edenhope area of Southwestern Victoria. By W. B. Emison, J. W. Porter, K. C. Norris and G. J. Apps. (Plates 19-21) 281 Contributions towards the cost of printing this Memoir have been made by — Dr R. M. Linsley (Paper 3); The Tasmanian Museum and Art Gallery (Paper 5); Dr J. A. Blake (Paper 11); Marine Studies Group, Ministry for Conservation (Paper 11); and Fisheries and Wildlife Division, Ministry for Conservation (Paper 12). INSTRUCTIONS TO AUTHORS Two copies of the manuscript, with any accom- panying Plates and Figures, should be sub- mitted initially to the Deputy Director, National Museum of Victoria. Authors should consult a recent number of the Memoirs to acquaint themselves with the format. Manuscripts must be typed on quarto paper, double-spaced, on one side of paper only, and with ample margins. Captions to Text Figures and Explanations of Plates must be attached to the MS as final pages. Underlining should be restricted to generic and specific names of bio- logical taxa. Measurements must be expressed in the metric system (SI units). 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They should then be photographed to reduce them to full page size, 15 5 cm x 21 cm, and submitted as glossy prints. They will be reproduced on metal, not type-set, and hence must be finally correct when submitted, since they cannot be corrected at the proof stage. GEOLOGY AND STRUCTURAL DEVELOPMENT OF THE CERBEREAN CAULDRON, CENTRAL VICTORIA By W. D. Birch,* A. J. W. GLEAr>ow,f B. W. Nottle,! J. A. Rossf and R. WHATELYf * Mineralogy Department, National Museum of Victoria t Geology Department, Melbourne University Abstract The Cerberean Cauldron, in Central Victoria, is a classic example of the cauldron sub- sidence phenomenon. The ring and radial fracture patterns and the interbedded volcanic and sedimentary units preserved enable the history of the cauldron's development to be traced in detail. Three cycles of acid magma represented by rhyolite-rhyodacite ash flows with associated tuffaceous sediments were interspersed with periods of sedimentation or eruptions of basic lavas. The earliest formations, constituting the pre-collapse phase, show variable thickness and distribution within the cauldron, whereas the thick Cerberean Volcanics, representing the major subsidence event overlap all other formations. The rhyolites and rhyodacites respectively from the three acid volcanic formations show similar petrographic features. Igneous activity in the region may have commenced with basining and ash-flow eruptions from along early formed faults or isolated vents. The ring and radial fractures were prob- ably initiated by a point explosion at depth in the chamber providing magma for the Cer- berean Volcanics. Eventual collapse of the central block was one of foundering under gravity, following removal of magmatic support due to substantial ash-flow eruptions. Activity culminated in high level intrusions of granodioritic magma. Introduction Upper Devonian vulcanism in Central Vic- toria is represented by a number of well-ex- posed subsidence complexes, occupied by part or all of a remarkably similar rhyolite-dacite sequence. The total volume of acid-intermedi- ate magma preserved is of the order of several thousand cubic kilometres. (A general outline of the geology of the Central Victorian province is given by Marsden, 1976.) The Cerberean Cauldron is the most spec- tacular complex, being a classical example of cauldron subsidence (in the sense of Smith and Bailey, 1948). Its importance lies in that its three cycles of acid volcanism may reflect rhythmic crustal movements and magmatism in the prevailing tectonic regime. The geometric centre of the Cerberean Caul- dron complex lies 96 km northeast of Mel- bourne. The volcanic rocks within the struc- ture now comprise the Cerberean Ranges, the high-level, roughly elliptical plateau forming part of Victoria's Eastern Highlands. The area was mapped in detail by Birch et al. (1970) and covers an area of about 1 150 square kilometres (Fig. 1). Earlier mapping work on parts of the cauldron had been carried out by Bell (1959), Thomas (1947), Hills (1929, 1932) and Whitelaw (1913). Additional mapping of the Acheron Cauldron, which adjoins the Cerberean along the latter's southern edge, was carried out by Dudley et al (1971) and Howard (1972). Mapping of the basement sediments, in particu- lar the Cathedral and Koala Creek Sandstone, was carried out by Vandenberg (1977). This paper outlines the general geology of the Cerberean cauldron and its structural de- velopment, and provides a brief petrography of the main rock types. The chemistry and pedo- genesis will be discussed in more detail in further papers, although these have been al- luded to by Birch and Gleadow (1974). Tectonic Setting The Central Victorian Trough was appar- ently a localized basin of sedimentation, pos- sibly representing a marginal sea within the Lachlan Geosyncline, at the southern end of the Palaeozoic Tasman Mobile Belt. The Trough is geologically bounded on east and west by complex up-faulted axial structures consisting of Cambrian sediments and green- stones with tholeiitic affinities — -the Mt Wel- lington and Heathcote axes respectively. Within this trough, marine geosynclinal conditions W. D. BIRCH, A. J. W. GLEADOW, B. W. NOTTLE, J. A. ROSS, and R. WHATELY Figure 1— The Cerberean Cauldon, showing its location and the positions of the larger scale maps of the sequence along the margin of the volcanic pile shown in Figs. 4 to 8. CERBEREAN CAULDRON were operative from at least the Lower Ordo- vician, but possibly as early as Lower-Middle Cambrian, up until Lower-Middle Devonian times. In the late Middle to early Upper Devonian, the sedimentary fill was folded on a NNW-SSE trend, and intruded by a dyke swarm with mainly lamprophyric and quartz dioritic rock types (the Woods Point Dyke swarm) whilst orogenic forces were still operative (Hills, 1959). Following deformation, large amounts of acid-intermediate magma were emplaced at high levels within the crust, forming a batho- lith-volcanic province fairly typical of circum- Pacific (Andean) magmatism. General Geology The Cerberean Cauldron is circumscribed by a circular, near vertical, partly dyke-filled ring fracture of 27 km diameter, in combina- tion with a radial fracture pattern focussing at the geometric centre. (See Figs. 1 and 2). The complex has been emplaced through the thick Lower Silurian-Lower or Middle Devonian bedrock of folded sandstones, siltstones and shales. The youngest basement rocks are the Cathedral Range and Koala Creek Sandstones, which outcrop on the western and southeastern sides of the cauldron volcanics respectively, in what appears to have been a northwest to southeast trending structural basin. They are apparently continuous beneath the volcanics (Clarke et al., 1970). Both units form prom- inent strike ridges within and subparallel to the ring fracture, but are apparently conform- able with underlying basement sediments (Dale, 1964; Bell, 1959; Hills, 1929). The units as a whole behaved competently during cauldron formation and have locally influenced the fracture pattern (Fig. 2). The sequence within the Cerberean Caul- dron can be divided into a number of alternat- ing volcanic and sedimentary formations, which have a general inward dip at relatively shallow angles (Fig. 3). These are shown in Table 1 and are based on the subdivision of Thomas (1947) and on more detailed mapping of Birch et al (1970). The five formations within the Taggerty Sub- group represent the pre-collapse phase in the cauldron's development. Wightmans Hill Conglomerate This is a terriginous formation comprised mainly of a high-energy, fluviatile conglomer- ate, with well rounded pebbles of indurated quartz sandstone derived from the Palaeozoic bedrock. The pebbles range up to about 30 cm in diameter, but average 3-8 cm. The formation overlies an irregular base- ment topography and hence outcrop distribu- tion and thickness are variable. In some places, quartzite pebbles on the surface are its only expression. It is thickest in the northeast (30 m) and most continuous in the Blue Range, east of Taggerty (Fig. 4) . Snobs Creek Volcanics Volcanic activity in the cauldron began with the emplacement of rhyolitic ash flow tuffs, of relatively small extent. The formation is best developed west of Snobs Creek in the north- west, where it reaches a maximum exposed thickness of 300 m (Thomas, 1947). Its out- crop elsewhere on the western side of the caul- dron is fairly continuous but it is generally less than 60 m thick (Fig. 4). Near Snobs Creek, the basal ignimbritic rhyolite, containing car- bonized plant remains, grades upwards into a rhyodacite. Overlying this is a band of poorly, or non-welded tuffaceous rocks, with more compact bands of biotite-rich rhyodacite. Tuffs rich in phenocrysts occur at the top of the formation (Thomas, 1947). The formation is restricted to the west or downthrown side of the Snobs Creek Fault apart from a small exposure on the eastern side of the cauldron (Fig. 5). Blue Range Formation This formation consists of sedimentary units of various types, some probably tuffaceous, and represents a definite break in the extrusive ac- tivity. The most significant unit is the 'Fish Beds' (Hills, 1929), which reaches a maximum development (about 130 m thick) in the Blue Range, north of Little River (Fig. 4). Here, well-bedded, red and yellow fine-grained, lacu- strine sandstones, and brown and greenish W. D. BIRCH, A. J. W. GLEADOW, B. W. NOTTLE, J. A. ROSS, and R. WHATFLY TABLE 1 Generalized sequence within the Cerberean Cauldron 5 o o > GO < s xj ft XI Oh 5 o o O O < INTRUSIVE PHASE ACHERON VOLCANICS Donna Buang Hypersthene Rhyodacite Warburton Quartz Rhyodacite CERBEREAN VOLCANICS Lake Mountain Rhyodacite Rubicon Rhyolite CENTRAL INTRUSIONS Granodiorite Porphyritic microgranodiorite RING DYKES, RADIAL DYKES Porphyritic granodiorite (equivalent to the Lake Mountain Rhyodacite). Porphyritic granite (equivalent to the Rubicon Rhyolite). NOT REPRESENTED Ignimbritic rhyodacite (uppermost unit in cauldron). Ignimbritic rhyolite, grading to rhyoda- cite at top. Lenticulite at the base. * 900 m 390 m ROBLEYS SPUR VOLCANICS TORBRECK RANGE ANDESITES BLUE RANGE FORMATION SNOBS CREEK VOLCANICS WIGHTMANS HILL CONGLOMERATE Porphyritic andesite, andesitic basalt, minor basalt in south, some inter- calated with underlying Ignimbritic rhyodacite flows. Interbedded non- welded tuff, tuffaceous sediments. Minor agglomerate. Ignimbritic rhyolite flows, associated tuff. 150 m Andesite basalt, andesite, minor basalt, basic and intermediate tuff, minor agglomerate. Shale and fine sandstone, with fish and plant remains. Some tuffaceous sedi- ments, minor quartzite and hornfels. Phenocryst-rich tuff. Non-welded biotite-rich tuff, bands of welded rhyodacite. Ignimbritic rhyodacite with biotite ap- parently grading into Basal ignimbritic rhyolite. 360m 120 m 285 m Basal quartzose conglomerate, minor quartzite, unconformable with base- ment. 30m * Approximate maximum thickness m metres. CERBEREAN CAULDRON 146° 00 ACHERON CAULDRON 37° 30' Late DEVONIAN Early SILURIAN DEVONIAN Volcanics of Marysville Ingeous Complex Central granodiorite intrusions Ring dyke Cathedral Beds, Koala Creek Beds Sandstone, claystone t sediments asement seaimen J_ Dip -* Fault - — ■ Thickening of volcanics 3K Eruption point Eruption point interpreted from gravity survey Figure 2 — Structural Map of the Cerberean Cauldron. A-B and C-D arc cross section lines for Fig. 3. W. D. BIRCH, A. J. W. GLEADOW, B. W. NOTTLE, J. A. ROSS, and R. WHATELY shales have yielded a fish fauna and plant re- mains of Upper Devonian age. (Hills, 1929, 1931). The formation is not well developed at Snobs Creek and only thin conglomerates are recorded by Thomas (1947). On the eastern side of the Cauldron, the stratigraphic position is occasionally represented by thin, compact brown or greenish shales containing mud pellets. Torbreck Range Andesites This is the most extensive of the formations in the Taggerty subgroup, and is the only known occurrence of basic vulcanism (apart from that of the Robleys Spur Volcanics) in any of the Victorian cauldron structures. The formation consists of uniform, dark fine- grained basaltic andesites with minor basic tuffs of various types and occasional agglo- merates. The greatest variety of rock types occurs at Snobs Creek (Thomas, 1947). The formation reaches its maximum thick- ness and continuity along the eastern side of the cauldron, and the outcrop pattern between the Conn Hill and Newman Plain Faults in- dicates a thickness of at least 380 m (Fig. 5). Robleys Spur Volcanics Two easily recognized and fairly persistent members characterize this formation. The older consists of dark, phenocryst-poor ash flow rhyolites (the Middle Rhyolite of Thomas, 1947) and the younger is of dark, fragmental rhyodacites (the Fragmental Toscanites of Hills, 1932). There is a wide variety of other rock types in the formation, including tuffs and tuffaceous sediments, particularly on the west- ern side of the cauldron. On the eastern side, the formation is generally less than 20 m thick and the rhyolitic member predominates. In the south of the Cerberean Cauldron, close to its junction with the Acheron Caul- dron, porphyritic andesites are interbedded with acid volcanics, e.g. in the cuttings of the Marysville-Cumberland Road up Robleys Spur and near Cumberland in the southeast of the cauldron (Fig. 6). Cerberean Volcanics This formation represents the catastrophic collapse phase of the central block and con- tributes about 80% of the volcanic rocks in the cauldron. The formation has been sub- divided into two mappable units but these have no definite structural boundary and represent almost continuous ash flow emplacement (Birch, in prep.). (1) Rubicon Rhyolite. The Rubicon Rhyolite (Nevadite of Hills, 1932 and Thomas, 1947) is a thick, phenocryst-rich, rhyolitic ash flow averaging 250 m in thickness and with an areal extent of about 400 km 2 in the Cerberean Cauldron. It is thicker on the western side (up to 390 m near Little River, Fig. 4), than on the east, where it may be absent (Fig. 5). In the north, near Snobs Creek, the Rubicon Rhyolite forms a prominent cliff-forming hori- zon. A lenticulite horizon occurs at the base, with the vitroclastic texture best developed in the Blue Ranges (Fig. 4). At Snobs Creek, a second lenticulite horizon occurs about half way up the rhyolite indicating at least two cooling units in this particular area. Xenoliths of hornfels, representing country rock, occur most commonly in the lenticulite horizons. (2) Lake Mountain Rhyodacite. The Lake Mountain Rhyodacite is the uppermost unit in the cauldron, and forms the high-level, dissec- ted plateau of the Cerberean Ranges. It prob- ably reaches the order of 1000 m in thickness at the centre of the cauldron (Fig. 3). There was no apparent gap in extrusive history be- tween the rhyodacite and the Rubicon Rhyo- lite and textural evidence in general suggests that the two rock types comprised the one cooling unit. The rhyodacite shows only minor mineralo- gical and textural variations. The most sig- nificant is the presence of lighter-coloured, flattened lenses or schlieren (with similar mineralogy to the host rhyodacite) which may occur at all levels and over its entire areal extent. The term 'Snobite' (Hills, 1932) was applied to the most heterogeneous ('hybrid') varieties of the rhyodacite. At one locality in the southeast, near the settlement of Cambarville (Fig. 6), the base of the rhyodacite contains small, greenish, rhyolitic, lens-like inclusions. Xenoliths with- CERBEREAN CAULDRON V Vertical Horizontal 1000 m 3km Figure 3 — Cross sections of the complex. KEY (for Figs. 3-3). F 7 "] w«h XAGGERTY SUBGROUP tmans Hill Formation. Snobs Creek Formation. Blue Range Formation. Torbreck Range Formation. Robleys Spur Formation. CERBEREAN VOLCANICS. Rubicon Rhyolite. Lake Mountain Rhyodacite. a Ring and Radial Lykes. Central Intrusion: □ Cathedral and Koala Creek Beds. Basement Siluro- Devonian Sediments Alluvium. in the rhyodacite include hornfels and the Rubicon Rhyolite. One piece of graphite was recorded and probably represents carbonized wood (by analogy with the Snobs Creek Rhyo- lite). Ring and Radial Dykes, Satellite Intrusions The circular, outer ring fracture is filled for about 70% of its extent, with the dyke ranging from garnetiferous porphyritic granodiorite (which is the more common) to porphyritic granite. In the north-east, the main ring dyke is complicated by several small satellite intrusions in which both rock types may occur, e.g. Chris- tie's Hill. In the southeast, the dyke undergoes a com- plex branching (Fig. 7) with fractures curving inwards or backwards on themselves. Further south, the dyke is a continuous body, trending sub-parallel to the strike ridges of the Koala Creek Sandstone. The fracture finally bends westward to cut the volcanics without dyke fill (or significant vertical and horizontal displace- ment) near Cumberland (Fig. 6). This con- trasts with the situation in the southwest, where the ring fracture cuts the volcanics and offsets them by 3000 m horizontally and 500 m verti- cally (Fig. 8). Detailed mapping on the northeastern side of the complex revealed an inner ring dyke actu- ally cutting the volcanic pile. In the best ex- posure, on the Barnewall Plains Road, the T1 D W. D. BIRCH, A. J. W. GLEADOW, B. W. NOTTLE, J. A. ROSS, and R. WHATELY Figure 4 — The geology in the Blue Range. chilled margin is of porphyritic granite, but the bulk of the dyke is of porphyritic grano- diorite (Fig. 5). In the extreme north, Thomas (1947) mapped thin branching porphyry dykes intruding the volcanics. Of the radial faults, only two definitely con- tain dyke fill. These are the Torbrecic and Conn Hill Faults (Fig. 5). Opposite the New- man Plains Fault, Whitelaw (1913) mapped a small, radial, protrusion from the outer ring dyke. Central Intrusions Intrusion of two texturally-distinct, high- level granodiorites, marked the final stages in the evolution of the Cerberean Cauldron. (1) Porphyritic Microgranodiorite. The older and less-developed is a fine-grained granodio- rite with occasional large orthoclase pheno- crysts. It forms a stock, intruding and meta- morphosing basement sediments and volcanics, southeast of Buxton (Fig. 8). The intrusion has a small extension to the southeast along the continuation of the outer ring fracture, and within the fault zone the Rubicon Rhyolite has been dragged down and incorporated with- in the intrusion (Fig. 8). The microgranodio- rite also occurs as thin dyke-like bodies intrud- ing the Lake Mountain Rhyodacite in the Keppels Falls area. (2) Granodiorite. The final high-level grano- diorite intrusion was emplaced into the base of the volcanic pile in the Junction region be- tween the Cerberean and Acheron Cauldron. The mass is elongated north-south in outcrop, and its eastern contact with the Lake Mountain Rhyodacite appears fault-controlled (Fig. 6). On the western side, the contact between granodiorite and volcanics is not well exposed, but a shallow westerly dip under the volcanics is suggested. A significant feature of the intrusion is a complex and not easily interpreted zone ('Hy- brid' zone of Hills, 1932), exposed in cuttings on the highest part of the Marysville-Cumber- land road, and extending south and west. Large blocks of hornfels, partially granitized, with less common andesite and rhyodacite, are veined by granodiorite and in places the rock Biotite is abundant in the Lake Mountain Rhyodacite, but less common in the Snobs CERBEREAN CAULDRON types are strongly sheared. Bedding preserved in some of the hornfels blocks suggests that little movement relative to the basement has occurred. This hornfels region is at a much higher level than the original surface on which the volcanic rocks in the Cerberean Cauldron to the north were deposited (Fig. 3). A few thin dykes of late-stage hornblende porphyrite have been described in the Snobs Creek area by Thomas (1947) and Birch et ah (1970). Petrography of the Major Igneous Rocks Rhyolites Quartz (euhedral to subhedral and em- bayed) and potassium feldspar (orthoclase to intermediate microcline ) are the dominant phenocrysts in the rhyolitic rocks, with less common plagioclase (oligoclase-basic ande- sine) and biotite. A small proportion (up to 5 modal per cent) of iron cordierite crystals and rare almandine garnets characterize the Rubi- con Rhyolite (see Birch and Gleadow, 1974), but these two minerals are absent in the Snobs Creek and Robleys Spur Rhyolites. All three rhyolites show remnant eutaxitic and vitroclastic textures, as evidence for an ash flow origin. These are best developed in the basal, phenocryst-poor region of the Rubicon Rhyolite. Dense welding, followed by devitri- fication and recrystallization has tended to ob- literate original ignimbritic textures in the quartzo-feldspathic groundmass in all three rhyolites. Both the Snobs Creek and Rubicon Rhyolites are pale grey in colour, but the latter is distinctive due to its high phenocryst content or prominent eutaxitic textures in the low phenocryst varieties. The Robleys Spur Rhyo- lites are dark and aphanitic, the dark colour being due to finely-divided magnetite in the groundmass. The Rubicon Rhyolite grades into the over- lying Lake Mountain Rhyodacite with increase in phenocryst content and modal proportion of biotite and plagioclase. figure 5 — The geology in the Barnewall Plain — Snobs Link Road area. Rhyodacltes Quartz and plagioclase phenocryst phases in the are the dominant rhyodacitic rocks. 10 W. D. BIRCH, A. J. W. GLEADOW, B. W. MOTTLE, J. A. ROSS, and R. WHATELY Figure 6— The geology in the Cumberland region. Creek and Robleys Spur Rhyodacite. Ortho- clase is uncommon in all three rhyodacites. Small quantities of hypersthene and almandine characterize the Lake Mountain Rhyodacite. The groundmass in all three rhyodacites is microcrystalline, and only in the Robleys Spur Rhyodacite are remnant vitroclastic textures preserved. However, the continuity of the Snobs Creek and Lake Mountain Rhyodacites with their underlying ignimbritic rhyolites im- plies ash flow origins for both. Recrystalliza- tion has occurred to the greatest extent in the Lake Mountain Rhyodacite due to autometa- morphic effects resulting from its considerable thickness. The three rhyodacites differ considerably in appearance, with the Snobs Creek Rhyoda- cite being pale in colour, the Robleys Spur Rhyodacite dark with abundant small pheno- crysts, and the Lake Mountain Rhyodacite dark grey and biotite-rich. Basaltic Andesites The basaltic andesites of the Torbreck Range Andesites are dark bluish-grey, uniform and finegrained. Prismatic phenocrysts of au- gite and occasional labradorite occur in a groundmass of small plagioclase lathes, prisms of augite and iron ore grains. Groundmass al- teration is widespread however, producing chlorite, actinolite, carbonate and iron oxides. The basaltic andesites interbedded with the rhyodacites of the Robleys Spur Volcanics near Marysville are dark grey, compact rocks with abundant sub-rectangular phenocrysts of plagioclase showing oscillatory zoning and less common augite, the latter usually altered to a pale green amphibole. The groundmass is similar to that in the Torbreck Range Ande- sites, although secondary biotite has developed in the sequence near Marysville, due probably to contact metamorphism from the effect of the nearby ring dyke mass. CERBEREAN CAULDRON 11 Intrusive rocks The porphyritic granite and granodiorite in the ring dykes are the intrusive equivalents of the Rubicon Rhyolite and Lake Mountain Rhyodacite respectively. The granite contains large phenocrysts of quartz, white to pink in- termediate microcline perthites and less com- mon saussuritized plagioclase and biotite in a fine-grained, greenish, sericitized, quartzo- feldspathic groundmass. Pinitized cordierite is rare. The granodiorite consists of quartz, pla- gioclase, biotite and orthoclase in a medium grained groundmass. It is characterized by reddish garnets, usually partly altered to bio- tite, up to 1 -5 cm across. The two rock types forming the central in- trusions have normal granodioritic mineralogy. However, the porphyritic microgranodiorite contains large, conspicuous orthoclase pheno- crysts up to 3 cm long, and sillimanite and al- mandine are unusual, if rare, constituents and are probably xenocrystic. The granodiorite forming the larger intrusion is generally me- dium-grained and sub-equigranular, although it tends to be more porphyritic in the veins within the complex hornfelsic zone at the junction between the Cerberean and Acheron cauldrons. Structural Development Structural pattern The Cerberean Cauldron subsidence area is basically a cylindrical block which has sub- sided along an integrated pattern of circular and radial faults (Fig. 2). Greater subsidence at the centre of the block has led to basining of the volcanic pile, i.e. all formations dip in- wards (Fig. 3). Differential movement has also occurred on the radial faults. Where the Cerberean Cauldron adjoins the Acheron Cauldron there is no clearly defined structural break. The volcanic sequence on the western side is apparently continuous between the two cauldrons. Figure 7 — The geology in the region of the complex branching of the ring fracture in the south-east of the Cerberean Cauldron. 12 W. D. BIRCH, A. J. W. GLEADOW, B. W. NOTTLE, J. A. ROSS, and R. WHATELY Early faulting The earliest known fault associated with the cauldron structure is Snobs Creek Fault, which is parallel to and probably controlled by the regional basement trends (Fig. 2). The fault was operative at least as early as the extrusion of the Snobs Creek Formation, since these rocks are restricted to the downthrown, or western side of the fault in the northwest. Of the overlying formations, only the Torbreck Range Formation changes in thickness across the fault. The much greater subsidence on the fault line in the northwest, compared to the southeast, where the vertical displacement is apparently reversed, indicates a hinging effect (Fig. 5). The inner ring dyke, exposed between pairs of radial faults in the northeast, represents an early-formed fracture. The interpretation of the geology between the Conn Hill and Newman Plain Faults (Fig. 9) requires this fracture being present as a scarp during emplacement of the Torbreck Range Andesites, since thick accumulations of basaltic andesites occur against it (Fig. 5). This scarp was apparently not present at the time between the Barnewall Plains and Torbreck Faults further north. Al- though interpretation of this ring fracture's history is complicated, it may represent part of the margin of a caldera, of about 9 km diameter, dipping inwards at 50°-60°, and centred on Snobs Creek Fault. The gravity survey by Clarke et at. (1970) indicates a probable thickening of the volcanic sequence in this area. Later faulting (1) Ring Fractures. The outer and most con- tinuous ring fracture is almost perfectly cir- cular, with a 27 km diameter. Minor branching is caused by radial and other faults, but only Snobs Creek Fault offsets it. The complex zone of arcuate fractures de- veloped in the southeast has probably been caused by the intersection of the southeastern end of the Snobs Creek Fault with the outer ring fracture, and influenced by the northwest- southeast regional basement trends. Further to the south, the fracture pattern has been shaped by the rigid block of Koala Creek Sandstones (Fig. 7). Minor fractures paralleling the main fracture occur inside it, but are only detectable where dyke-filled. These include the dykes intruding the volcanics in the north (Thomas, 1947) and a partly dyke-filled fracture concentric with the outer ring dyke in the east. A gravity survey by Clarke et al. (1970) near Taggerty, shows an outward dip of 70° for the ring dyke, confirming earlier observa- tions by Hills (1959) and Thomas (1947). The offset of the ring fracture by Snobs Creek Fault in the north also indicates an outward dip. A further gravity and magnetic survey (Clarke et al 1970) near Buxton, indicates an anomalous body dipping outwards at 50°. If this is the sub-surface extension of the ring dyke, its shallow dip could explain the marked thickening of the dyke near Buxton (Fig. 8). In the southeast, where the ring fracture in- tersects the volcanic sequence with consider- able horizontal and vertical displacement (see Fig. 8), the Rubicon Rhyolite does not change markedly in thickness but has been dragged down across it. Thus, the major subsidence on the ring fracture must have postdated extru- sion of the Rubicon Rhyolite. (2) Radial Fractures. The mapping by Birch et al (1970) disclosed a set of eight ra- dial fractures, which focus at the geometric centre of the outer ring fractures (Fig. 2), implying a close relationship between all frac- tures. Movement on the radial fractures oc- curred late in the eruptive history of the Lake Mountain Rhyodacite, as they may offset its base (Figs. 5 and 8). Stream lineaments follow fault lines in the rhyodacite, across the top of the Cerberean Ranges. The Torbreck Fault (Fig. 5) was probably active at an earlier time, as the volcanic se- quence changes across it. Considerable drag on early formations is evident on this fault, and is also shown by the magnetic survey of Clarke et al (1970) near the Barnewall Plain Fault. Source Vents The feeder vents for the Snobs Creek Vol- canic were probably associated with Snobs CERBEREAN CAULDRON 13 Figure 8 — The geology in the Mt Margaret-Little River area. Creek Fault, as the formation is thickest close to the fault, which was active at the time of emplacement. The suspected caldera now rep- resented by the inner ring dyke may have been a feeder for the thick Torbreck Range Ande- sites. A coarse, basic agglomerate in the south- west of the cauldron (Fig. 8) with which a magnetic anomaly is associated (Clarke et al. 1970), indicates a likely eruption point of a more minor character. Because of the wide- spread occurrence of the basaltic andesites, it is likely that more vents such as this existed, but have been hidden by later volcanic erup- tions. Eruption points for the Robleys Spur For- mation are probably in part represented by agglomerates, for example, one at Barkers Gully in the north (Thomas, 1947) and one near Cumberland Junction in the southeast (Fig. 6). 14 W. D. BIRCH, A. J. W. GLEADOW, B. W. NOTTLE, J. A. ROSS, and R. WHATELY J[ Inner Ring Dyke Lake Mountain Rhyodacite Rubicon Rhyolite Torbreck Range Andesites ] Basement Sediments Figure 9— Block diagram illustrating the origin of the outcrop pattern between the Conn Hill and Newman Plain Faults. CERBEREAN CAULDRON 15 Large scale ignimbritic eruptions generally take place from fissures associated with vol- canic subsidence structures (Branch, 1967; Roberts, 1967; Ross and Smith, 1961). In the case of the Cereberean Volcanics, the most likely source of the ash flow eruptions is the ring fracture, as it contains intrusive equiva- lents of both the Rubicon Rhyolite and the Lake Mountain Rhyodacite. Tuffs with rhyo- litic fragments are included in the ring dyke at Cumberland Junction. The inner ring dyke may also have been a feeder as it contains in- trusive equivalents of both these rock types. Source vents may also have been situated along radial faults. The gravity survey by Clarke et al. (1970) indicated several anomalies which may represent buried vents (Fig. 2). A large early-formed central vent may also have de- veloped, evidenced by the focussing of the radial faults. Nature of the subsidence The subsidence of the Cerberean Cauldron can be divided into two components — a basin- ing and a ring fracture subsidence. Early basin- ing is suggested by decreasing inward dip of the volcanics progressively up the sequence (Tho- mas, 1947) and by the deposition of the Blue Range Formation under lacustrine conditions. Across the southeast, gravity data (Clarke et ah 1970) indicates a line of thickening of the volcanics (Fig. 2), suggesting a warp marking the edge of the main cauldron basin. To the south and east of this line, the volcanics are relatively flat lying (Clarke et al 1970). By restoring the downwarped basin to its original state, a diameter of about 31 km is obtained for the circular block, compared to the present ring dyke of 27 km diameter. The well documented outward dip of the ring dyke means that the 4 km difference between these diameters represents stretching of the central block. Most of this stretching must have oc- curred before the tensile stress was relieved by failure along the ring fracture. The major sub- sidence along the ring fracture postdates the Rubicon Rhyolite, but may have taken place progressively throughout emplacement of the Cerberean volcanics. A certain amount of bas- ining occurred after emplacement of the Lake Mountain Rhyodacite as the base shows some inward dip. Mechanism of the subsidence The mechanism of subsidence of the Cer- berean Cauldron has undoubtedly been one of foundering under gravity, following withdrawal of magmatic support due to the extrusion ot the Cerberean Volcanics. The relatively co- herent subsidence of the downthrown block in- dicates very rapid evacuation of the magma chamber, as discussed by Williams (1941), and is borne out by the ash flow emplacement of the Cerberean Volcanics as one or two large cooling units. The fracture pattern and stretch- ing of the foundered block suggest that the roof of the magma chamber behaved as a cir- cular plate with a thickness considerably less than its diameter. The timing and cause of formation of the ring and radial fractures are problematical, but there are two possibilities. 1. Eruption of magma for the Cerberean Vol- canics was initiated through central vents in the cauldron area, until enough magma had been withdrawn to enable gravitational collapse of the central block along ring and radial faults. 2. Pressure build-up in the magma chamber led to explosive failure of the roof rocks along ring and radial faults. These would relieve the pressure and act as feeders for the eruption of the Cerberean Volcanics. Subsidence would then occur as the magma chamber was emptied. In evaluating these possibilities, the follow- ing evidence must be considered: 1. The geometry of the fracture pattern is highly regular and is apparently little in- fluenced by regional basement trends. 2. The region was apparently under tension. 3. Chemical evidence (Birch, 1975) suggests that a volatile phase could have been gene- rated at the top of the Rubicon Rhyolite magma at some interval prior to eruption. 4. The ring fracture dips steeply outwards (at the surface). The first three lines of evidence suggest that a 'point' explosion at depth was responsible 16 W. D. BIRCH, A. J. W. GLEADOW, B. W. NOTTLE, J. A. ROSS, and R. WHATELY for the fracture pattern. It is difficult to im- agine purely gravitational collapse producing a perfectly circular ring fracture through a linea- ted crustal plate. A fracture pattern produced by an explosion at depth would be extended and enlarged by a rebound effect. Eruptions from those fractures which extended through to the surface could then permit the wedge- shaped fragments to subside into the magma chamber, at the same tune initiating or increas- ing eruptions along the ring fracture. The evidence from the dip of the ring frac- ture is inconclusive. While inward dips have been ascribed to pressure build-up by a num- ber of workers (Anderson, 1936; Williams, 1941; Robson and Barr, 1964; Roberts, 1967; Smith and Bailey, 1968), the theoretical treat- ment of the problem of the fracture patterns derived from either withdrawal of magmatic support or upward magmatic pressure is by no means rigorous. For example, Robson and Barr (1964) suggest that steeply outward dipping ring fractures may result from either mechanism. Emplacement of high level intrusions Following the volcanic episode, with its associated dyke intrusions, came a culminat- ing period of intrusion into the base of the volcanic pile. Such high-level extrusive ac- tivity is typically the final magmatic event in similar complexes throughout the world (ci. Branch, 1966; Kingsley, 1931; Oftedahl, 1952; Jacobson et ah, 1958). The intrusion proceeded in two stages and was largely controlled by pre-existing lines of weakness. The first phase is represented by the porphyritic microgranodiorite. While its intrusion along the main Cerberean ring frac- ture in the southwest was permissive, its em- placement in the Keppels Falls area, along the continuation of the main boundary fault of the Acheron Cauldron may have produced local- ized gneissic textures in the Lake Mountain Rhyodacite. The second-phase granodiorite is more ex- tensive, forming a relatively flat-roofed intru- sion. The fault controlling the intrusion's eastern boundary is continuous to the south with the main boundary fracture of the Acher- on Cauldron (Dudley et at. 1971). The ap- parent shallow westerly dip of the western mar- gin of the intrusion may explain the high degree of contact metamorphism of the se- quence on Robleys Spur, east of Marysville. Intrusion probably occurred by quiet stop- ing, as the magma apparently caused no resur- gent vulcanism, yet cooled within a kilometre of the surface. The region of granitized horn- fels described previously was probably a struc- tural high at the time of cauldron formation both to the north (Cerberean) and south (Acheron). That thin volcanic sequences may have been deposited on it is suggested by the occurrence of isolated, flat-lying, metamor- phosed remnants of the Lake Mountain Rhyo- dacite, forming the highest part of the topo- graphy (Mt Stinton and Mt Grant, Figs. 2 and 3). Acknowledgements This work was completed as part of the authors' B.Sc. (Hons.) thesis, in 1970. We are grateful for the assistance and advice of Dr R. J. W. McLaughlin and Dr A. Cundari of the Geology Department, University of Mel- bourne. Bibliography Anderson, E. M„ 1936. The formation of cone sheets, ring dykes and cauldron subsidences. Proc. Roy. Soc. Edinburgh, 56: 128-157. Bell, G., 1959. Jamieson, Victoria. 1: 63 360 sheet. (Geological Survey of Victoria). Birch, W. D., 1975. Mineralogy, Geochemistry and Pedogenesis of some Victorian Palaeozoic Rhyo- lites. Unpub. Ph.D. thesis, University of Mel- bourne. Birch, W. D. and A. J. W. Gleadow, 1974. The genesis of garnet and cordierite in acid volcanic rocks: Evidence from the Cerberean Cauldron, Central Victoria, Australia. Contr. Mineral, and Petrol. 45: 1-13. Birch, W. D., A. J. W. Gleadow, B. W. Nottle, J. A. Ross and R. Whately, 1970. An inves- tigation of the Upper Devonian Cerberean Caul- dron Complex in Central Victoria. Unpub. B.Sc. (Hons.) thesis, University of Melbourne. Branch, C. S., 1966. Volcanic Cauldrons, Ring Com- plexes and associated granites of the Georgetown Inlier, Queensland. Bull. Bur. Miner. Res. Geol. Geophys. 76. CERBEREAN CAULDRON 17 Branch, C. D., 1967. The source of eruption for pyroclastic flows; cauldrons or calderas. Bull. Vole, 30: 41-50. Clarke, G. F., P. B. Edwards, D. R. Eyles, A. G. Power and H. T. Wood, 1970. Geophysical studies of the Cerberean Cauldron. Unpub. Man., University of Melbourne. Dale, G. R., 1964. The geology of the Cathedral Range area (Victoria). R.M.I.T. Fellowship Thesis (unpub.). Dudley, R., J. Dudley and A. Rossiter, 1971. The geology of the Acheron Cauldron, Victoria. Un- pub. B.Sc. (Hons.) thesis, University of Mel- bourne. Hills, E. S., 1929. The geology and palaeontology of the Cathedral Range and the Blue Hills in north-western Gippsland. Proc. Roy. Soc. Vic, 41: 176-201. , 1931. The Upper Devonian fishes of Vic- toria, Australia, and their bearing on the strati- graphy of the State. Geol Mag. 68: 206-231. -, 1932. The geology of Marysville, Victoria. Geol Mag. 69: 145-166. , 1959. Cauldron subsidence, granitic rocks and crustal fracturing in south-eastern Australia. Geol. Rund. 47: 543-561. Howard, R. T., 1972. Acheron Cauldron and Black Range. Unpub. B.Sc. (Hons.) Thesis, University of Melbourne. Jacobsen, R. R. E., W. N. Macleod and R. Black, 1958. Ring complexes in the younger granite pro- vince of northern Nigeria. Mem. Geol. Soc. Lon- don, 1. Kingsley, L., 1931. Cauldron subsidence of the Ossi- pee Mountains, New Hampshire, Amer. J. Set, 22: 130-168. Marsden, M. A. H., 1976. Upper Devonian-Carboni- ferous. In 'Geology of Victoria', Geol. Soc. Aust. Spec. Pub. No. 5, Ch. 5. Oftedahl, C, 1952. Cauldron subsidences of the Oslo region. Int. Geol. Cong., 18th Session, Gt. Brit., 13: 205-213. Roberts, J. L., 1967. Ignimbrite eruptions in the Glencoe Cauldron Subsidence. GeoL JL, 5: 174- 184. Robson, G. R. and K. G. Barr, 1964. The effect of stress of faulting and minor intrusions in the vicinity of a magma body. Bull. Vole, 27: 314- 329. Ross, C. S. and R. L. Smith, 1961. Ash-flow tuffs, their origin, geologic relations and identification. Prof. Paper. U.S. GeoL Surv., 366. Smith, R. L. and R. A. Bailey, 1968. Resurgent cauldrons. Mem. Geol, Soc. Amer. 116: 613- 662. Thomas, D. E., 1947. The geology of the Eildon Dam project. Mem. Geol. Surv. Vic. 16. Vandenberg, A. H. M., 1977. Warburton, Victoria. 1: 250 000 sheet (Geological Survey of Vic- toria) . Whitelaw, O. A. L., 1913. Unpublished map of region west of Cerberean Cauldron. (Geologi- cal Survey of Victoria). Williams, H. 1941. Calderas and their origin. Bull. Calif. Univ. Dept. Geol. Sci. 25: 239-346. GASTROPODS FROM THE EARLY DEVONIAN BELL POINT LIMESTONE, CAPE LIPTRAP PENINSULA, VICTORIA By C. B. Tassell Albany Residency Museum, Albany, Western Australia Abstract One new genus and five new species of gastropods are described from the late Early Devonian Bell Point Limestone, on Cape Liptrap Peninsula, 150 km south-east of Melbourne. Six other gastropod taxa are recorded. The new genus Bassotrochus is assigned to the family Omphalotrochidae. Together with Orecopia murrayi sp. nov., also from the Bell Point Lime- stone, Bassotrochus angulatus sp. nov. represents the earliest known occurrence of this family to date. The other species are Tropidodiscus waratahensis sp. nov., Straparollus (Euomphalus) sp., ? Mourlonia sp., Gyronema bellense sp. nov., Murchisonia (Murchisonia) bassensis sp. nov., M. (Murchisonia) sp. A, M. (Murchisonia) sp. B and M. (Hormotomina) sp. Also described are Micromphalus clarkei (de Koninck) from near Yass, N.S.W. and an indeterminate form from the Waratah Limestone, Cape Liptrap Peninsula. This shallow water gastropod fauna occurs as both coquinite and non-coquinite assem- blages associated with a diverse invertebrate community which includes corals, stromatopor- oids, bivalves and brachiopods. Introduction The Bell Point Limestone occurs as a small faulted block on the eastern side of Cape Lip- trap Peninsula which is approximately 150 km south-east of Melbourne. Exposed on this peninsula are sections of the Waratah Bay axis, one of a number of such structural axes occurring in Victoria. This axis has been the site of considerable tectonic activity and as a consequence some major faults occur on the eastern side of the peninsula. Besides the Bell Point Limestone, certain other units are also only exposed in a few small areas. These in- clude the Early Devonian Waratah Limestone and the Early Ordovician Digger Island Lime- stone, The limestones at Cape Liptrap have been known for some time, Ulrich (1875) being the first to mention them in print. However, it was not until recently that the stratigraphy of the area has become adequately known. The most recent account of the geology being that of Singleton (1968). McCoy (1877) described the first fossil Palaeopora inter stincta ( Wahl ) from the Waratah Bay area. In 1898 Etheridge noted a dorsal valve of a spirifer, some crinoid frag- ments and a new coral Tryplasma. Since then some elements of the limestone fauna including corals, ostracods and brachiopods have been described. Although abundant at Bell Point, gastro- pods have only been mentioned in passing un- til now. Lindner (1953, p. 82) noted 'a trocho- form gastropod is the only fossil in the lowest 75 feet'. Above this spiriferid brachiopods oc- cur together with the gastropod. Higher in the section 'several small species of gastropods, in- cluding turreted and turbinate forms' occur. Similarly, Teichert (1954) listed gastropods amongst the abundant fauna of the Bell Point Limestone at Bell Point. Talent (as cited by Singleton, 1968) listed Bellerophon and Coelocaulus amongst the gastropods occurring in the Waratah Lime- stone. The gastropods listed for the Bell Point Limestone include Tropidodiscus, Amphelissa isisensis and 'a new Trochid'. Bounded by faults and in close proximity to the large Waratah fault, the Bell Point Lime- stone has been subjected to some stress. Partly as a result of this many of the gastropods are distorted, in addition to the crushing and fragmenting of the shells associated with the initial compaction of the sediment. As many of the specimens are either crushed, fragmented or distorted the measure- ments given can only be considered as a gene- ral indication of the specimens' original shape. All measurements are in millimetres and the following symbols relating to the measurements have been used: c, number of spiral cords; Hap, height of aperture; Ht, total height of shell; L, length measured at the selenizone in 19 IQ C, B. TASSELL the bellerophontids, Wap, width of aperture; Wh, total number of whorls in shell; Wt, total width of shell. All specimens used in this study are lodged in the palaeontological collection of the Na- tional Museum of Victoria. Age of the Faunas When Talent (1965) discussed the Bell Point Limestone he commented upon the simi- larity of the brachiopod fauna to that in the Buchan Caves Limestone. Such a similarity is also found amongst some of the other faunal elements of the two limestones (Vandenberg et at., 1976). Philip and Pedder (1967) recog- nized their coral-conodont Fauna D in both the Buchan Caves Limestone and the Taemas Limestone. This coral-conodont fauna is now considered to be Early Emsian in age (Strusz, 1972). In the older Waratah Limestone, Talent (1965) recognized a fauna containing many elements which showed considerable similarity to that found in the Coopers Creek Formation. The coral-conodont Fauna B of Philip and Pedder which has been found in both the Waratah Limestone and the Coopers Creek Formation is now considered to be Late Sie- genianin age (Strusz, 1972). As with previous studies of middle Palaeo- zoic gastropods in southeastern Australia, this study makes little contribution toward age de- terminations of the limestones. Certainly Mur- chisonia (Murchisonia) sp. A from the Bell Point Limestone is very similar to Murchisonia (Murchisonia) turris de Koninck known to oc- cur in the Emsian 'Receptaculites' Limestone at Taemas, N.S.W. Similarly the indeterminate cyrtoconoid gastropod from the Waratah Lime- stone would appear to have an affinity with the poorly preserved Michelia sp. from the simi- larly aged Toongabbie Limestone. What is of considerable interest is the pre- sence of the genera Orecopia, Bassotrochus and Straparollus (Euomphalus) in the gastro- pod fauna. Both Orecopia and Bassotrochus are members of the Omphalotrochidae, a fam- ily known principally from the Late Palaeo- zoic. The presence of these genera at Bell Point constitutes the earliest occurrence of the family to date. Their presence and that of Micromphalus clarki in the Late Silurian sedi- ments of the Yass Basin, indicate an affinity not previously suspected of the Australian gas- tropod fauna with younger Devonian gastro- pod faunas of the Northern Hemisphere (Tas- sell, 1976). Palaeoecology The Bell Point Limestone is a well bedded dark muddy limestone within which minor amounts of lighter grey limestone and black shale occur. Lindner (1953) considered that it was at least 140 feet thick. Although there has been some disagreement about the depth of de- position of the Early Devonian sediments else- where in the Melbourne Trough it is generally agreed that the carbonate sediments occurring on the Waratah Bay axis were deposited in shallow water (Singleton, 1968; Vandenberg etal. 1976). Part of the prolific and diverse fauna of the Bell Boint Limestone has been described in the past (Hill, 1954; Krommelbein, 1954; Talent, 1956; Pedder, 1965). The fauna includes stromatoporoids, tabulate and rugose corals, brachiopods, bivalves, rostroconchians, gastro- pods and ostracods. The faunas in this unit are preserved in two ways, either as coquinite assemblages or non-coquinite assemblages. Al- though less frequent the coquinite assemblages are extremely prominent, for example, assem- blages of Orecopia murrayi and Bassotrochus angulatus thickly cover surfaces sometimes ex- ceeding 25 sq m in area. A characteristic of these coquinite assemblages is their low faunal diversity. This is true for any such assemblage whether it is composed of gastropods, bivalves or any other faunal component. Similar, al- most monospecific, coquinite assemblages have been reported in Eastern North America (Mo- Ghee, 1976). Coquinas composed largely of O. murrayi are the most frequently occurring gastropod as- semblages in the limestone. They consist of complete specimens of O. murrayi ranging in diameter from 4 mm upwards, as well as large quantities of angular shell fragments of BELL POINT LIMESTONE GASTEROPODA 21 the same species. Both the fragments and the complete shells lack evidence of abrasion and transport. S. (Euomphalus) sp. and B. angula- tus also occur frequently in O. murrayi coquin- as. Similarly O. murrayi occurs in some of the coquinas composed of B. angulatus or the pre- sently undescribed bivalves. In contrast to the spectacular coquinas of 0. murrayi and B. angulatus the coquinas or bedding plane concentrations of Tropidodiscus waratahensis are quite restricted, rarely ex- ceeding a sq m in area. Unlike O. murrayi which occurs in association with bivalves in some coquinas, those of T. waratahensis contain numerous brachiopods. Again, evidence of transport or abrasion is lacking. The non-coquinite assemblages possess a much greater faunal diversity. For example M. (Murchisonia) bassensis occurs in the light grey limestone with rugose corals, bivalves, bra- chiopods, rostroconchians as well as occasional individuals of O. murrayi. M. (Murchisonia) sp. B found in a similar limestone is associated with rugose corals, ostracods, bivalves, bra- chiopods, rostroconchians and infrequent ex- amples of T. waratahensis. The composition of this gastropod fauna differs considerably from any other non-platy- ceratid fauna of similar age known from south- east Australia. Unlike the Lilydale fauna which is dominated by turbiniform and high spired gastropods, notably 'Cyclonema' and some murchisoniaceans, the Bell Point Limestone fauna is dominated absolutely by the medium- sized rotelliform O. murrayi. High spired and turbiniform gastropods form only a small com- ponent of the fauna in numerical terms al- though they account for much of the fauna's diversity. Systematic Descriptions Family BELLEROPHONTIDAE McCoy, 1851 Subfamily TROPIDODISCINAE Knight, 1956 Genus Tropidodiscus Meek and Worthen, 1866 Type Species: Bellerophon curvilineatus Con- rad, 1842; Lower Devonian; Schoharie, New York. Tropidodiscus waratahensis sp. nov. (Plate 1, figure 1) Diagnosis: small form of genus with subdued dorsal crest and fine closely spaced growth lines. Description: Small form with angular dorsal crest; whorl profile gently rounded between dorsal crest and edge of the umbilicus, then rounds more strongly into the umbilicus; um- bilici deep and wide; aperture sub-triangular with a ridge formed by the dorsal crest of pre- ceding whorl on the inner lip; inner lip thin; thick strongly prosocyrt outer lip passes back- wards obliquely from the suture to form a moderately deep v-shaped sinus at the dorsal crest, narrow convex selenizone raised above former edges of the slit; fine closely spaced col- labral lines. Dimensions: L Wh P46963 75 — P46964 3 2 P46965 4 3 + P46966 7 3+ P46967 8 — Location of Types: National Museum of Vic- toria. Holotype, P46963. E. D. Gill Coll. Material: Holotype and 20 other specimens. Discussion: Besides being substantially smaller than the type species, T. waratahensis has finer, more closely spaced collabral lines. T. centrifugalis (Chapman) from Lower Devon- ian mudstones at Killara and Loyola in the Melbourne Trough is of similar size to T. waratahensis. However, its prominent dorsal crest and widely spaced, frequently foliaceous collabral lines distinguish it from the Bell Point Limestone species. An undescribed species from the Lilydale Limestone is smaller and has more prominent collabral lines than T. warata- hensis. Talent (1963) described Tropidodiscus sp. A from the Kilgower Member, Tabberabbera. This species does have fine closely spaced pro- socyrt collabral lines similar to those of T. waratahensis. Poor preservation of the slightly larger Tabberabbera forms precludes further comparisons. C. B. TASSELL Family EU0MPHAL1DAE de Koninck, 1881 Genus Straparollus Montfort, 1810 Subgenus Straparollus (Euomphalus), J. Sowerby, 1814 Type Species: Euomphalus pentangulatus J. Sowerby, 1814; Lower Carboniferous; near Dublin, Ireland. Discussion: The variability of specific charac- ters in this genus has been commented upon by numerous authors (Knight, 1934; Batten, 1966). The latter author cited the example of 5. (E) acutits (J. Sowerby) which possesses the high spired shell of S. (Straparollus) and the well developed angulate whorl of S. (Euom- phalus). The limited material from the Bell Point Limestone provides a further example of this variability. Straparollus (Euomphalus) sp. (Plate 1, figures 15,21) Description: Medium to large low spired gas- tropod with angular whorl profile; sutures dis- tinct but variable in form during growth; nu- merous whorls with profile which varies with growth and the degree of development of the upper keel; initially whorls gently rounded to nearly flat between sutures; with growth, profile flat to gently concave and inward sloping be- tween the upper suture and the variability de- veloped upper keel at the junction of the upper and outer whorl surfaces; the junction of the outer and basal whorl surfaces particu- larly in the late growth stages forms a pro- minent narrow thickened peripheral frill; be- tween the upper keel and the basal frill the whorl profile is gently rounded; base gently rounded to circum-umbilical swelling where it becomes more strongly arched; narrow distinct umbilicus; aperture sub-elliptical; short thick- ened columellar lip slightly oblique outwards; parietal lip thin; outer lip of moderate thick- ness extends forwards gently from the upper suture to just above the peripheral frill where it rounds gently before crossing the base in a strongly oblique manner; the columellar and outer lips on the base are strongly concave; where a keel is present a shallow concave flexure of the lip may be developed on it; the closely spaced growth lines range from fine to moderately coarse on both the outer and basal whorl surfaces; the growth lines continue into the umbilicus. Dimensions: Ht Wt Hap Wap Wh P47503 24 59 7 18 5 P47564 25 53 — — 34- P47505 — — — — 6 P47506 15 51 — — 5+ Location of Types: National Museum of Vic- toria. Figured specimen P47503. Collected T. A. Darragh, 1971. Materials: One figured specimen and eight other specimens all of which are either crushed or distorted. Discussion: The small number of specimens and their poor state of preservation precludes detailed comparisons wtih other members of the genus and the formal naming of the spe- cies. The Bell Point form differs considerably from S. (Euomphalus) pentangulatus. The latter is a large discoidal gastropod with a wide um- bilicus and thin lips which contrasts with the low spire, narrow umbilicus and thickened lips of the Bell Point form. The shape of both the aperture and lips together with the distinct peripheral frill of S. (Euomphalus) sp. also serve to distinguish it from the type species. The Bell Point form can be distinguished quite readily from the other members of the genus previously described from Victoria. The low spire, narrow umbilicus, aperture shape, thickened lips and relatively few whorls dis- tinguish it from the dextrally coiled S. (E.) northi (Etheridge) from the Lilydale Lime- stone. S.(S.) serpenteus Talent from the mud- stones of central Victoria differs from the Bell Point form in its smaller size, trochiform shape, more numerous whorls, wider umbili- cus, apertural shape, growth line characters and whorl profile. Talent (cited by Singleton, 1968) included the gastropod Amphelissa isisensis in the faunal list for the Bell Point Limestone. Of all the forms known from this limestone, 5. (Euomphalus) sp. most closely resembles this species. A. isisensis, the type species of the genus, was described by Etheridge (1921) from BELL POINT LIMESTONE GASTEROPODA 23 the Isis River area, Scone district, N.S.W. This gastropod occurs in one of two faunas in the Timor Limestone Member regarded by Voi- sey and Packham (1969) as Givetian. Unfortunately the quality of Etheridge's il- lustrations and the nature of his description makes comparison difficult. Etheridge (p. 2) noted that 'as the individual specimens vary much in their state of preservation it is difficult to select features of a constant character other than those of the strongly angular and upstand- ing whorls, deeply seated sutures and nearly flat bases'. Subsequently Knight et ah (1960) synonymized Amphelissa with the subgenus S. (Euomphalus), Although poorly preserved, in each case the similar sized S. (E.) isisensis and S. (Euom- phalus) sp. do possess certain similarities. Both are low spired with 5 to 6 whorls. Both possess relatively flat bases with a narrow umbilicus. There are also some differences between the two forms. While S. (E) isisensts appears al- ways to possess a distinct angular keel and deep distinctive sutures this is not always the case with 5. (Euomphalus) sp. Similarly the generally distinct peripheral frill of 5. (Euom- phalus) sp. does appear to be more variably developed on S. (E.) isisensis. Certainly it can be clearly seen in one of Etheridge's illustra- tions (plate 1, fig. 1) accompanying his original description. Other details, notably those of the apertural region are lacking for S. (E.) isisensis. Austerum carinatum the type species of the genus Austerum was described by Hei- decker (1959), from Givetian limestones near Charters Towers, north Qld. This large, low spired species possesses a number of charac- teristics similar to those of the Bell Point form. Both are large, low spired gastropods with sub-elliptical apertures, thickened lips and concave basal and collumellar lips on the basal surface. There is also a flexure on the keel. The shallow umbilicus, persistence of the keel through most growth stages, consistent deep sutures and lack of a peripheral frill all dis- tinguish the north Qld. form from S. (Euom- phalus) sp. If specimens P47512 — 15 from near Arthur's Peak in the Fanning River Dis- trict, north Qld. are indicative of the variations in this species then there is considerable varia- tion in the height of the shell (pi. 1, figs. 16, 19). These specimens are distinctly trochiform in shape, another feature which distinguishes them from 5. (Euomphalus) sp. A re-examina- tion of the type material of A. carinatum is required. Whether this results in this genus be- ing synonymized with Straparollus (Euompha- lus) or establishes a subgenus of Straparollus characterized by features such as its thickening of the columellar and outer lips remains to be seen. Certainly the presence of at least two Devonian low spired euomphalids forms with substantially thickened lips in eastern Australia is of interest. Genus Micromphalus Knight, 1945 Type Species: Micromphalus turris Knight, 1 945 ; Lower Carboniferous; Elizabethtown, Kentucky, United States of America. Range: Late Silurian to Lower Carboniferous. The presence of a species of this genus near Yass extends the lower limit of its range from Middle Devonian to Late Silurian. Distribution: North America and Australia. This is the second record of the genus in Aus- tralia; Roberts et ah ( 1967) noted Micrompha- lus sp. amongst a fauna from the Frasnian Westwood Member in the Bonaparte Gulf Ba- sin of northern Western Australia. Micromphalus clarkei (de Koninck), 1876 (Plate 1, figures 20, 23) 1 876 Euomphalus (Omphalotrochus) clarkei de Koninck, p. 41, pi. 1, figs. 7, 7a-7b. Diagnosis: Large low spired form of genus with strongly embracing whorls and narrow shoulder. Description: Large, low spired trochiform gas- tropod with each whorl strongly embracing that above; distinct, moderately narrow shoulder; flat to strongly rounded; gently rounded to flat whorl profile from edge of shoulder to angular basal whorl periphery; initially sutures im- pressed, with growth become canaliculate; base rounded; small umbilicus present; aper- ture circular or nearly so; reflexed columellar lip; thin parietal inductura, extends forward of aperture; moderately thick irregularly pro- socline outer lip; from upper suture the outer 24 C. B. TASSELL lip passes backwards to below the shoulder where a distinct but variable shallow concave flexure or sinus is developed; the outer lip then continues more gently to the whorl peri- phery; the base is gently rounded; shell of moderate thickness; growth lines strongly fo- liaceous. Dimensions: Ht Wt Hap Wap Wh P47516 26 58 — — 4+ P47517 39 59 — — 5 P47519 32 49 13 14 4+ P47520 36 62 — — 5 Location of Types: 1. Euomphalus (Omphalo- trochus) clarkei, the specimen figured by de Koninck was apparently destroyed when the Garden Palace in Sydney was burnt on Sep- tember 22nd, 1882. 2. Micromphalus clarkei, National Museum of Victoria. Hypotypes P47517 and P47519, A. J. Shears- by Coll. Type Locality: 1. Euomphalus (Omphalotro- chus) clarkei, Yass District, NSW. 2. Micromphalus clarkei, Limestone or Der- ringullen Creek, near Yass, NSW. Stratigraphic Range: Although Shearsby's lo- cality data was vague the nature of preserva- tion of the material from Limestone Creek would suggest that it comes from the Bow- spring Limestone, a unit containing silicified gastropods and brachiopods (Dr K. S. W. Campbell pers. comm.) Brown (1941) has re- corded the occurrence of this gastropod in the Bowspring Limestone, near Yass. This unit was considered by Link (1970) to be Middle Lud- lovian in age. Material: 2 hypotypes and 4 other specimens. Discussion: When Knight (1945) erected the genus Micromphalus, he noted that M. turris possessed features similar to both Orecopia and Straparollus. That a form should exhibit features suggestive of a position intermediate to the families Euomphalidae and Omphalotro- chidae is consistent with Yochelson's (1956) contention that the omphalotrochids were de- rived from the euomphalids. Af. clarkei differs from the type species in possessing a lower spire, narrower shoulder, considerably stronger embracing whorls, more variable sutures and being considerably larger. Both possess the distinctive prosocline outer lip with a shallow sinus and narrow umbilicus. De Koninck, when describing M. clarkei, was aware of its similarity not only with the genus Euomphalus but also with the then re- cently established genus Omphalotrochus. As a consequence he placed the species from near Yass in the genus Euomphalus and the sub- genus Omphalatrochus, Family OMPHALOTROCHIDAE Knight, 1945 Genus Orecopia Knight, 1945 Type Species: Platyschisma ? mccoyi Walcott, 1884; Upper Devonian; Nevada, United States of America. Discussion: The presence at Bell Point of this genus extends its known distribution. Pre- viously it was confined to Europe and North America. The Bell Point occurrence also ex- tends the lower limit of the generic range from Upper Devonian to Lower Devonian. Orecopia murrayi sp. nov. (Plate 1, figures 6, 8, 12, 13) Diagnosis: Rotelliform member of genus fre- quently lacking umbilicus, with an ovoidal aper- ture and variably developed sinus in the outer up. Description: Medium rotelliform gastropod with a broad shallow rounded sinus in the outer lip; whorls each embracing that above; whorl profile flat to gently arched between su- tures; sutures sharp, incised, shallow and ir- regular; basal periphery strongly rounded; base flat to gently rounded; small variably de- veloped umbilicus sometimes present aperture ovoidal; columellar lip thickened, and strongly oblique outwards thins towards junction with outer lip; parietal inductura of moderate thick- ness, occasionally developed forward of aper- ture; moderately thick outer lip with u-shaped sinus of variable depth; from upper suture the outer lip passes backwards before rounding in sinus and passing forwards to the periphery BELL POINT LIMESTONE GASTEROPODA 25 where it turns roundly before continuing back- ward, obliquely across the base; below rounded sinus broad forward projection vari- ably developed at or just above the periphery; secondary deposits variably developed in peri- pheral angle; growth lines fine to foliaceous on both outer whorl surface and base; when um- bilicus is open growth lines continue into it; occasional growth rugae occur; no other sculp- ture developed. Dimensions: Ht Wt Hap Wap Wh P46987 15 29 — — 6+ P46988 — 35 5 11 P46989 — , 15 — P46992 . — 23 4 8 P46993 — — 6 8 P46994 8 15 — — 8 P46996 11 17-5 3 — 6+ P46999 13 27 — . — Location of Types: National Museum of Vic- toria. Holotype, P46987, Paratypes, P46988, P46989 and P46990. Collected C. B. Tassell, 1975. Material: Holotype, 3 paratypes and more than 100 other specimens, most of which are crushed or fragmented. Discussion: O. mccoyi (Walcott) is quite vari- able in shape, Knight (1945, pi. 80, figs, la-g) figured forms ranging from rotelliform to bee- hive like. In contrast O. murrayi is typically rotelliform. Comparison between the Bell Point species and the more rotelliform ex- amples of O. mccoyi reveals few differences. 0. murrayi is generally lower spired with a more ovoidal aperture resulting from a longer more outwardly oblique columellar lip. The fre- quent absence of an umbilicus, lack of an an- gulation on the upper whorl surface, and the apparently more variable depth of the sinus in the outer lip also distinguish O. murrayi from the type species. 0. murrayi has the peripheral angle thick- ened by secondary deposits. As a consequence the thinnest and weakest section of the base is at the inner edge of the peripheral thickening. When crushed as is frequently the case at Bell Point, the base is broken and fragmented most severely at the inner edge of the peri- pheral thickening. A similar pattern of fractur- ing occurs on the base of Scalaetrochus lind- stromi Etheridge from the Lilydale Limestone (Tassell, 1976). Typically the fragmented thin outer shell layer on the base has been lost. Bassotrochus gen. nov. Type Species: Bassotrochus angulatus sp. nov.; Lower Devonian, Bell Point Limestone, Bell Point, Victoria. Description: Trochiform gastropod with dis- tinct peripheral frill; whorl profile gently con- vexly arched to nearly flat; base gently rounded to flat; broad shallow sinus developed in up- per half of outer lip; broad anterior projection developed beneath sinus and above periphery; shell thickened by secondary deposits. Stratigraphic Range: Lower Devonian. Distribution: Bell Point, Victoria. Discussion: This genus was established with some hesitation because of the poor preserva- tion of the material of the type species B. angulatus. But the distinctive sinus, albeit shal- low, on the upper part of the outer lip and the protruding lower section of the outer lip clearly indicate its familial relationships. The shallow sinus, peripheral frill, whorl profile and lack of an umbilicus all serve to distinguish this genus from any of the other trochiform genera of this family. Of the genera currently assigned to the family Omphalotrochidae, Babylonites from the Permian of North America most closely resembles this genus. These two tro- chiform gastropods have a relatively shallow sinus and weakly developed anterior projection in the outer lip when compared to Omphalo- trochus and Orecopia. Both Bassotrochus and Babylonites possess a thickened periphery. In either genus a shallow depression can be de- veloped in the thickened periphery which tends to accentuate the peripheral frill. However, Bassotrochus angulatus lacking an umbilicus but possessing a convex whorl pro- file differs considerably from the species of Babylonites which possess a distinctly concave to nearly flat whorl profile and an umbilicus. The broad anterior projection beneath the sinus on the outer lip of Bassotrochus reaches its most anterior development above the peri- phery, not at the periphery as in Babylonites. 26 C. B. TASSELL Although variably developed the peripheral frill of Bassotrochus is generally sufficiently distinct to distinguish it from the blunter peri- phery of Babylonites. The thickened columel- lar lip and its more strongly prosocline nature also distinguish B. angulatus from the Ameri- can species. The existence of two genera of the family Omphalotrochidae in the Bell Point Lime- stone is of considerable interest. The genus Orecopia from the Upper Devonian of North America and Europe has been known for some time (Pedder, 1966). Its occurrence at Bell Point enlarges its range to include the Lower Devonian. The presence of Orecopia and Bassotrochus at Bell Point not only ex- tends the range of the family but increases the diversity of the family in the Devonian. How- ever, the continuing absence of forms during much of the Carboniferous means that Yochel- son's (1956, p. 201) comment that 'it is im- possible to construct any meaningful family phylogeny at this time' is still valid. The generic name Bassotrochus is derived from the trochiform shape of the type species and the close proximity of the Bell Point Limestone to Bass Strait. Bassotrochus angulatus sp. nov. (Plate 1, figures 7, 14, 17, 18, 22) Description: Medium to large trochiform gastropod with thickened peripheral frill; su- tures distinct, variable from adpressed to being overhung by periphery of preceding whorl; whorl profile gently convex to nearly flat from upper suture to near periphery; shallow de- pression frequently developed in thickened periphery emphasizes basal peripheral frill; base gently rounded to flat; umbilicus appar- ently lacking; aperture ovoidal; thickened pro- socline columellar lip with very broad shallow sinus; thins towards junction with outer lip; parietal lip thin or wanting; outer lip of moderate thickness with a very shallow broad sinus in the upper part; from the upper suture the upper lip passes backwards a very short distance before rounding in the shallow broad sinus and continuing forwards to just above the peripheral frill where it turns gently be- fore continuing backwards obliquely across the edge of the peripheral frill and the base; below sinus a broad anterior projection vari- ably developed above the periphery; anterior projection may continue further forward than the contact of the outer lip with the upper su- ture; thick secondary deposits developed later in periphery; growth lines typically closely spaced and foliaceous on both the outer whorl surface and base; no other sculpture de- veloped. Dimensions: Ht Wt Hap Wap Wh P47479 7 — — — — P47480 17 38 — — 4+ P47482 12 31 4 8 5+ P47483 16 47 — — 4+ P47485 24 56 — — 5+ P47486 20 45 — — 6+ P47489 19 32 — — 5+ Location of Types: National Museum of Vic- toria. Holotype P47480. Paratypes P47481, P47482, P47502, P47498. Collected C. B. Tassell 1975. Material: Holotype, 4 paratypes, 1 hypotype and more than 90 other specimens all of which are crusted, distorted or fragmented. Discussion: Both B. angulatus and O. murrayi occur together in abundance in poorly sorted coquinas within the Bell Point Limestone. In each instance many of the shells have been broken and fragmented. However, the two species can be readily distinguished by the dif- ferences in their overall shape, nature of their sutures and whorl peripheries, the degree of development of the parietal lips and the posi- tion of the anterior projection beneath the sinus on the outer lip. In addition an umbili- cus is frequently developed in O. murrayi. Family EOTOMARIIDAE Wenz, 1938 Subfamily EOTOMARIINAE Wenz, 1938 Genus Mourlonia de Koninck, 1883 Type Species: Helix carinatus J. Sowerby, 1812; Lower Carboniferous; Mountain Lime- stone, near Settle, Yorkshire, Great Britain. ? Mourlonia sp. (Plate 1, figure 9) Description: Small to medium gastropod with prominent raised selenizone bordered by two BELL POINT LIMESTONE GASTEROPODA 27 strong spiral cords at whorl periphery; seleni- zone located at about mid-whorl; basal and apertural regions unknown; from upper suture to selenizone collabral lines prosocyrt; beneath selenizone prosocyrt initially; regular collabral lines strongly developed above and below selenizone; no spiral sculpture developed. Dimensions: Ht wt wh P46975 — 14 4 Location of Types: National Museum of Vic- toria. Figured specimen P46975. Collected C B. Tassell 1975. Material: Figured specimen. Discussion: The incomplete preservation of the specimen precludes its confident assignment to the genus. The specimen from Bell Point differs considerably from M. carinata (J. Sow- erby). Above the selenizone of the larger type species the fine closely spaced collabral lines are weakly prosocyrt to prosocline in contrast to the strong prosocyrt lines of the Bell Point form. The selenizone of the type species with its weaker bordering spiral cords is located lower on the whorl than that of the Bell Point form. The larger M. striata (J. Sowerby), type spe- cies of Ptychomphalina synonymized with the genus Mourlonia by Knight et al. (1960) has strongly developed collabral lines above and below a selenizone located high on the whorl in later growth stages. The selenizone is bor- dered by prominent spiral cords. The colla- bral lines, position of the selenizone and the cords bordering the selenizone are similar to those developed on the Bell Point form. M. talboti (Dickens) described by Batten (1972) from the Permian of Malaya is similar in size to the Bell Point form. The Malayan species also has strong collabral lines and a selenizone bordered by threads located higher on the whorl surface. Family HOLOPEIDA Wenz, 1938 Subfamily GYRONEMATINAE Knight, 1956 Genus Gyronema Ulrich, 1897 Type Species Trochonema (Gyronema) pul- chellum Ulrich and Scofield, 1897; Middle Or- dovician; Chatfield, Minnesota, U.S.A. Range: Ordovician to Middle Devonian. Distribution: North America, Europe, Africa and Australia. The occurrence of a species of this genus at Bell Point extends the generic range to include Australia. Gyronema bellense sp. nov. (Plate 1, figures 10, 11) Diagnosis: Typical form of genus with numer- ous elements of spiral sculpture of only one order and thin inner and outer lips. Description: Medium, turbiniform gastropod with numerous elements of spiral sculpture; whorls rounded between impressed sutures; periphery probably at mid-whorl; base rounded; small variably developed umbilicus sometimes present; aperture circular; no aper- tural emargination present; columellar lip slightly thickened; thin peripheral inductura developed; outer lip moderately thin; straight and oblique backwards from the upper suture; fine to slightly foliaceous, prosocline collabral lines present; sculpture composed of regularly spaced spiral cords. Dimensions: Ht Wt Hap Wap Wh C P46941 — 15 — 4 — 7+ P46945 16 15 3+ 10+ P46954 — 57 28 25 — 10+ Location of Types: National Museum of Vic- toria. Holotype P46945. Paratype, P46941. Collected C. B. Tassell 1975. Material: Holotype, paratype and 1 3 other specimens which are all crushed or broken. Discussion: The holotype is a crushed incom- plete specimen. Some apertural details are pre- served as are numerous spiral sculptural ele- ments and evidence of its turbiniform shape. The paratype shows clearly the straight colla- bral lines. The species from the Bell Point Limestone is in many ways similar to G. pulchellum. Both possess thin lips, fine growth lines, a similar number of strongly developed cords and pro- minent concave interspaces between the cords, although in a number of specimens from Bell Point the development of the prominent cords has been enhanced by subsequent weathering. ci 28 C. B. TASSELL However, G. bellense can be quite clearly dis- tinguished from the type species by its very much larger size and differing development of the cords. In terms of size G. bellense is comparable with 'Cyclonema' australis Etheridge from the Lilydale Limestone. G. bellense can be dis- tinguished from this species by its thinner lips, lack of growth rugae, single order of cords and relatively fewer cords. Family MURCHISONIIDAE Koken, 1896 Genus Murchisonia D'Archiac and DeVerneuil, 1841 Subgenus Murchisonia (Murchisonia) D'Archiac and DeVerneuil, 1841 Type Species: Muricites turbinatus Schlotheim, 1820; Middle Devonian; Stringocephalus lime- stone, near Gladbach in Bergischen, Germany. Murchisonia (Murchisonia) bassensis sp. nov. (Plate 1, figure 4) Diagnosis: Small form of subgenus with well rounded whorl profile selenizone in upper half of whorl; prominent, well-developed collabral lines. Description: Small, high-spired numerous whorled gastropod with a selenizone bordered by two fine cords above the rounded mid- whorl periphery; whorl rounded; impressed sutures moderately deep; base rounded; lacking um- bilicus aperture sub-circular; gently arcuate columellar lip of moderate thickness; no sug- gestion of canal-like constriction at junction of columellar and outer lips; parietal inductura thin; outer lip with angular sinus that forms a slit which generates the selenizone; depth of angular sinus increases with growth; from the upper suture to the selenizone the nearly straight outer lip passes posteriorly with only a slight obliquity; below the selenizone it passes forwards to the edge of the base before con- tinuing backwards across the base; flat seleni- zone located above mid- whorl periphery; with collabral lines becoming finer but remain strongly developed; no sculpture. Dimensions: Ht Wt Hap Wap Wh P46969a 4 1 7 P46969b — — 0-5 05 — P46971 4 1*5 — — 1 P46970 a 3 1 — — 8 P46970b — 15 0-6 1 — Location of Types: National Museum of Vic- toria. Holotype P46969a. Paratype P46969b. Collected C. B. Tassell 1975. Material: Holotype, paratype and at least 30 other specimens. Discussion: The principal differences between M. (M.) bilineata (Dechen), the type species, and this species are the latter's small size, well rounded whorl profile and selenizone located above mid-whorl. The species from the Bell Point Limestone also possesses more strongly developed collabral lines and weaker spiral cords bordering the selenizone than the type species. A number of other similar sized members of the subgenus are known from Victoria. In most cases they are inadequately known because of poor preservation or lack of specimens. As a result any comparisons that can be made are limited. M . (Murchisonia) bassensis can be dis- tinguished from a number of undescribed mur- chisoniacians from the Lilydale Limestone by its well-rounded whorl profile, strong collabral lines and possession of a selenizone located above mid-whorl. All the specimens from Lilydale possess prominent cords bordering the distinctly concave selenizone in contrast to the subdued cords of M . (M.) bassensis. Chapman (1912) mentioned the presence of a small member of the genus in the Buchan Limestone. From thin sections it was seen to have an angular whorl profile and an umbili- cus. Other details, notably the presence of a selenizone, were obtained from a poorly pre- served specimen P 12873. Typically the genus Murchisonia lacks an umbilicus. Whether the specimens in the thin section were the same as the P12873 is open to question, in any case, poor preservation precludes comparison with M. (Murchisonia) bassensis. BELL POINT LIMESTONE GASTEROPODA 29 Murchisonia (Murchisonia) sp. A. (Plate 1, figure 5) Description: Medium, high spired gastropod with a selenizone on a slightly raised narrow keel at the angular periphery; periphery just above mid- whorl; whorl face flat to slightly concave above and below the selenizone; su- tures moderately deep; base rounded; umbilical region unknown; columellar lip unknown; mod- erately thick outer lip with deep angular sinus that forms a slit at the periphery which gene- rates a selenizone from the upper suture to the selenizone the prosocyrt outer lip passes pos- teriorly with a strong obliquity; below the sele- nizone it passes forwards to the base less strongly; collabral lines fine; no sculptural ele- ments present. Dimensions: P46596 P46957 Ht 17 18 Wt 7 Wh 7+ 8+ Location of Types: National Museum of Vic- toria. Figured specimen P46956. Collected CB.Tasselll975. Material: Figured specimen and 6 other speci- mens which are all crushed or broken. Discussion: The poor preservation of the speci- mens limits comparison. The form from the Bell Point Limestone differs from the type species principally in having a selenizone raised upon a keel on a more angular whorl peri- phery. In this respect it is very similar to M. (Murchisonia) gracilicrista Linsley from the Middle Devonian Anderdon Limestone, Michi- gan. However, M. (Murchisonia) gracilicrista has a mid-whorl peripheral selenizone. The only other form of similar age and shape known from Australia is M. (Murchi- sonia) turris described by de Koninck (1876) from the Yass District. Study of material from the Taemas limestones reveals some differences between M. (Murchisonia) turris and the spe- cies from Bell Point. M. (Murchisonia) turris has a more rounded whorl profile with a con- cave selenizone located on keel at the peri- phery. From the upper suture the outer lip does not pass backwards as strongly as in M. (Murchisonia) sp. A. Murchisonia (Murchisonia) sp. B (Plate 1, figure 3) Description: Small, many whorled, high spired gastropod with a selenizone bordered by two prominent coarse cords at the sub-angular mid-whorl periphery; whorl profile sub-angu- lar; whorl surfaces flat above and below seleni- zone; impressed sutures; base rounded; lacking umbilicus; gently arcuate columellar lip of moderate thickness; parietal inductura thin or absent; flat selenizone located at or just below mid-whorl; no sculpture or collabral lines present. Dimensions: P46976 a P46976 b P46977 P46978 P46980 Ht 4 7 11 3-6 4 Wt 2 4 5 Hap Wap 2-5 Wh 5 6 7 5 6 Location of Types: National Museum of Vic- toria. Figured specimen P46976. Collected C. B. Tassell 1975. Material: Figured specimen and at least 30 other specimens. Discussion: M. (Murchisonia) bilineata (De- chen), the type species, is considerably larger than M. (Murchisonia) sp. B, from the Bell Point Limestone. Both species possess a dis- tinctive selenizone bordered by two prominent spiral cords at an angular mid-whorl peri- phery. These features distinguish them from M. (Murchisonia) bassensis which occurs at Bell Point. The poor preservation of the Bell Point form precludes more detailed comparison be- tween it and the type species as also with some undescribed forms from the Lilydale Limestone, similar in size, shape, whorl profile and selenizone development. Subgenus Murchisonia (Hormotomina) Grabau and Shimer, 1909. Type Species Murchisonia maia Hall, 1861; Middle Devonian; Columbus limestone, near Dublin, Ohio, U.S.A. Murchisonia (Hormotomina) sp. Description: Medium high spired gastropod with a median spiral thread on mid-whorl sele- ct 30 C. B. TASSELL nizone; whorl profile rounded; impressed su- tures; base unknown; apertural details poorly known outer lip thick with a shallow concave slit that generates a slightly elevated selenizone bordered on each side by prominent cords; sculpture absent. Dimensions; Ht 48 Wh 8 P46968 Location of Types: National Museum of Vic- toria. Mentioned specimen P46968. Collected C.B. Tassell 1975. Material: Mentioned specimen. Discussion: The poor preservation of the speci- men from Bell Point Limestone and the type species prevents effective comparison of these similar sized forms. The only similar described form from Victoria is Gyrodoma etheridgei (Cresswell) from the Lilydale Limestone. This large high-spired gastropod also has a me- dium spiral thread developed on a mid-whorl selenizone in some instances. However, this species is characterized by the presence of nu- merous fine spiral sculptural elements of two or more orders. High-spired gastropod gen. et sp. indet. (Plate 1, figure 2) Description: Small to medium cyrtoconoid gas- tropod; numerous whorls with gently convex whorl profile between shallow impressed su- tures; no other details preserved. Dimensions: Ht 7 Wt 4 Wh 6 P46974 Location of Types: National Museum of Vic- toria. Figured specimen, P46974. Type Locality: Waratah Limestone about 100 m west of Bird Rocks, near Walkerville South. Stratigraphic Range: The Waratah Limestone is considered by Strusz (1972) to be Pragian. Material: Figured specimen and one other specimen. Discussion: Michelia sp. described by Talent and Philip (1956) from the similar-aged Toon- gabbie Limestone, Marble Creek is similar to the form from the Waratah Limestone. Talent (as cited by Singleton, 1968) included Coelocaulus in a faunal list for the Waratah Limestone. This genus had been synonymized with Michelia by Knight et al. (1960). Thus it is possible that the unidentified form may be a member of the genus Michelia. Acknowledgements I wish to thank Mr T. A. Darragh of the National Museum of Victoria, Melbourne for his discussion and criticism of this work; Dr O. P. Singleton and Dr P. J. Coleman for their helpful comments, criticisms and dis- cussions; Margaret Tassell for the photographs and discussion throughout this study. References Batten, R. L., 1966. The Lower Carboniferous gas- tropod fauna from the Hotwells Limestone of Compton Martin, Somerset, pt. 1. Palaeont. Soc. 509: 1-52. , 1972. Permian gastropods and chitons from Perak, Malaysia. Bull Amer. Mus. Nat. Hist. 147(1): 5-44. Brown, T. A., 1941. The Stratigraphy and structure of the Silurian and Devonian rocks of Yass-Bown- ing district, New South Wales. /. Proc. R. Soc. N.S.W., 74: 312-341. Chapman, F., 1912. Reports on fossils — Middle Devonian of the Buchan district. Rec. geol. Surv. Vict. 3(2): 218-222. Etheridce, R., 1898. Record of Fossils. Rep. Prog. geol. Surv. Vict. 9: 125-126. , 1921. Palaeontologia Novae Cambriae Meridional is — Occasional descriptions of New South Wales fossils — No. 8. Rec. geol. Surv. N.S.W. 10(1): 1-11. Heidecker, E., 1959. Middle Devonian molluscs from the Burdekin Formation of North Queensland. Pap. Dep. Geol. Univ. Qd. 5(2): 3-8. Hill, D., 1954. Devonian corals from Waratah Bay, Victoria. Proc. R. Soc. Vict., 66: 105-118. Knight, J. B., 1934. The gastropods of the St. Louis, Missouri Pennsylvanian outlier: vii. The Euom- phalidae and Platyceratidae. /. Paleont. 8(2): 139-166. Knight, J. B., 1945. Some new genera of Paleozoic gastropoda. Ibid, 19(6): 573-587. Knight, J. B., R. L. Batten and E. L. Yochelson, 1960. Descriptions of Paleozoic Gastropoda In Moore, R. C. (Ed.), Treatise on Invertebrate Paleontology: Part I. Mollusca (1), Univ. Kan- sas Press. BELL POINT LIMESTONE GASTEROPODA 31 Koninck, L. G. de, 1876. Recherches sur les Fossiles Paleozoiques de da Nouvelle — Galles du Sud (Australie). Mem. Soc. Roy. Sci. Liege. 2, 6. (Translated, 1898, as: Descriptions of the Pal- aeozoic fossils of New South Wales (Australia). Mem. geol. Surv. N.S.W., Palaeont. 6.). Krommelbein, K., 1954. Devonische Ostracoden aus der Gegend von Buchan und von der Kuste der Waratah Bay, Victoria, Australien. Sencken- bergiana 35: 119-229. Lindner, A. W., 1953. The Geology of the coastline of Waratah Bay between Walkerville and Cape Liptrap. Proc. R. Soc. Vict. 64: 77-92. Link, A. G., 1970. Age and correlations of the Siluro- Devonian strata in the Yass Basin, New South Wales. /. geol. Soc, Aust. 16(2) : 711-722. McCoy, F., 1877. Record of Fossils. Rep. Prog. geol. Surv. Vict. 4: 156. McGhee, G. R., 1976. Late Devonian benthic marine communities of the central Appalachian Alleg- heny Front. Lethaia., 9: 111-136. Pedder, A. E. H., 1965. A revision of the Australian Devonian corals previously referred to Micto- phyllum. Proc. R. Soc. Vict., 78(2) : 201-220. , 1966. The Upper Devonian gastropod Orecopia in western Canada. Palaeontology 9(1): 142-147. Philip, G. M. and A. E. H. Pedder, 1967. A corre- lation of some Devonian limestones of New South Wales and Victoria. Geol. Mag. 104: 232- 239. Roberts, J., P. J. Jones and E. C. Druce, 1967. Palaeontology and correlations of the Upper Devonian of the Bonaparte Gulf Basin, Western Australia and Northern Territory. In Oswald, D. H., Ed. International Symposium on the De- vonian System, Calgary, 1967. 2 vols. Alberta Society of Petroleum Geologists, Calgary. Singleton, O. P., 1968. Geology of South Gippsland. In McAndrew, J., and Marsden, M. A. H. (Eds.), Regional guide to Victorian geology, School of Geology, University of Melbourne, Publication No. 1. Strusz, D. L., 1972. Correlation of the Lower De- vonian rocks of Australasia. /. geol. Soc. Aust. 18(4): 427-455. Talent, J. A., 1956. Devonian brachiopods and pele- cypods of the Buchan Caves Limestone, Vic- toria. Proc. R. Soc. Vict,, 68: 1-56. -, 1963. The Devonian of the Mitchell and Wentworth Rivers. Mem. geol. Surv. Vict. 24: 1-118.78 pi. 1965. The Stratigraphic and diastrophic evolution of central and eastern Victoria in middle Palaeozoic times. Proc. R. Soc. Vict. 79: 179-195. Talent, J. A. and G. M. Philip, 1956. Siluro-De- vonian Mollusca from Marble Creek, Thomson River, Victoria. Ibid. 68: 57-71. Tassell, C. B„ 1976. A Revision of the Gastropod fauna of the Lilydale Limestone (Early Devon- ian) of Victoria. Mem. natn. Mus. Vict. 37: 1-22. Teichert, C, 1954. Note on Devonian Limestones between the Bluff and Bird Rock, Waratah Bay, Victoria. Proc. R. Soc. Vict. 66: 103-104. Ulrich, G. H. F., 1875. Observations on the Waratah Bay Limestone. Rep. Prog. geol. Surv. Vict. 2; 125-126. Vandenberg, A. G. M., M. J. Garratt and D. Spen- cer-Jones, 1976 Silurian-Middle Devonian In Douglas, J. G. and Ferguson, J. A. (Eds.), Geology of Victoria. Spec. Publ. geol. Soc. Aust. 5. Voisey, A. H. and G. H. Packham, 1969. Sediments of the Tamworth Group. In Packham, G. H. (Ed.), The Geology of New South Wales. A geol. Soc. Aust. 16(1): 1-654. Yochelson, E. L., 1956. Permian Gastropoda of the south-western United States: I. Euomphalacea, Trochonematacea, etc. Bull, Amer. Mus. Nat. Hist. 110: 173-260. Explanation of Plate PLATE 1 Figure 1 — Tropidodiscus waratahensis, P46963, ho- lotype, x3. Figure 2 — High-spired gastropod gen. et sp. indet., P46974, x3. Figure 3 — Murchisonia (Murchisonia) sp. B., P46976, x5. Figure 4 — Murchisonia (Murchisonia) bassensis, P46969, holotype, x4. The aperture vis- ible is of another specimen of this species. Figure 5 — Murch ison ia (Murch ison ia) sp . A . , P46956, x2. Figure 6—Orecopia murrayi, P46990, paratype, xH. Basal view. Figure 7 — Bassotrochus angulatus, P47502, para- type, x2. Figure 8 — Orecopia murrayi, P46987, holotype, xl. Figure 9 — ? Mourlonia sp., P46975, xli. Figure 10 — Gyronema bellense, P46941, paratype, x2. Basal view. Figure 11 — Gyronema bellense, P46945, holotype, xl. Figure 12 — Orecopia murrayi, P46988, paratype, xl. Basal view. Figure 13 — Orecopia murrayi, P46987, holotype, xl. Figure 14 — Bassotrochus angulatus, P47498, para- type, xl. Figure 15 — Straparollus (Euomphalus) sp., P47503, xl. Basal view. Figure 16 — Austerum carinatum Heidecker, P47513, xl. Fanning River district, Queensland. Figure 17 — Bassotrochus angulatus, P47482, paratype, xli. Basal view. 32 C. B. TASSELL Figure 1 $—Bassotrochus angulatus, P4748 1 , paratype, Figure 21— Straparollus (Euotnphalus) sp. P475 1 1 , x l- xr,. Basal view. Figure 19— A usterum carinatum Heidecker, P47514, Figure 22— Bassotrochus angulatus, P47480, holo- xl. Fanning River district, Queensland. type, xl. Figure 20— Micromphalus clarkei (de Koninck), Figure 23— Micromphalus clarkei (de Koninck), P47517, xf. Derringullen Creek near P47519, x$. Basal view. Derringullen Yass, N.S.W. Creek, near Yass, N.S.W. MEM. NAT. MUS. VICT. 39 PLATE THE OMPHALOCIRRIDAE: A NEW FAMILY OF PALAEOZOIC GASTROPODA WHICH EXHIBITS SEXUAL DIMORPHISM By Robert M. Linsley Department of Geology, Colgate University, Hamilton. New York, USA Abstract A careful study of the gastropod Hypomphalocirms rugosits (gen. et sp. nov.) from the Middle Devonian of Northeastern Michigan demonstrated that in this population two distinct morphotypes are present. Both morphotypes persisted throughout the entire sixty feet of the limestone and over fifty miles of lateral distribution. This suggests that they might possibly be male and female dimorphs. Tn order to check this as a possible explanation, re- lated species were studied from Germany, Manitoba and Australia. In all cases two morpho- types were present in an approximate one-to-one ratio, but only in Australia was a large enough sample present to be of significance (one hundred specimens from Australia, com- pared to a dozen specimens from Germany and Manitoba). This study has resulted in the erection of a new family (the Omphalocirridae), one new genus {Hypomphalocirms), one resurrected genus {Liomplwlus, Chapman) and one new species {Hypompltalocimts rugosus). The genus ArctamphaLus Toemachov, 1926, is sup- pressed as a synonym of Omphalocirrus. The discovery of multispiral opercula in place in three of the genera demonstrate conclusively that these snails are dextral rather than sinistral as has frequently been suggested. The functional significance of the dimorphism is considered a mode of life established for the organisms and the probable evolutionary sequence suggested. Sexual Dimorphism in Gastropoda difference by itself will be unrecognizable in Sexual dimorphism of shell shape, size and fossil populations. However if there are mor- radular form are all well-known phenomena Phologic differences whtch allow one to sort among living archaeogastropods and caenogas- the population into two samples, then pre- tropods (Baker, 1926; Lamy, 1937; Robert- sumably a size distinction could be made. son, 1971; Sohl, 1969). Most commonly the In the case of the Omphalocirridae a variety difference takes the form of a larger average of criteria were discovered which allowed the (or maximum) size for the female. Differences mature members of the population to be in form generally are expressed in a greater divided into two subsets. The most ubiquitous rate of expansion of the generating curve in distinction is the type of coiling present. All the female which usually results in a relatively members of this family are characterized by lower spired trochoid shell (a 'fatter' shell), discoidal shells, but mature members can nor- Both of these differences are generally thought mally be designated either as orthostrophic or to be adaptive to accommodate the bulky eggs hypestrophic. Other features that may be used produced by the female. Radular differences to differentiate the two subsets are the degree may reflect food differences as in the case of of development of collabral ornamentation, the the archaeogastropod Tricolia (Hiloa) variabil- degree of development of a keel or spines, the is (Pease) where the female feeds preferentially roundness of the whorl profile, and the inclma- on brown alga (Padina) while the male lives on tion of the outer whorl face. Sorting of the top of the female's shell preceding and during population on the basis of the above features, the mating season, and presumably the radular either singly or in combination, makes it pos- differences in the male allow it to feed on the sible to show that these subsets also exhibit epiphytes on the female's shell (Robertson, a distinction of the size of the individuals in 1971, p. 77). tnese subsets. It is assumed in this paper that Obviously radular differences will be of no the subsets of the population containing the help in sexing fossil gastropods. Similarly size largest individuals represent the females. 33 34 ROBERT M. LINSLEY Systematic Paleontology Class GASTROPODA Cuvier, 1797 Subclass PROSOBRANCHIA Milne Edwards, 1848 Order ARCHAEOGASTROPODA Thiele, 1925 Suborder MACLUR1TJNA Cox and Knight, 1960 Superfamily EUOMPHALACEA dc Koninck, 1881 Family Omphalocirridae new family Type genus Omphalocirrus, Ryckholt, 1860 Description: Large, dextral, discoidal gastro- pods varying from gently orthostrophic to gently hypertrophic; whorl profile varying from subcircular to subtriangular, frequently with a marked circumbilical keel or spines on base; base broadly phaneromphalous; aper- ture subcircular. Early whorls filled with septa at maturity. Operculum disc-shaped, multi- spiral. All known members exhibiting varying degrees of sexual dimorphism. Discussion: The Omphalocirridae are readily distinguished from the Helicotomidae and Omphalotrochidae in being discoidal, while most members of the latter two families pos- sess elevated spires. The Omphalocirridae also differ from these two families in having an abundance of septa filling the early whorls. The Omphalocirridae resemble the Euomphalidae in these two characters of septation and discoi- dal form, but are distinguished from the Euom- phalidae and all other Euomphalacea in hav- ing a calcareous, multispiral operculum and exhibiting sexual dimorphism. As is true of all Euomphalacea the Ompha- locirridae possess an aperture whose plane passes through the axis of coiling Cradial aper- ture'), and some of the genera possess an an- gulation near the upper whorl face that is in- terpreted as exhalent. They resemble the Om- phalotrochidae in that some members of both groups possess a second angulation near the base that resembles a siphonal canal or an in- halent angulation. The uniqueness of the form of these angulations suggests that they were developed independently in each of the two groups. In each of these groups this combina- tion of features suggest that only a single gill exists (Knight, Batten and Yochelson, 1960, p. 1196) and supports the contention that this is the case for the entire superfamily. The Omphalocirridae also resemble the gas- tropods presently placed in superfamily Orio- stomatacea (Suborder Thochina, Knight, Bat- ten and Yochelson, 1960, p. 1245) in that both groups have radial apertures and multispiral, calcareous opercula. It is probable that the Oriostomatacca should be transferred to the Euomphalacea and, if it could be demon- strated that they exhibit sexual dimorphism, to the Omphalocirridae. Genus HYPOMPHALOCIRRUS gen nov. Type species: H, rugosus sp. nov. Description: Large, dextral gastropods with spire very low to hyperstrophically depressed; whorl profile subtriangular with flattened up- per, outer and lower faces joining at marked angulations; base broadly phaneromphalous; the juncture between the outer whorl face and the base typically developing canal-like projec- tions which may also occur at juncture be- tween upper and outer whorl faces. Earlier whorls filled with septa at maturity. Opercu- lum disc-shaped, multispiral. Sexual dimor- phism strongly developed. Discussion: The most abundant and obvious fossil in the Rogers City Limestone is the large euomphalid gastropod previously known as Omphalocirrus manitohensis Whiteaves (Eh- Iers and Radabaugh, 1938; Ehlers and Kes- ling, 1970). Close study of about 200 speci- mens from the collections of the University of Michigan Museum of Paleontology, the U.S. National Museum and Colgate University show that this euomphalid not only is not conspeciiic with Whiteaves species but that both the Rogers City material and the material from Manitoba are sufficiently distinct from Ompha- locirrus goldfttssi to warrant the establishment of a new genus (also see Yochelson, 1966, p. 41). This genus is most readily characterized by the subtriangular whorl profile and the angular meeting of the outer whorl face with the base and upper whorl face. It is obviously closely OMPHALOCIRRIDAE 35 related to Omphalocimis but this latter genus has a rounded whorl profile. Both of these genera possess the unusual feature of canal- like projections at the juncture of the base and outer whorl face. In addition it bears close similarity to the Australian genus Liomphalus, even though the latter lacks the canal-like pro- tuberances. Two species are assigned to this genus. One (H. rugosus) is herein described for the first time, while the second is Whiteaves species H. manitobensis (Whiteaves) from the Winni- pegosis Limestone of Manitoba, Canada. In addition the specimen described from the An- derdon Limestone of the Michigan Basin (Lin- sley, 1968) probably also belongs to this genus, but it is too poorly known to describe formally. Hy omphalocimis rugosus sp. nov. (Plates 2-6) Herein designated type species. Omphalocirrus manitobensis Ehlers and Rada- baugh 1938 Omphalocimis manitobensis Anonymous Omphalocirrus manitobensis Ehlers and Kes- ling 1970 Description: Large, discoidal, dextral gastro- pods with sub-triangular whorl profile, with angular base bearing canal-like projections or a keel. Nuclear whorls smooth, simple, dextral. Whorl profile sub-triangular in shape with outer whorl face varying from vertical (parallel to axis of coiling) to inclined outward at a 45° angle. Upper whorl face varying from flat and horizontal to roundly dipping into spire; outer whorl face flattened to broadly arched, basal whorl face gently rounding into umbili- cus. Junction of upper and outer whorl faces joining at an acute angle forming a smooth keel or distinctly undulating keel. Junction of outer and lower whorl faces typically sharp, occasionally rounded and bearing a series of canal-like projections or flanges which are sometimes replaced by a single or double keel in the adult. Upper suture varying from indis- tinct to very sharp and deep with considerable overhang of ultimate whorl; umbilical suture sharp and deeply incised, occurring just out- side of the umbilical ridge of the preceding whorl. Upper shell surface flattened in youth- ful stages, but varying in the adult from flat- tened to a considerably depressed spire. Shell widely phaneromphalous with umbilical angle varying from 120° to 180°, depending on the degree of hyperstrophism. Growth lines on upper whorl face generally opisthocline, begin- ning perpendicular to suture and culminating at a 45° angle at the periphery; growth lines prosocyrt on outer whorl face, continuing onto base with backward obliquity and backward concavity; normal course of growth line de- flected both by undulations on shoulder and canal-like projections on the base. Ornament consisting of strong collabral ornamentation in youth, giving way to simple growth lines at maturity; in mature forms, interruptions in growth frequently resulting in rugose appear- ance; outer whorl face sometimes developing roughened appearance in adult, even develop- ing low canal-like projections. Shell very thick at junctures of the three whorl faces, thin at centres of each whorl face, resulting in a subcircular inner whorl profile and a subtri- angular outer whorl profile. Septa abundant and evenly spaced in the early whorls of ma- ture individuals. Operculum circular, disc- shaped multispiral, expanding counterclock- wise, all volutions visible on outer surface; upper surface of each opercular volution in- clined inward with slight rounding towards preceding whorl; outer face of opercular whorl parallel to axis of coil, resulting in deep, sharp suture separating each volution; early opercu- lar volutions (first ten or eleven) increasing in size logarithmically; later volutions (last five or ten) more irregular, approximately maintaining constant size and thus increasing in an archi- median rather than logarithmic spiral; inner surface of operculum poorly known. Holotype measuring 102-6 mm in width and 36-4 mm in height. Largest Paratype measur- ing 149 mm in width. Holotype— UMMP 22377; Figured Paratypes— UMMP 22375, 22378, 22379, 22380, 22383, 22384, 57888, 57889. USNM 102939, 213754, 213755 213756, 213757, 213758, 213759, 213760, 213761, 213762, 213763, 213764, 213765, 213766, 213767, 213768, 213769, 213770, 213772, 213773, 213774, 213775, 213776, 213777, 213778, 213779, 213780, 213781, 213782, 213783. Unfigured Paratopes— UMMP 22381, 22382, 22385. USNM 213771. 36 ROBERT M. LINSLEY Discussion: Hypompiialocirrus rugosis is a very large gastropod (150 mm) that rivals the size of the largest Macluritida. Among Paleozoic Gastropoda only Straparollus grandis (Ko- ninck ( 1 30 mm ) Pithodea amplissa, Ko- ninck (140 mm) from the carboniferous of Belgium, Arctomphalus grandis Tolmachotf (now Omphalocirrus goldfussi) (170 mm) and a Carboniferous euomphalid (210 mm) re- ported by Yochelson (1966) as being in the British Geological Survey and Museum can compare in size with this new species. The diagnostic features of this species are the sub-triangular whorl profile, the tendency to develop flanges at the upper angulation of the female, and canal-like projections on the base. However there is considerable variation of each of these features within the population of about two hundred individuals that were studied. Females of H. nigosus can be distinguished from females of H. manitobensis (Whiteaves) by the presence of a second row of canal-like projections which the latter bears on the upper angulation. The whorl profile of H. manito- bensis has the triangular profile tilted so that the outer whorl face is subparallel to the axis of coiling and the periphery of the shell is at the midpoint of the rounded whorl face rather than at the upper angulation as it is in H. rugo- sus. H. manitobensis has assumed an almost isotrophic coiling and thus appears to converge on a bellerophont-like form. The single, poorly preserved specimen of Hypompiialocirrus described from the Ander- don Limestone (Linsley, 1968) is not well enough known to warrant a formal description. The chief distinction is the bundling of growth lines found in this species. Five or six growth lines are subparallel to each other and all participate in making up a single basal projec- tion. The next set of prominent growth lines then diverge sharply from the preceding set to form the next basal projection. Within the population of females of H. rugosus the degree of hyperstrophism is still highly variable ranging from USNM 213781 (pi. 6, fig. 5) where the upper surface of the penultimate whorl strikes the ultimate whorl about one-third the way down the ultimate whorl, to UMMP 213758 (pi. 4, fig. 7) where the upper surface of the ultimate whorl strikes the ultimate whorl two-thirds the way down the ultimate whorl. The geometry of the whorl profile is such that variation in the amount of hyperstrophism also is associated with other changes in the whorl profile. As the degree of hyperstrophism increases the entire triangular whorl profile of the shell rotates so that the outer whorl face varies from being inclined almost 45° to the axis of coiling in the male (see USNM 213775, pi. 2, fig. 4) to almost parallel to the axis of coiling in the female (see UMMP 22377, pi. 4, fig. 2). The round- ness of the upper whorl face also seems to in- crease with increased hyperstrophism (UMMP 22377 and USNM 213758, pi. 4, figs. 6, 7) and of course umbilical depth decreases as hy- perstrophism increases. UMMP 22377 (pi. 4, figs. 5, 6) has an umbilical angle of 180° while USNM 213759 (pi. 6, fig. 3) has an umbilical angle of about 150° and USNM 213758 (pi. 4, fig. 7) is so depressed that one could talk about an 'umbilical spire'. Other features, equally variable, do not seem to be related to the degree of hyper- strophism. The proportionate size of the outer whorl face may be rather small (UMMP 22379, pi. 6, fig. 8) or very great USNM 213759 (pi. 6, fig. 3). Some features are ob- viously related to age. Immature forms of males are very difficult and sometimes impos- sible to distinguish from those of females. Im- mature forms tend to have well-developed col- labral ornamentation (UMMP 22375, pi. 5, fig. 7, early whorls of USNM 213760, pi. 5,' fig. 3). Immature specimens (UMMP 22375, pi. 5, fig. 7) also tend to have a smooth junc- ture between the upper and outer whorl faces. Rarely this feature will be carried into maturity in the females (UMMP 57888, pi. 5, fig. 4 and USNM 213781, pi. 6, fig. 5). More fre- quently this upper juncture develops into a scalloped flange at maturity, either with a gentle fluting (UMMP 22379, pi. 5, fig. 6 and pi. 6, fig. 1) or in a few cases very pronounced, almost grotesque fluting (USNM 213772, pi. 6, fig. 4 and USNM 213754, pi. 6, figs. 10, 11). In the holotype (UMMP 22377, pi. 4, fig. 6) the ultimate whorl is well fluted until the last OMPHALOCIRRIDAE 37 i volution, when the fluting ceases. There seems to be some tendency for the degree of fluting to be related to the amount of sculpture present on the outer whorl face. In general the greater the fluting, the more sculpture. The sculpture varies tremendously from raised growth lines (USNM 213754, pi. 6, fig. 11) to small ridges perpendicular to the growth lines (USNM 213759, pi. 6, fig. 3) large bumps (USNM 213765, pi. 6, fig. 7) or even canal- like foldings of the growth lines (USNM 213772, pi. 6, fig. 4). The canal-like projections on the basal an- gulation are a most persistent feature of the females. Their position on the base is depen- dent upon the amount of hyperstrophism and the subsequent rotation of subtriangular whorl profile. Where hyperstrophism is not great and outward rotation of the outer whorl face mini- mal, the projections will appear to be in the middle of the whorl as seen from the base (UMMP 22378, pi. 6; fig. 2 and USNM 213760, pi. 5, fig. 3). When hyperstrophism is strongly developed and outward rotation of the outer whorl face is substantial (UMMP 22377, pi. 4, fig. 5) the projections will appear to be at the periphery of the whorl as seen from the base. The shape of these projections varies from the typical rather slender ones (UMMP 22377, pi. 4, figs. 5, 8 and USNM 213764, pi. 5, fig. 2) to very broad, spade-like projections (USNM 213767, pi. 5, fig. 5). One specimen, UMMP 22380 (pi. 6, fig. 9) is the most aberrant of the collection. It has a very swollen, rounded whorl profile and thus seems to closely resemble Omphalocirrus goldfussi. In all other respects, such as the sculptured, outwardly rotated outer whorl face, etc. it resembles H. rugosus and I have thus treated merely as one aberrant member of a very diverse population. The males of Hypomphalocirrus rugosus may be readily distinguished from the fe- males by their flat upper whorl face and by the fact that the spire is flush, with the upper whorl faces continuous with each other rather than the depressed spire so typical in the fe- males. The males exhibit less variation than the females. The spire profile is very constant, maintaining an essentially flat upper surface. The outer whorl face occasionally shows a faint, revolving ornamental groove just below the periphery (USNM 213779, pi. 3, fig. 4, USNM 213773, pi. 3, fig. 6 and USNM 213778, pi. 3, fig. 9). The outer whorl face is generally flattened, but a few individuals, es- pecially USNM 213761 (pi. 3, fig. 12) and USNM 213756 (pi. 3, fig. 13) exhibit a very rounded whorl profile. The most marked varia- tion in the males occurs on the base relative to the circumbilical ridge. The ridge develops very early at the end of the second volution. During this early stage canal-like projections develop very much like those of the females. However in the males the projections rarely persist beyond the third whorl. An exception is USNM 213773 (pi. 3, fig. 6) which maintains very large, fully developed spines through ma- turity. In most mature individuals (USNM 213775, pi. 3, fig. 8 and USNM 213766, pi. 3, fig. 5) the mature whorls have only a rela- tively sharp ridge, perhaps with only faint suggestions of protrusions (USNM 102939, pi. 3, figs. 1, 11). In at least one example (USNM 213757, pi. 3, fig. 3) a double ridge results. In one other case (USNM 213756, pi. 3, fig. 13) there is no sign of a keel at all, but rather the base is very rounded and the growth lines are unusually rugose at maturity. Dr G. Arthur Cooper of the U.S. National Museum kindly brought to my attention a collection of gastropods from the Miami Bend Formation of Indiana (Cooper and Phelan, 1966). Most of the collection consists of stein- kerns which are generically and specifically indeterminate, but a few molds were collected which can be identified. One of these (USNM 213769, pi. 2, fig. 6) is a partially complete im- pression of the spire of a male Hypomphalo- cirrus rugosus. This adds one more small piece of evidence to their mass of data supporting their claim for a correlation of the Miami Bend Formation with the Rogers City Lime- stone. The wide latitude of variation present in this species is in part explained by the morpholo- gical differences between the males and fe- males. The existence of these two subsets of this population can best be seen in the graph 38 ROBERT M. LINSLEY in Fig. 1 , which plots size against a measure of the degree of hyperstrophism (umbilical width divided by umbilical depth). This plot demon- strates that though the youthful individuals are inseparable, the adults separate into two dis- tinct populations. This is, of course, perfectly consistent with an interpretation of sexual di- morphism. Using the degree of hyperstrophism as a criterion for establishing two populations, many other distinctions were discovered which correlate with this character. The follow- ing paragraphs are formalized descriptions of the females and males. Description of females: Large, shallowly hy- pertrophic gastropods with well-developed canal-like projections at the junction of the outer whorl face with the base. Nucleus smooth, simple and dextral. Whorl profile subtriangu- lar in shape, outer whorl face tilted in slightly from perpendicular. Upper whorl face flattened near periphery, gradually curving downward near preceding whorls and curving downward abruptly at suture, so that inner edge of upper whorl face overhangs upper whorl face of preceding whorl; upper whorl face adjoining outer whorl face at periphery with a marked keel; keel smooth in youthful stages, but at maturity keel tending to develop pronounced periodic undulations; outer whorl face gently rounded, sloping inward and downward from peripherally located junction with upper whorl face; outer whorl face adjoining base forming a keel which develops canal-like projections at a very early stage; canal-like projections gene- rally becoming larger and better developed with maturity; basal whorl face flattened, proceed- ing upward towards umbilical suture from its junction with outer whorl face. Upper suture fairly deep, sharply incised with ultimate whorl slightly overhanging penultimate whorl; um- bilical suture sharp and very deep, occurring outside of the canal-like projections of pre- ceding whorl. Early whorls depressed below mature whorls resulting in hyperstrophic whorl profile. Degree of hyperstrophism quite vari- able, but generally increasing with maturity. Shell widely phaneromphalous, with umbilical angle varying from 150°to 180° depending on degree of hyperstrophism. Growth lines op- isthocline on upper whorl face, beginning per- pendicular to suture and proceeding with a forward obliquity and marked backward con- cavity; at periphery growth lines variable de- pending on the position relative to flange; pre- sence of flange causing growth line to bend backward from its opisthocline course, mak- ing it subparallel to suture once again; growth lines continuing on outer whorl face in proso- cyrt fashion; at junction of outer whorl face and base, growth lines interfered with by canal-like projections; growth lines bending back rather strongly in a U-shape to partici- pate in formation of spine, bending forward in an V shape in front of completed spine, continuing uninterruptedly where no spine is present. Ornament consisting of collabral orna- ment in youth, giving way to growth lines in adult; growth lines becoming crowded during spine formation causing adult to have rugose appearance; flanges or undulations developing along periphery in adult; canal-like projections around base tending to become very strong in adult; outer whorl face tending to become roughened in adult, developing low, rugose, and in extreme cases, even canal-like projec- tions. Shell thickness exceedingly variable, be- ing thin at centre of each whorl face and be- coming very thick at junctures of whorl faces, resulting in a subtriangular outer whorl pro- file and subcircular to elliptical inner whorl profile. Septa abundant in earlier whorls in mature shells, unevenly spaced at about eight to twelve per whorl. Holotype measuring 102-6 mm in width and 36-4 mm in height. Largest paratype measur- ing 149 mm in width. Holotvpe — XJMMP 22377; Figured Paratvpes — UMMP 22375, 22378, 22379, 22380, 22383. USNM 213760, 213764, 213758, 213754, 213765, 213772, 213776, 213755, 213768, 213767. Unfigured Para- types— UMMP 2238 J, 22382, 22385. Description of males: Large, flat-topped ortho- strophic gastropods with a subtriangular whorl profile and sharp angular base bearing either a keel or canal-like projections. Nuclear whorls smooth, simple, dextral. Whorl profile re- sembling an equilateral triangle with upper whorl face horizontal or perpendicular to axis of coiling. Upper whorl face flat, bending OMPHALOCIRRIDAE 39 downward very slightly at the surface. Outer whorl face joining upper whorl face at a sharp angle at shell periphery, forming a sharp smooth keel. Outer whorl profile flattened to gently rounded; typically with slight concavities just below peripheral keel and just above ba- sal keel, occasionally gently rounded from keel to keel. Basal whorl face joining outer whorl at sharp angulation and progressing flatly or with gently rounding to umbilical su- ture. Upper suture varying from indistinct to sharp with slight overhang of inner whorl edge over preceding whorl; umbilical suture sharp, frequently deep, normally just outside circumbilical ridge. Upper shell surface flat- tened, not depressed. Base widely and deeply phaneromphalous; umbilical angle varying be- tween 120° to 145°. Growth lines on upper whorl face beginning perpendicular to the su- ture and proceeding opisthoclinally towards the periphery, intersecting periphery at 45° angle; growth lines prosocyrt on outer whorl face, continuing with strong collabral orna- mentation, normally reducing to faint growth lines in adult; strong angular keel at periphery at junction of upper and outer whorl faces; faint revolving groove sometimes developed on outer whorl face, just below periphery. Cir- cumbilical ridge developing early with well- developed canal-like projections; in maturity, circumbilical ridge normally reverting to single or double ridge, rarely retaining canal-like pro- jections to maturity. Shell thick, particularly at keels, resulting in subtriangular outer whorl profile and subcircular to elliptical inner whorl profile. Septa present through much of early portion of mature shell. Shell width of largest paratype 110 mm. Androtvpe—USNM 102939; Paratypes—UMMP 22382, 22384. USNM 213775, 213770, 213762, 213763, 213777, 213757, 213778, 213779, 213773, 213766, 213774, 213769, 213761, 213756, 213780. Un figured Paratype — 213771. Discussion continued; Briefly, the main fea- tures of each sex can be summed as follows. The females tend to be larger than the males, a feature similar to so many modern gastro- pods and presumably related to the greater bulk of eggs of the female. The canal-like pro- jections on the base of the males usually de- generate into a keel while in the female these projections become larger in the adults, pre- sumably remaining functional throughout its life. Assuming that these do indeed facilitate in- current water, this would again be consistent with the increased oxygen needs of the female in producing the bulky egg masses. In addition the increased hypcrstrophicity of the female would appear to be due to, or at least accom- panied by, the rotation of the whorl profile. This rotation brings the outer whorl face into a sub-vertical position (parallel to the axis of coiling) thus bringing both incurrent and excurrent openings into a peripheral position which would again facilitate water movement through the mantle. The male, with less critical circulatory demands tends to keep the in- current opening at a slightly less favourable position in order to better position the anus at the extreme periphery. The male also tends to loose the incurrent canals, which were ap- parently useful during the rapid growth of youth, but no longer so crucial to success in the adult. It is possible that the collabral ornamenta- tion of the immature shells served to strengthen those shells. As the shell increased in size it also increases in thickness, apparently to the point where the collabral ornamentation lost its functional significance and so disappears on the adults. Hypomphalocirms manitobensis (Whiteaves) Euomphalus manitobensis Whiteaves 1890, p. 100, pi. 6, figs. 2-26. Omphalocirrus manitobensis Whiteaves 1892, p. 327, pi. 43, figs. 5-7. (Plate 7, figures 1-8) Description: Large, slightly hyperstrophic gas- tropods with one or two rows of canal-like extensions at boundaries of outer whorl face. Nucleus poorly known, apparently smooth and dextral. Whorl profile varying from subtri- angular in the females, to suboval in shape in the males. In females the upper whorl face meeting outer whorl face to form a keel which may bear long canal-like projections, then proceeds roundly down to suture; outer whorl face of females broadly rounded 40 ROBERT M. LINSLEY and meeting lower whorl face at a keel which also may bear canal-like protuberances similar to those on top; basal whorl face continuing roundly up to the umbilical suture to complete the rounded equilateral triangle. Whorl profile of males more rounded with less distinct keels, bearing canal-like projections only on base. Upper suture deep, sharply incised. Umbilical suture distinct, deeper in females than in males. Aperture, as determined from observations of growth lines, gently opisthocline on upper whorl face, bending back at periphery to form upper rows of canal-like projections, continu- ing on outer whorl face in prosocyrt fashion and on to the base with a gently backward con- cavity, again involved with canal formation at the juncture of the base with the outer whorl face. Males with greater hypcrstrophism than females; base of males being relatively flat while females deviating only slightly from dis- coidal. Ornament consisting of strong collabral ornament in males with one row of canal-like projections on the base; ornament in females consisting only of fine growth lines with two rows of canal-like projections both above and below the outer whorl face. Outer whorl face sometimes roughened in adult females. Shell moderately thick, particularly at junc- tions of whorl faces. Septa fairly abundant in mature individuals. Operculum thin, and disc- shaped, circular in outline, multispiral, ex- panding counterclockwise with all volutions visible on outer surface. Lectotype—GSC 4173; Picsiotypes—GSC 4163a, 4174, 4175, 4176, 4177; Ifvpotvpes—USNM 213782, 213783. Discussion: Yochelson (1966, p. 41) has al- ready treated the problems attendant on this species. Whiteaves (1890, p. 100) original specimens of H. manitohensis are steinkerns from the Dawson Bay Limestone. These are generically indeterminate. Subsequently, Whiteaves (1892, p. 327, pi. 43, figs. 5-7) re- described this species and reassigned it to the genus Omphalocirms, but this was done prim- arily on the basis of material obtained from the Winnipegosan Limestone which underlies the Dawson Bay Limestone. There is no cer- tain way of knowing whether or not the speci- mens from these two units are conspecific. The above description is based on material from the Winnepegosan Limestone as our knowledge of Hypomphalocirrus manitohensis at this time is based entirely on those specimens. The collection available for study of this species consists of fifteen specimens collected by Linsley and Cottrell in 1968 from locality 1 8 of the Field Guide to Devonian Outcrops of Southwestern Manitoba (McCabe, 1967). This locality is about one mile west of the Narrows, on Lake Winnepegosis. In addition, Whiteaves types (GSC 4174, 4176, 4177) from the Geo- logical Survey of Canada at Ottawa were kindly made available for study. While seven- teen specimens do not constitute a large enough collection on which one might base a definitive study, they do suggest that two distinct mor- photypes are present and may best be regarded as males and females. The females (pi. 7, figs. 3, 5, 6, 8) are characterized by a triangular whorl profile and have well-developed canal-like protrusions at both boundaries of the outer whorl face. As this form is regarded as having evolved from //. rug&sus, it would seem a natural develop- ment for the undulating shoulder of the Rogers City females to have developed into the true canals of the Manitoba form, thus perfecting the current flow in and out of the mantle cavity. The females of H. manitohensis also resemble the females of H. rugosus in having a sub- vertical outer whorl face, a fair degree of hy- perstrophicity, and rugosities on the outer whorl face. The males of the Winnepegosis Formation differ markedly from the Rogers City forms. The males of //. manitohensis are more hyper- strophic than the females which suggests that the degree of hyperstrophism is important only as it affects the position of the incurrent or ex- current water streams. The basal canal-like ex- tensions on the males of H. manitohensis are positioned only slightly more peripherally than their H. rugosus counterparts. The rest of the whorl profile of the H. manitohensis males is much more rounded than either the females or the Rogers City forms. However they resemble the males of the Rogers City forms by having much stronger collabral ornament than that possessed by the females. OMPHALOCIRRIDAE 41 Omphalocirrus goldfussi (Archiac and Verneuil) — (Plate 8) Knight (1941, pp. 45, 213) and Yochelson (1966, p. 42) have both offered excellent des- criptions and synonymies of Omphalocirrus goldfussi; (Archiac and Verneuil, 1842) and Arctomphalus grandis Tolmachoff, 1926. I am familiar with this genus only through their works and through a careful study of the plastotype material made available to me from the US National Museum. This study has led me to the conclusion that 'Arctomphalus gran- dis' is the sexual dimorph (probably male) of Omphalocirrus goldfussi. Plate 8, fig. 7 is an illustration of the lectotype of Arctomphalus grandis (Specimen A-19229, Paleontologisk Museum of Oslo). Unfortunately the outer volution, represented by the broken line in the figure, is a steinkern and only an impression of the basal suture of the inner whorls remains. However comparison of this remnant with some German material of Omphalocirrus gold- fussi (see pi. 8, figs. 4, 5, 9) illustrates that all of these specimens are similar in that they are hyperstrophic, have strong collabral ornament and have a circumbilical ridge which develops periodic nodes which are only occasionally de- veloped into canal-like projections. Collec- tively, these features suggest male characters of Hypomphalocirrus rugosus and H. manito- bensis. The other types of Omphalocirrus gold- fussi (PL 8, figs. 1-3, 6, 8) are quite distinct from those described above. They are all or- thostrophic, have much weaker collabral orna- mentation and have well-developed, apparently functional canal-like extensions which appear in early youth and persist through maturity. In addition this morphotype has a very pro- nounced swollen form due to a greater rate of expansion of the generating curve. All of these features are consistent with the interpre- tation of this form as the female. To date no opercula have been found asso- ciated with O. goldfussi, but I fully expect multispiralled opercula to occur in the same beds and with great fortune to be found in a life situation, well back from the aperture. It would also be beneficial if a study could be made on a large population of this species to ascertain whether or not the dimorphism ap- parent in the half-dozen specimens at my dis- posal holds true for the entire population. Genus Liomphalus Chapman, 1916, p. 90 Type species: Liomphalus australis by original designation Description: Large, discoidal, septate gastro- pods with low to hyperstrophically depressed spire. Outer whorl profile varying from sub- circular to subtriangular with circular inner whorl profile. Strong circumbilical ridge with no canal-like projections. Operculum disc- shaped, multispiral. Sexual dimorphism weakly developed. Discussion: Like the other members of the family Omphalocirridae, this genus exhibits sexual dimorphism. However compared to the other genera, the sexual dimorphism of Liom- phalus seems poorly developed, as though it were an incipient feature. This feature is con- sistent with its lower stratigraphic position and presumed ancestry to the other members of the family. Liomphalus also resembles the other members of this family by being septate, dis- coidal gastropods which bear a disc-shaped, multispiral calcareous operculum (see also Yochelson and Linsley, 1972). It can readily be differentiated from other members of this family by the presence of a circumbilical ridge without any suggestion of nodes or canal- like extensions on it. Liomphalus northi (Etheridge) 1890 Plate 9, 10 1890 Oriostoma northi Etheridge, p. 64, pi. 9, figs. 6-7 1894 Oriostoma northi Etheridge, p. 151, pi. 9, fiss. 1-4 1894 Euomphalus (Oriostoma) northi Etheridge; Cresswell, p. 157 1913 Euomphalus northi (Etheridge); Chapman, p. 227 1916 Euomphalus northi (Etheridge); Chapman, p. 90 1916 Liomphalus australis Chapman, p. 90, pi. 4, figs. 32-33 1959 Straparollus (Euomphalus) northi (Etheridge); Philip and Talent p. 50, pi. 7, figs. 1-21, pi. 8, figs. 1-2 1972 Oriostoma northi Etheridge; Yochelson and Lin- sley, p. 8, pi. 1, fig. 6, pi. 2, figs. 1-5 1976 Straparollus (Euomphalus) northi Etheridge; Tassell, p. 9, pi. 1, figs. 7, 8, pi. 2, fig. 11, pi. 3, figs. 1,2,7,8 42 ROBERT M. LINSLEY Description: Shells large, discoidal, varying from flatly orthostrophic to slightly hypertro- phic with distinct circumbilical ridge. Nuclear whorls unknown, presumably simple, smooth and dextral. Whorl profile subrectangular in youthful stages, changing to subelliptical or subtriangular in adults. Upper whorl face vari- able, ranging from broadly arched and con- fluent with upper whorl face, to gently arched and meeting outer whorl face of a distinct keel; marked keel separating upper and outer whorl faces typically present in immature forms, but disappearing in many members of the adult population; outer whorl face strongly rounded, joined lower whorl face at pronounced circum- bilical ridge; lower whorl face with slight con- cavity inside circumbilical ridge, continuing flatly to umbilical suture. Both upper and um- bilical sutures shallow but distinct; depth of sutures varying with degree of spire depression with deeper suture generally present on flattest surface. Shells varying from slightly ortho- strophic to slightly hypertrophic, sometimes bordering on isostrophic, particularly in imma- ture shells. Growth lines orthocline on upper whorl face, opisthocyrt on outer whorl face and slightly prosocline on lower whorl face, opisthocyrt on outer whorl face and slightly prosocline on lower whorl face with gently prosocyrt bendings over keels when present. Ornament of youthful forms consisting of strong collabral lines and two keels above and below the outer whorl face; in adult forms the basal collabral ornamentation and keel invari- ably persist, but the upper collabral ornamen- tation gives way to fine, closely spaced growth lines in one-half the population; the keel sep- arating the upper and outer whorl face fre- quently disappearing in one-half the adult population. Shell thinnest near parietal lip, sometimes present only as a parietal inductura; shell becoming thicker near outer lip and reaching maximum thickness (6 or 7 mm) at circumbilical ridge. Septa abundant in early portion of whorls of adult shells, becoming more widely and unevenly spaced in more mature whorls; may occasionally be placed within one-half whorls distance from the aper- ture. Opiculum thick, circular, disc-shaped, multispiral, expanding counterclockwise. Shell width of largest hypotype measuring 98 mm. Figured Hvpotypes—NMV PI 107, P28373, P28498, P28499, P28707, P28708, P287Q9, P28710, P287I1, P28712, P28713, P28714, P287I5, P287J6, P28717, P287l8and P28719. Description of males: Shell large, discoidal, slightly hyperstrophic. Nucleus unknown, pre- sumably normal dextral. Whorl profile varying from subelliptical to distinctly subtriangular, typically with marked bilateral symmetry con- verging to isostrophism. Upper whorl face typically gently arched to the keel-like shoul- der separating upper and outer whorl faces; keel-like shoulder occasionally absent with upper and outer whorl faces roundly confluent with one another. Outer whorl face rather strongly convex between upper keel-like shoulder and circumbilical ridge; lower whorl face flattened to umbilical shoulder. Upper suture distinct but not sharply incised; um- bilical suture slightly deeper, also sharply in- cised. Shell basically hyperstrophic but vari- able in degree. Growth lines orthocline on up- per whorl face, opisthocyrt on outer whorl face and slightly prosocline on lower whorl face with gently prosocyrt areas over each keel. Ornament normally consisting of strong colla- bral ornament and two keels. Normally with strong collabral ornament on upper and lower whorl faces with weaker collabral ornament occasionally absent on upper and outer whorl faces. Normally with keel-like shoulder separating upper from outer whorl faces and with circumbilical ridge separating outer and basal whorl faces; upper ridge occasionally ab- sent. Septa common in early whorls, more rare in mature whorls, but present to within one- half volution of aperture. Operculum as in orthostrophic individuals. Shell width of largest hypotype measuring 70 mm. Ifypotvpes—NMV PJJ07, P28707 t P28710, P28713, P28714, P28715, P28717, P28718 } P28719 t P28708. Description of females: Shell large, discoidal with depressed spire. Nucleus unknown. Whorl profile subelliptical with long axis of ellipse perpendicular to axis of coiling, upper whorl face broadly rounded, usually continuing without interruption to outer whorl face; outer whorl face more sharply rounded. Joining OMPHALOCIRRIDAE 43 lower whorl face at marked circumbilical keel; lower whorl face with slight concavity inside circumbilical ridge, then flattened before rounding into umbilical suture. Upper suture shallow but distinct; umbilical suture more shallow but also distinct. Shells basically or- thostrophic tending towards planispiral, apical angle varying from 200° to 230°. Umbilical angle varying from about 130° to 160°. Growth lines closely spaced and very fine on upper whorl face, generally orthocline becom- ing very slightly opisthocline on outer whorl face and very slightly prosocline on base re- sulting in a slight prosocyrt phase over cir- cumbilical ridge. Growth lines becoming strong collabral ornamentation as they cross circumbilical ridge onto base. Ornamentation consisting of growth lines on upper and outer whorl faces, collabral ornamentation on base, with circumbilical ridge invariably present and shoulder separating upper and outer whorl faces only rarely present. Shell thinnest near parietal lip, sometimes present only as parie- tal inductura, occasionally with some degree of thickness to it. Shell becoming thicker (up to four or five mm) near outer lip and reaching maximum thickness at circumbilical ridge. Septa numerous through early portion of shell, becoming more widely spaced in mature region, but occasionally present to within one- half whorl of the aperture. Operculum thick, circular, disc-shaped, multispiral, expanding counterclockwise (for more complete descrip- tion, see Yochelson and Linsley, 1972). Shell width of largest hypotype measuring 98 mm. Hypotypes—NMV P28498, P28499, P28373, P28709, P28711, P28712 and P28716. Discussion: Chapman based the genus Liom- phalus on some very poorly preserved speci- mens from the Lilydale Limestone which are undoubtedly abraded specimens of Etheridge's species 'Oriostoma northi'. As a result his original species name of Liomphalus australis Chapman is invalid and should be Liomphalus northi (Etheridge). However, the generic name 'Liomphalus' is still a valid and available name, and since the new evidence presented in this paper supports the removal of this species from either Oriostoma or Straparollus 1 have utilized the genus Liomphalus. This species can be easily differentiated from other members of the Omphalocirridae by the fact that its base lacks any nodal swellings or canal-like extensions. In addition, the sexual dimorphism while present, is not as well de- veloped as in the descendant, more advanced members of the family. The adult males of L. northi tend to preserve juvenile features such as collabral ornamentation on the upper whorl face in addition to a strong keel separating the upper and outer whorl faces. In addition the males develop only a modest degree of hyper- strophism. The upper surface of the females however lose both the keel and the collabral ornament so that their upper whorl face flows continuously into the outer whorl face and has only closely-spaced, fine growth lines on it. The females also tend to be slightly ortho- strophic. However none of these sets of characters are inviolate. In the collection of 104 specimens available of the National Mu- seum of Victoria, Melbourne, there were oc- casional hypcrstrophic forms with a rounded upper whorl face (PI. 10, fig. 12, NMV P28713) or a faint keel which faded out (PI. 10, figs. 9, 10), but the great majority of the members of this large population could be separated with ease^ However, this incipient sexual dimorphism, the lack of canal-like projections and the low stratigraphic position of L. northi are all con- sistent with the interpretation that this form is ancestral to Omphalocirrus and Hypomphalo- cirrus. Stratigraphic Distribution The oldest member of the family Omphalo- cirridae is Liomphalus northi of the Lilydale Limestone, Lower Devonian Yeringian of Australia. It is considered to be late Siegenian byStrusz(1972). Although the next known occurrence of a member of this family is delayed until the Anderdon Limestone of the Michigan Basin, it is probable that further examination of gastro- pods presently assigned to the family Euom- phalidae will uncover other genera and species that should be transferred to this group. The 44 ROBERT M. LINSLEY Andcrdon form (Linsley, 1968) of Hypom- phalocirrus is unfortunately known only from a single fragment of an immature specimen. The Anderdon find represents the first known occurrence of this family in North America. The Anderdon represents a limited facics on the southeastern flank of the Michigan basin and is thought to be Eifelian (see Fagerstrom, 1961, and Linsley, 1968). It is considered to be late Sieyenian by Strusz (1972). The next occurrence of a member of this family is Hypomphalocirrus rugosus from the Rogers City Limestone of the North East cor- ner of the Michigan Basin. This unit has been placed in the lowermost Givetian (Cooper and Phelan, 1966, Ehlers and Kesling, 1970, Lins- ley 1973). Although this unit does not contain the brachiopod Sfringocephalus, it does contain other members of the Stringocephalus fauna such as Atrypa arctica, Subrensselandia n. sp., Liromytilus attenuatus, Buechelia tyrellii, Car- inatina dysmorphostrota (Ehlers and Kesling, 1970, p. 29) and Straparollus (Straparollus) cottrelli and S, (Eitomphalm) hofjmani (Lins- ley and Yochelson, 1973). The other occurrence of Hypomphalocirrus rugosus in the Miami Bend Formation of In- diana adds to the already substantial array of evidence put forth by Cooper and Phelan (1966) that this unit is directly correlative with the Rogers City Limestone. The Winnipegosis Formation of Manitoba is also generally considered to be of Lower Givetian age (Ehlers and Kesling, 1970, p. 29, Cooper and Phelan, 1966, p. 28, Baillie, 1951, p. 59) and is generally correlated by these authors as the equivalent of the Rogers City Limestone. However a study of the species of Hypomphalocirrus present in these two for- mations suggests that the Rogers City form is ancestral to the Winnipegosis form and thus must slightly antedate the latter. This view is also supported by the study of the middle Devonian carrier shell, Straparollus (Linsley and Yochelson, 1973) which suggests that the Rogers City species are ancestral to the Winni- pegosis species. Most certainly the two forma- tions are close in age to each other, but the evidence on hand suggests that the Rogers City-Miami Bend Formations are very slightly older than the Winnipegosis Formation. The last known occurrences of members of this family are those of Omphalocirrus gold- fussi from the Givetian Stringocephalus beds of Europe and the lone occurrence of the same species (ex-Arctomphalus grandis) from Goose Fiord, Ellesmere Land, from the Blue Fiord Formation of Eifelian age (McLaren, 1963, pp. 324-328). General Considerations The genus Omphalocirrus, to which Hypom- phalocirrus is closely related, has had a check- ered career in the literature, with no one being quite certain whether to place the canal-like protrusions on the 'top' or on the 'bottom'. Even within the past thirty years the beast has been flipped three times. Knight ( 1 94 1 , p. 213) considered the canal-like projections to be the base of the organism, but without much conviction for he was not at all certain 'that the orientation employed . . . should not be reversed' (Knight, 1941, p. 213). In 1952 (p. 37) he reversed it, indicating that he con- sidered Omphalocirrus to be a sinistrally coiled Macluritacean, and that the canal-like projec- tions served an exhalent function. This view was retained in the Treatise (Knight, Batten and Yochelson, 1960, p. 1189). In 1966, E. L. Yochelson (1966, p. 43, 45) broke from the Treatise triumvirate and flipped Omphalocirrus once again, placing the projections on the base, assuming them to be purely ornamental with no function, and transferring the genus from the Macluritacea to the Euomphalacea. In Whiteaves original description of H, manitohensis (Whiteaves, 1890, p. 100) he de- scribed a steinkern assigned to that species that had an operculum associated with it. Considerable doubt has always surrounded this association. The operculum in this fossil is positioned well back into the shell, away from the aperture and tilted at an angle that would be unusual for an intact operculum (pi. 7, fig. 2). It was impossible to demonstrate that this was not just an operculum that had floated into an empty shell. Added to this is the fact that Whiteaves original description of H. mani- OMPHALOCIRRIDAE 45 tobensis (Whiteaves, 1890) was based on stein- kerns from the Dawson Bay Formation that are generically indeterminate and our present understanding of the species is based on ma- terial collected from the underlying Winne- pegosis Formation and assigned by Whiteaves to H. manitobensis. To further complicate matters no opercula of a multispiral form were known either from the Euomphalacea or the Macluritacea, and the operculum ascribed by Whiteaves to H t manitobensis did not fit our understanding of either of these groups. I have been fortunate enough to find oper- cula in place both in H. manitobensis (pi. 7, fig. 1) from the Winnipegosis Formation of Mani- toba and in H. rugosus from the Rogers City Limestone of Michigan. In both instances the operculum is multispiral, set well back away from the aperture and slightly tilted to the axis of the cone of revolution of the whorl, al- though not as tilted as in Whiteaves specimen. Thus there can now be no doubt that Hypom- phalocirrus does indeed possess a multispiral operculum. The discovery also lends credence to Whiteaves claim that his steinkerns (one with a multispiral operculum) from the Daw- son Bay Formation are at least related to the material from the Winnipegosis Formation, if not actually conspecific. In both opercula as seen in place within the aperture, the volutions grow outward in a counter-clockwise fashion thus demonstrating conclusively that Hypomphalocirrus, and by in- ference Omphalocirrus, are dextral shells, and that the canal-like projections are really on the base of the shell. Thus the orientation of this enigmatic genus is no longer in doubt and it can finally come to rest, 'spines' down. However, the solution of the problem of the orientation of the shell of the Omphalocirridae, is only the solution of the orientation for our descriptive purposes. Saying that the beast has finally come to rest 'spines down' only relates to the orientation of figures on a plate for the purposes of comparative morphology and has nothing necessarily to do with the orientation of the shell to the substrate while the organism was living. It allows us to invoke the concepts 'dextral', 'sinistral', 'orthostrophic' and 'hyper- strophic' and to ascertain homologous por- tions of the shell (inhalent, exhalent, etc.). But the orientation of the shell during life remains as much of an enigma as ever. In the majority of modern gastropods, i.e. those with either high spires or elongated aper- tures (cones, mitres, volutes, terebras, etc.) the 'base' of the shell is actually the leading edge or anterior-most portion of the shell relative to the living animal. In low-spired modern gastropods (turbonids, naticids, trochids, etc.) or discoidal shells (planorbids, etc.) the 'outer lip' forms the leading edge of the shell. Thus in this second group the 'base' of the shell is the left side and the spire projects back and to the right side of the animal. In all of these gastro- pods the shell is held dorsally over the body by the columellar muscle and the plane of the aperture is parallel to the substrate. If the Omphalocirridae are 'typical' gastro- pods then the shell would be held in a dorsal position with the coiled portion resting on the operculum of the upper surface of the pos- terior portion of the foot (the metapodium). However the Omphalocirridae are unlike all of the above-mentioned gastropods in one im- portant aspect. In all of the aforementioned snails the plane of the aperture is tangent to the preceding whorls (a tangential aperture) while in the Omphalocirridae the plane of the aperture transects the axis of coiling of the shell (a radial aperture) with the result that a great portion of the earlier volutions projects beyond the plane of the aperture. This would seem disadvantageous for a gastropod whose aperture is parallel to the plane of the sub- strate. Perhaps the Omphalocirridae have a very different relationship to the substrate. In fact it has been suggested (Linsley and Yochel- son, 1973) on the basis of completely different criteria that some euomphalids [Euomphalus (Euomphalus), E. (Straparollus) and E. (Ser- pulospira)] lived base down with the plane of the aperture perpendicular to the substrate. In this position there would no longer be a need for the aperture to be tangent to the preceding whorls and indeed it seems to be a general character of most euomphalids to have the plane of the aperture intersect at or near the axis of coiling of the shell. 46 ROBERT M. LINSLEY This would be an adaptation suitable for a sedentary gastropod, perhaps a deposit feeder or even a filter feeder. The organism might possibly be capable of hoisting his shell into the normal position for short journeys to new feeding grounds. More probably the shell would just be dragged along during locomo- tion. It would be expected that the majority of the time would be spent with the shell lying flat on the substrate and the animal largely withdrawn inside the shell, protruding only to obtain food. For such a habit an operculum would be a distinct advantage. Whether the Omphalocirridae were filter feeders and assumed a living position of the shell lying flat on the substrate, or whether they were active grazers and thus carried the shell erect over their back is not known. Possibly the nature of their spines might afford some clue. During his investigation of the genera Om- phalocirrus and Arctomphalus, Yochelson (1966, p. 53) remarked that 'the fact that the spines develop gradually from nodes would seem to imply that they served no vital func- tion'. Close examination of these canal-like protuberances in Hypomphalocirrus belie that statement. As many as ten to twelve growth lines are involved in making one of the projec- tions, the last of which curve backward in a U-shaped fashion, and upward to form the rim of the canal-like projection (see pi. 4, figs. 5, 8). The next growth line does not bend up- ward but proceeds across the shell along the normal shell contour, bending forward at the position of the protuberance, thus leaving an opening ('tremata) which I suggest remains functional until the appearance of the next pro- jection, whereupon it is filled in with secondary calcite deposits, very much as are the tremata of Haliotis. However, these canal-like projections can- not be called 'tremata', because tremata (as found in Tremanotis, Polytremaria and the Haliotidae) are associated with the exhalent function and normally mark the position (or the eventual position) of the anus. In Hypom- phalocirrus the basal position of these projec- tions necessitates that we regard them as in- halent in function, thus resembling the inhalent (siphonal) canals of many of the modern Caenogastropoda and I propose that they be called 'inhalent tremata'. Hypomphalocirrus is not unique among Devonian gastropods in the development of a portion of the aperture for a specialized incurrent function. In the Raphistomatinae alone, both Tylozone and Buechella show well developed siphonal canals and the lower lip of Arlzonella most probably marks the position of the inhalent currents as does the plethospirid Diplozone. Even in the Euomphalidae, Pleuronotus and Diploconula have a basal angulation which would most probably result from a mantle fold which would make an incurrent stream more effective by being localized. It is probable that the advent of more ad- vanced and efficient groups during the Devon- ian, such as the Neritacea, the Palaeotrochacea and the Paleozygopleuridae, which would prob- ably be more efficient, increased the pressure on the older, more conservative groups such as the Macluritacea, Euomphalacea and Pleu- rotomariacea. One temporary solution could be in improvement of the efficiency of water cur- rents as evidenced by Omphalocirrus, Hypom- phalocirrus, Tylozone et ah It was a solution that apparently was not tried again until the Pennsylvanian when we find Knightites, Cyclio- scapha, and Straparollus (Amphiscapha) de- veloping similar canals or angulations. The primary water current in Hypomphalo- cirrus thus entered the mantle cavity by way of the incurrent canals on the base, passed the osphradium and left gill and then passed out of the mantle cavity, past the anus which was undoubtedly located at the angulation formed by the junction of the upper and outer whorl faces. In females of //. manitobensis this upper angulation is also equipped with a row of canal-like projections which mirror the form of those on the lower angulation, but which served an excurrent rather than an incurrent function. It is possible that the peripheral un- dulations in the female H. rugosus are antece- dent to this upper row of projections. If the interpretation of the current arrange- ment in Hypomphalocirrus given above is cor- rect, this adds support to the concept (Knight, OMPHALOCIRRIDAE 47 Batten, Yochelson, 1960) that the Macluritidae and the Euomphalidae suffered a reduction or elimination of the right gill, for there is no mor- phological evidence in the shell that a compar- ably effective water current would flow over the right gill which, in Hypomphalocirrus would have to be positioned under the upper whorl face. Therefore, if the spines are indeed func- tional as incurrent modifications, then the shell of the Omphalocirridae must have either been carried in an upright position or if lying flat on the substrate with the left side bearing the spines (the 'base') uppermost. This latter posi- tion would thus be 'upside down' compared with our normal way of depicting these shells. A Summary of Sexual Dimorphism and the Evolution of the Omphalocirridae Our present knowledge of this family would suggest that Liomphalus first developed the diagnostic features of this family and then gave rise to two stocks, one European culminating in Omphalocirrus and one North American, furnishing the Hypomphalocirrus lineage. The development of the operculum of this group has already been fully treated (Yochel- son and Linsley, 1972). Evolutionarily the thick operculum of Liomphalus northi is set at the apertural margin, while in Hypomphalocir- rus rugosus and H. manitobensis the operculum is much thinner and drawn well into the aper- ture, as far back as one-fourth volution. It also tends to be slightly rotated in this genus, which is a nice adaptation to allow a circular opercu- lum to fit snugly inside a slightly elliptical whorl profile. The operculum of Omphalo- cirrus goldfussi is unknown but most probably exists and has not been associated with that form. The varying reasons for the presence of septa in gastropods has also been discussed re- cently (Yochelson, 1971). In the case of the Omphalocirridae it seems unlikely that the septa are for strengthening the shell or for protection against breakage of the early whorls. Rather the septa would appear only to serve the purpose of body shortening. We do not know if this body shortening would result in any mass decrease. If the early chambers were filled with water, as is most probable, then pre- sumably the mass of water would closely ap- proximate the mass of flesh. Without a siph- uncle and the complex gas-exchange cells of the cephalopods it seems most unlikely that there would be any increase in buoyancy gained by the secretion of septa in gastropods. The features which are involved in the sexual dimorphism of this family, i.e. size differential, collabral ornamentation, differential develop- ment of keel or canals, and hyperstrophism, are all present in Liomphalus northi although they are not as fully differentiated in this species as they are in subsequent species. In Liompha- lus northi the males tend to be hyperstrophic, smaller, with collabral ornamentation on the upper whorl face and a keel separating the upper and outer whorl faces. The females are more orthostrophic, larger, and have no colla- bral ornamentation or keel on the upper whorl face. In all cases where I have had a sufficiently large population to study, I have assumed that the largest individuals are the females. The one exception to this rule is Omphalocirrus goldfussi. In the six examples I have seen from Germany, those that I have called fe- males are indeed the largest individuals. How- ever, this makes Tolmachoff's specimen from Ellesmere Land a male yet this specimen is the largest individual in the entire family. How- ever, as outlined below, I have other reasons for believing that this is a correct interpreta- tion, so until someone has had an opportunity to study a large population of this species I will continue to consider Tolmachoff's specimen as a male. In the Middle Devonian members of the family, the best indicator of sex is the rela- tive development of the canal-like extensions, the inhalent tremata. In the Rogers City form, the female has very well developed, obviously functional, canal-like inhalent tremata on the base. In addition the junction between the upper and outer whorl face has developed a series of undulations that are presumed to act as a canal, or at least a means of localizing the excurrent flow. The male generally lacks both of these traits. The females from the Winni- pegosis Limestone have developed canal-like 48 ROBERT M. LINSLEY extensions both on the top and bottom of the outer whorl face, while the males have inhalcnt tremata developed only on the base. This would seem an obvious improvement over the Rogers City Form and is the main reason that I consider Hypomphalocirrus rugosus ancestral to //. manitobensis. As mentioned earlier I believe the basal canals to be incurrent, while the upper canals or shoulder mark the posi- tion of the anus and is thus essentially excur- rcnt. It is presumed that the female needs better mantle circulation to enable it to pro- duce the more bulky egg masses. The males in both of these cases have less well developed aids to water circulation. In H. rugosus the adult males typically have only a keel where the incurrent canals of the female are located and have an even upper shoulder in place of the undulating shoulder of the female. The males of //. manitobensis have only a single set of canals at the incurrent position instead of the double set of both incurrent and excurrcnt found in the females. Thus the entire evolu- tion of the North American lineage was ac- companied by an improvement of water cir- culation in the mantle, with the females con- sistently better than the males. A similar situation exists with the European stock of Omphalocirrus goldfussi. In this species the female has very well developed in- halent canals whereas in the males, the canals arc not always well developed and are fre- quently only suggested by nodal swellings on the basal keel. The Australian representative deviates from the patterns set by its descendents in that the male of Liomphalus northi possesses keels above and below while the female has only the lower keel. However these keels have no ex- pression on the inner surface of the aperture and therefore may not be functional relative to water currents as are the well-developed keels of its descendents. I would suggest that this is merely a case of pre-adaptation, where the keels of the ancestral form were selected for one function (possibly strengthening the shell) and secondarily proved useful for a completely different function (isolating water currents). As a result it would not have the same selective pressures relative to the sexes in Liomphalus as it subsequently had in the more highly specialized offspring. The matter of hyperstrophism is also a per- plexing problem in the Omphalocirridae. In Liomphalus northi, Omphalocirrus goldfussi and Hypomphalocirrm manitobensis the males are more hyperstrophic than the females. This leads me to the conclusion that it is not the mode of coiling that is significant in itself, but the effect the coiling has on the positioning of the outer whorl face. In the genus Hypom- phalocirrus which has a triangular whorl face, the outer whorl face of female is oriented so that it is sub-parallel to the axis of the shell. This places both inhalent and exhalent portions of the aperture on the very periphery of the shell, both placed as far from the axis of coil- ing as possible. If the shell is positioned in a recumbent position, inhalent tremata up, then they are in the perfect position to obtain clean water while the anus of Hypomphalocirrus ru- gosus would be placed adjacent to the sub- strate, while excurrent canal of H. manitobensis would be thrust down into the substrate. Pos- sibly they buried their faces as does the modern detritus feeder Xenophora (Shank, 1969, p. 6). In the case of Omphalocirrus goldfussi and Liomphalus northi the whorl profile of the female tends to be more inflated than that of the male, presumably to accommodate the bulkier egg masses. The male, on the other hand, has a less inflated slightly more angular external whorl profile (the internal whorl cross section being subcircular). It is possible that these differences may dictate the hyperstrophic coiling of the males and the ortho-isostrophic coiling of the female. The final feature is one that is possessed by all members of this family and that is col- labral ornamentation in the adult males. In all members the immature forms have strong col- labral ornamentation, but only in the males is this juvenile feature carried into the adults. It is probable that this feature serves to strengthen the shell on juveniles and males. It is possible that this becomes unnecessary in females because they possess thicker shells, but I have no direct confirmation of this from OMPHALOCIRRIDAE 49 actual observations on shells (a particularly difficult task when ninety per cent of the fauna studied consists of latex impressions). Summary and Conclusions All four species of the Omphalocirridae, Omphalocirrus goldfussi, Liomphalus northi, Hypomphalocirrus rugosa and H. manitoben- sis exhibit considerable dimorphism in their adult shells. The expressed dimorphism is com- pletely consistent with an interpretation of sex- ual dimorphism in that both dimorphs are al- ways present in all localities where extensive collections have been made and in a more or less one-to-one ratio. All species except Liom- phalus northi are characterized by having func- tional inhalent tremata developed as canal-like extensions of the base. These inhalent tremata are consistently better developed in the mor- photypes that have been considered as females and it is suggested that this is related to the improved circulation necessary to aerate the bulky egg mass of the females. Acknowledgements This study has been made possible by a series of grants extended to the author by the Sloan Foundation and the Colgate Research Council. These grants were awarded for the summers of 1967, 1968, 1969, 1970 and for a sabbatical semester of 1971 which enabled me to go to Australia to study the fauna of the Lilydale Limestone as represented in the col- lections at the National Museum of Victoria, Melbourne. I am particularly indebted to Mr Edmund Gill and Mr Thomas Darragh of the National Museum of Victoria for opening their facilities and collections to me and for ex- tending the hospitality of the entire museum. I would also like to thank Dr Gary Batt of the Department of Zoology, University of Auckland for assisting with the photography of the Australian material. For German material of Omphalocirrus gold- fussi I am indebted to Dr Urlich Jux of the University of Cologne for taking the author to a number of collecting sites in the Paffrath area and making available comparative ma- terial for study. Dr Hugh R. McCabe of the Department of Mines and Natural Resources of the Province of Manitoba was most kind in discussing Mani- toba Geology with the author and in accom- panying the author to outcrops of the Winni- pegosis Formation. The author was first introduced to the Rogers City Formation of Michigan by Dr G. M. Ehlers and Dr R. V. Kestling of the University of Michigan Museum of Paleonto- logy. Both of these men have contributed enor- mously to the author's understanding of the geology of this area and to his understanding of the Rogers City Limestone in particular. I am deeply indebted for the many hours spent with them discussing both geological and bio- logical aspects of this problem. Access to the Calcite Quarry, Michigan Limestone Opera- tions, U.S. Steel Corporation at Rogers City, Michigan and to the Preque Isle Corporation Quarry, formerly Lake of the Woods Quarry, north of Alpena, Michigan, managed by Mr Roy Hutchinson for a consortium of steel com- panies, has always been graciously granted. I would also like to express my deep appre- ciation to four former students who have been of great assistance in this problem. Mr John Cottrell and Mr John Hoffman spent one summer with me under the auspices of a Col- gate Research Council Undergraduate Re- search Participation grant. They were inde- fatigable field workers and spent many hours preparing specimens and discussing various aspects of the problems. Cottrell was also able to accompany me to the Manitoba field area near Lake Winnipegosis. Dr Harold B. Rollins and Mr William Arendt also accom- panied me to the Rogers City area for field work and in addition Rollins was of inestimable value for the photographic work he did on the Rogers City material. Dr R. V. Kesling of the University of Michi- gan Museum of Paleontology, Dr G. Arthur Cooper of the U.S. National Museum and Dr Digby McLaren of the Canadian Geological Survey at Ottawa were very kind in giving the author access to the collections of their re- spective institutions. Lastly I greatly appreciate the many hours 50 ROBERT M. LINSLEY of stimulating discussion of this problem pro- vided by Dr John Morton, Department of Zoology, University of Auckland, Dr C. M Yonge, Department of Zoology, University of Edinburgh and Dr E. L. Yochelson, U.S. Geological Survey. Abbreviations used in this paper are as fol- lows: USNM — United States National Mu- seum, Washington, D.C.; UMMP — University of Michigan Museum of Paleontology, Ann Arbor, Michigan; GSC — Geological Survey of Canada, Ottawa, Ontario, Canada; NMV— National Museum of Victoria, Melbourne, Vic- toria, Australia. References Anonymous, 1960. Fossils found in Limestone of the Calcite Quarry of Michigan Limestone opera- tions of United States Steel Corp. at Rogers City, Michigan, U.S. Steel Corp. Archiac, E. J. A. d\ and E. P. de Verneutl, 1842. On the fossils of the older deposits in the Rhen- ish Provinces, preceded by a general survey of the fauna of the Palaeozoic rocks, and followed by a tabular list of the organic remains of the Devonian system in Europe. Trans, Geo!. Soc. Lond.,6; 303-410, pis. 25-37. Baillie, Andrew D„ 1951. Devonian geology of Lake Manitoba-Lake Winnipegosis area: Province of Manitoba, Dept. Mines & Nat. Resources, Mines Branch, publ. 49-2. Baker, H. B., 1926. 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Mas., 1, no 3, pp. 60-67. Fagerstrom, J. A., 1961. The Fauna of the Middle Devonian Formosa Reef Limestone of South- west Ontario, /. Paleont., 35, no. 1, pp. 1-48, pi. 1-14. Knight, J. Brooks, 1941. Paleozoic Gastropod Geno- types, Spec. Pap. Geol. Soc. America, 32, pp. 1-510, pis. 1-96. , 1952. Primitive Fossil Gastropods and their Bearing on Gastropod Evolution, Smith- son. Misc. Collns. 117, (13): 1-56, pi. 1-2. Batten, R. L., and Yochelson, E. L., 1960. Treatise on Invertebrate Paleontology (Moore, R. C„ editor) Part I, Mollusca 1. Geo- logical Society of America, Inc., Boulder, Colo- rado and University of Kansas, Lawrence, Kansas, xxiii and 351 pp. Koninck, L. G. de, 1881, Faune du calcaire car- bonifere de la Belgique, 3* partie, Gasteropodes: Mus. roy. H'tstoire nat. Belgique, Ann. sew paleont., 6, pp. 1-170, pi. 1-24 (Bruxelles). Lamy. E., 1937. Sur le Dimorphisme Sexuel des Coquilles, Journ. Conchyliol., 81, pp. 283-301. Linseey, Robert M., 1968. Gastropods of the Middle Devonian Anderdon Limestone, Bull. Amer. Paleont., 54, no. 244, pp. 333-465, pi. 25-39. , 1973. Paleoecological Interpretation of the Rogers City Limestone (Middle Devonian, Northeast Michigan), Contrib. Mus. Paleont. Univ. Michigan, 24, no. 11, pp. 119-123. and E. L. Yochelson, 1973. Devonian Carrier Shells (Euomphalidae) from North America and Germany U.S. Geo!. Surv. Prof. Paper 824, pp. 1-26, pi. 1-6. McCabe, H., 1967. Field Guide to Devonian Out- crops of Southwestern Manitoba. McLaren, D. J., 1963. Southwestern Ellesmere Is- land-Goose Fiord to Bjorne Peninsula, p. 310- 338 in Fortier, Y. A. and others: Geology of the North-Central part of the Arctic Archi- pelago, Northwest Territories (operation Frank- lin), Mem. Geol. Surv. Canada, 320. Milne Edwards, Henri, 1848. Note sur la classifica- tion naturelle des moilusques gasteropodes, Ann. Sci. Nat., ZooL, ser. 3, 9, pp. 102-112 (Paris). Philip, G. M. and L A. Talent, 1959. The gastro- pod genera Liomphalus Chapman and Scalaetro- chus Etheridge, /. Paleont. 33, pp. 50-54. Robertson, Robert, 1971. Sexually Dimorphic Archaeogastropods and Radulae, Annual Re- ports for 1970 of the American Malacological Union, pp. 75-78. Ryckholt, P. de, 1860. Revue des generes qui com- posent la famille des Haliotidae, D'Orbigny, Jour. Conchyliol., 2e ser. 4, pp. 183-188. Shank, Paul, 1969. The Timorous Carrier Shell, Close Observation of the Xenophora conchvlio- phora Born. New York Shell Club Notes 151 PP. 5-7. OMPHALOCIRR1DAE 51 Sohl, Norman F., 1969. Gastropod Dimorphism, in Westermann, G.E.G. editor, Sexual Dimorphism on Fossil Metazoa and Taxonomic Implications, Intern. Union of Geol. Sciences, Ser. A, 1, pp. 94-100. Strusz, D. L., 1972. Correlation of the Lower De- vonian Rocks of Australasia. /. Geol. Soc. Aits*. 18(4): 427-455. Tassell, C. B., 1976. A Revision of the Gastropod Fauna of the Lilydale Limestone (Early Devon- ian) of Victoria, Mem. Natn. Mus. Vic. 37: 1-21. Thiele, Johannes, 1925. Mollusca — Weichtiere: Handbuch der Zoologie, gegriindet W. Kuken- thal, 5, pp. 15-256 (Berlin, Leipzig). Tolmachoff, I. P., 1926. On the fossil faunas from Per Schie's Series D from Ellesmere Land with exception of brachiopods, corals, and cephalo* pods: Report of the second Norwegian Arctic expedition in the Tram' 1898-1902, 38, pp. 1-106, 8 pis. Whiteaves, J. F., 1890. Descriptions of some new or previously unrecorded species of fossils from the Devonian rocks of Manitoba, Trans. R. Soc. Canada, VIII, sec. 4, no. 3, pp. 93-110. , 1892. The fossils of the Devonian Rocks of the islands, shores, or immediate vicinity of Lakes Manitoba and Winnipegosis, GeoL Surv. Canada, Contrib. to Cana. Paleont., 1, pt. 4, no. 6, pp. 255-359, pis. 33-47, Ottawa. Yochelson, E. L., 1966. A Reinvestigation of the Middle Devonian gastropods Arctomphalus and Omphalocirrus, Arbok norsk Paralinst. 1965, pp. 37-48. , 1971. A new Upper Devonian gastropod and its bearing on the problems of open coiling and septation, Smithsonian Contr. Paleobiology, 3, pp. 231-241, pis. 1,2. and R. M. Linsley, 1972. Opercula of two gastropods from the Lilydale Limestone (Early Devonian) of Victoria, Australia, Mem. Natn. Mus. Vict. 33, pp. 1-14, pis. I-II. Explanation of Plates PLATE 2 Hypomphalocirrus rugosus n. sp. All figures are of males. All specimens except Figure 6 from the Rogers City Limestone. Figures 1-3 — Basal, apical and apertural views (xl-8) of paratype UMMP 22384, from the beach at Rockport Note flat upper sur- face, strong collabral ornamentation and 45° inclination of outer whorl face. Figure 4 — Apertural view of paratype USNM 213775 (xO-75) showing subtriangular whorl pro- file with flat upper whorl face and in- clined outer whorl face. Figure 5 — Apical view of paratype, UMMP 22383 (xl) exhibiting strengthening of collabral ornamentation near the suture. Figure 6 — Apical view (xll) of paratype USNM 213769 from the Miami Bend Formation in Ohio. Figure 7 — Apical view (xl-75) of paratype USNM 213770. In this specimen the collabral or- namentation is very strong in the youth- ful stage and becomes diminished with increasing age. Figure 8 — Apical view (x0-75) of paratype USNM 213762. Again the collabral ornament be- comes weaker with increased age. Figure 9 — Cross-section (xl-4) of paratype USNM 213763 showing numerous irregularly spaced septa. PLATE 3 Hypomphalocirrus rugosus n. sp. All figures are of males from the Rogers City Limestone. Figures 1, 11 — Oblique basal (xl-5) and basal (x0-8) views of Paratype, USNM 102939 show- ing flange-like projections of early whorls giving way to a continuous keel on the ultimate whorl. Figure 2 — Apertural view of Paratype USNM 213777 (xO-75). Figure 3- — Basal view (x0-65) of Paratype USNM 213757. Note continuous keel on ultimate whorl. Figure 4 — Adapertural view (0-6) of Paratype USNM 213779 showing slight suggestion of a spiral groove just below the shoulder. Figure 5 — Oblique basal view (xO-6) of Paratype USNM 213766 very large male with un- usually well developed flange-like projec- tions on the base. Figure 6 — Oblique basal view (x0-6) of Paratype USNM 213773 showing well developed spiral groove just below shoulder. Figure 7 — Basal view (x0-9) of Paratype USNM 213774. Figure 8— Apical view of Paratype USNM 213775 (x0-6) showing well developed colla- bral ornamentation. Figure 9 — Oblique basal view (xO-9) of Paratype USNM 213778. Note sharp single keel on base and smooth outer whorl face. Figure 10 — Basal view (xO-9) of Paratype UMMP 57889, a mature specimen with well- developed keel and deep umbilicus. Figure 12, 14 — Basal (xO-9) and oblique basal (xO-8) views of Paratype, USNM 213761. This specimen is unusual because of the round- ness of the base. Note also the well- developed flanges and the strong collabral ornamentation. Figure 13 — Basal view (xO-6) of mature specimen, Paratype USNM 213756 with a rounded base with no flanges or keel. 52 ROBERT M. LINSLEY Figure 15 — Basal view (xO-6) of Paratype USNM 213780. PLATE 4 Hypomphalocirrus rugosus n. gen. and sp. All figures are of females from the Rogers City Limestone. Figures 1,2, 5, 6, 8 — Oblique apical (xO 6), apertural (x0-8), basal (x0-8), apical (xO-8) and oblique basal (xO-8) of holotype UMMP 22377. A very large mature female show- ing well-developed basal flanges, undulat- ing shoulder, high degree of hyperstro- phism and rugose, near vertical outer whorl face, all typical features of the adult female. Figure 3 — Apical view (xO-6) of paratype UMMP 22378 showing smoothness of early whorls and the undulating shoulder. Figure 4 — Outer surface (xl-25) of operculum, para- type USNM 213768 showing the multi- spiral form typical of the opercula of this family. Figure 7 — Oblique apical view (xO-75) of Paratype USNM 213758, the most extremely hyper- strophic individual in the entire collec- tion. PLATE 5 Hypomphalocirrus rugosus n. gen. and sp. All figures are of females from the Rogers City Limestone. Figure 1— Apical view (xO-75) of Paratype, USNM 2 1 3755. Note the degree of hyperstro- phism and undulating shoulder on this large specimen. Figure 2 — Oblique basal view (x0-7) of Paratype, USNM 213764 showing well-developed projecting flanges. Figure 3— Basal view (x0-85) of Paratype USNM 213760. This mature specimen illustrates the well-developed basal flange-like pro- trusions of the female, but the collabral ornamentation is better developed than is typical. Figure 4 — Apical view (xO-6) of Paratype UMMP 57888. This very large female has only moderate hyperstrophism and no develop- ment of the undulating shoulder. Figure 5— Basal view (xl-0) of Paratype USNM 213767 showing strongly developed, spade-like basal flanges. Figure 6 — Apical view (xl-1) of Paratype UMMP 22379, showing undulating shoulder. Figure 7— Apical view (x3 0) of Paratype UMMP 22375. Very immature specimens such as this are very difficult to sex. This in- dividual has the degree of hyperstrophism which suggests a female, but the well- developed collabral ornamentation sug- gests a male. PLATE 6 Hypomphalocirrus rugosus n. gen. and sp. All figures of females from the Rogers City Limestone. Figures 1, 8 — Oblique basal (x0-7) and apertural (xl-25) views of Paratype UMMP 22379. A thin individual with undulating shoul- der, flanges and roughened outer whorl face. Figure 2 — Apertural view (xO 6) of Paratype UMMP 22378 showing strong hyperstro- phism. Figure 3 — Oblique basal view (x0-5) of Paratype USNM 213759 with an unusually high outer whorl face. Figure 4 — Oblique basal view (xO-8) of Paratype USNM 213772 showing well-developed undulations on shoulder and the rugose outer whorl face. Figure 5 — Apical view (xO 9) of Paratype USNM 213781. This specimen is a female on the basis of its hyperstrophism, but exhibits the male characters of strong collabral ornamentation and smooth shoulder. Figure 6 — Oblique basal view (x0-7) of Paratype USNM 213776 showing strong flanges on base and undulating shoulder. Figure 7 — Oblique apical view (xO 75) of Paratype, USNM 213765 showing exceedingly ru- gose outer whorl face- Figure 9 — Basal view (xO 6) of Paratype UMMP 22380. This specimen has an unusually round whorl profile and thus greatly re- sembles Omphalocirrus goldfussi. Figures 10, 11 — Oblique apical (x0-7) and oblique basal views (x0-9) of Paratype, USNM 213754 showing rugose outer whorl face and very strong undulating shoulder. PLATE 7 Hypomphalocirrus manitobensis (Whiteaves) Figure 1— Oblique apical view (xl-8) of Hypotype, USNM 213783 showing operculum set of an oblique angle inside the ultimate whorl. Coiling of multispiral operculum indicates this is orthostrophic coiling. Figures 2, 4— Whiteaves cotype, GSC 4174. The break in this steinkern marks the position of the operculum. This operculum is placed at a greater angle to the axis of the whorl than that found in Fig. 1, in- dicating that there was some rotation after death. Figure 3 — Oblique basal view of Whiteaves Plesio- type GSC 4176, a female showing full development of both rows of spines. Also note rugose outer whorl face. Figures 5, 6, 8 — Apertural, basal and apical views of Whiteaves Plesiotype GSC 4177a. Note the two rows of spines and the pseudo- OMPHALOCIRRIDAE 53 isostrophic coiling typical of the female. Figure 7 — Basal view of hypotype USNM 213782, an immature male showing well-developed basal inhalent tremata, collabral orna- mentation and hypertrophic coiling. All figures Verneuil) Figures 1, PLATE 8 of Omphalocirrus goldfussi (Archiac and 2, 9— Oblique basal (xO 8), basal (xO-75) and apical (xO-7) of latex cast (from USNM) of topotype figured by Knight (1941) of a specimen from the Puzo Col- lection, Ecole de Mine. This specimen is presumably a female characterized by being orthostrophic and having well-de- veloped inhalent tremata present through- out its growth. Figures 3, 6— Basal (xlO) and oblique basal (xO-8) views of latex cast (from USNM) of Holotype which is housed in the De Ver- neuil Collection at the Ecole national superieure des Mines, Paris, France. This is presumably a female showing the weak collabral ornamentation of the base. Figures 4, 5 — Oblique basal and basal views (xl-8) of a plaster cast (from USNM) of a specimen from Sotenich, Germany, Strin- gocephalus Limestone (Givetian). The original of this specimen is from Hum- boldt Universitat, Berlin, Germany. This is presumably a male and is thus hyper- strophic, with non-functional inhalent tre- mata and strong collabral ornamentation of the base. Figure 7 — Basal view (xl-0) of plaster cast (from USNM) of holotype of 'Arctomphahis grandis* TolmachofT, 1926, specimen No. A 19229 in the Paleontologist Museum of Oslo, Norway. The total size of this specimen is indicated by the dashed line, but the adult whorls are only represented by a steinkern. The inner whorls indicate the features typical of the male Omphalo- cirrus. Figure 8 — Basal view (xLl) of plaster cast (from USNM) of a specimen from Vilmar, Germany. Upper Stringocephalus zone. The original of this specimen is in the Humboldt Universitat, Berlin, Germany. This specimen is hypertrophic and shows the other male characters of incipient nonfunctional inhalent tremata and strong collabral ornamentation. PLATE 9 Liomphalus northi (Etheridge). All figures of females from the Lilydale Limestone, Cave Hill Quarry, Lily- dale, Victoria, Australia. Figures 1-3 — Apical, basal and apertural views (x2) of an immature female Hypotype NMV P28716, At this young age the females exhibit many of the male characters such as collabral ornament and a faint keel on top as well as bottom. Note however that it is orthostrophic rather than hypertro- phic as is the case with the males (cf. PI. 10, Fig. 5). Figure 4 — Apical view (xO-7) of Hypotype NMV P28709, showing rounded upper whorl face with no keel and no collabral orna- ment. Figures 5, 7, 8 — Oblique apical, apical and basal views (xO-7) of Hypotype, NMV P28711, a large female showing strong keel and collabral ornamentation on the base and no keel and no collabral ornamentation on the top. Figures 6, 9, 10 — Oblique apical (xO-7), basal and apical (x0-5) views of Hypotype, NMV P28712. Figure 11 — Cross section (xO-7) of Hypotype, NMV P28499 showing septal filling of early whorls. i Figures 12, 13 — Basal (xO-8) and apical (xl) views of Hypotype NMV P28708. Figures 14, 17 — Basal and apical views (xO-7) of Hypotype, NMV 28498. Note flat spire and depressed umbilicus. Figures 15, 16 — Basal and apical views (xO-5) of Hy- potype, NMV P28373, the largest speci- men in the collection. PLATE 10 Liomphalus northi (Etheridge). All figures of males from the Lilydale Limestone, Cave Hill Quarry, Lily- dale, Victoria, Australia. Figures 1-3 — Apical, basal and oblique apical views (xl) of Hypotype, NMV P28714, a typi- cal male showing collabral ornamentation and keel on both top and bottom. Also note the degree of hyperstrophism. Figures 4-6 — Apical, apertural and basal views (x2) cf Hypotype NMV P28719. This is a small male showing the degree of hyper- strophism typical of the males. Figure 7 — Basal view (xl) of Hypotype NMV P28707 showing very strong basal keel. Figures 8, 11 — Apical and oblique views (xO-7) of Hypotype, NMV P28717. On this strongly hyperstrophic male the upper keel be- comes lost on the ultimate whorl, but the collabral ornament continues all of the way. Figures 12, 17— Apical (xO 9) and basal views (xl) of Hypotype, NMV P28713. This male is unusual for though it is markedly hyper- 54 ROBERT M. LINSLEY strophic it has lost the keel and collabral ornament on the top. Figures 13, 1 6 — Apical (xl ) and oblique apical (xO 7) views of Hypotype, NMV P1107. This large male exhibits all of the typical features: keel and collabral ornamenta- tion top and bottom, and hyperstrophism. Note massive, multispiral operculum in place in the aperture. Figure 14 — Apical view (xO-7) of Hypotype, NMV P28710.This specimen still has the keel on top, but has lost the collabral orna- mentation. Figure 15— Apical view (xO 7) of Hypotype, NMV 28718. This crushed male shows strong collabral ornamentation and well- developed keel. IO0r 80 70 60 *>. &> < CO ,30 iao wo SHELL WIDTH (mm) Figure 1 — A plot of shell width on the abscissa ver- sus the width of the umbilicus divided by the depth of the umbilicus, a sensitive measure of the degree of hyperstrophicity. Note that in the larger individuals the cloud of points divides into two groups of about equal numbers. This is considered as strong evidence in favour of sexual dimorphism. MEM. NAT. MUS. VICT. 39 PLATE 2 \ -tf'f " I'r"*^. ~y i y» 3 ^ - ^pSSjj r. .. * 1 ■Jf _ £*f ^r : : Si! ,< 4' m. W I ' "9^ • i MKM. NAT. MUS. VICT. 39 PLATE 3 MEM. NAT. MUS. VICT. 39 PLATE 4 MEM. NAT. MLS. VICT. 39 PI. A I I 5 ■? ) 2 / / 4 >!'-%. % , :? J*~ "SJ -' k 1 -*1 ,;-. MEM. NAT. MUS. VICT. 39 PLATE 6 m MEM. NAT. MUS. VICT. 39 PLATE 7 iif^ MEM. NAT. MUS. VICT. 39 PLATE 8 Ml M. NAT. MUS. VICT. 39 PLATE 9 MEM. NAT. MUS. VICT, 39 PLATE 10 PROPLEOPUS CHILLAGOENSIS, A NEW NORTH QUEENSLAND SPECIES OF EXTINCT GIANT RAT-KANGAROO (MACROPODIDAE: POTOROINAE) By Michael Archer,* Alan Bartholomai* and Larry G. Marshall! * Queensland Museum t Princeton University Abstract ^ Propleopus chillagoensis is described as a second species in the genus. The genus is rediagnosed. The new species differs markedly in details of premolar and molar morphology from the Pleistocene P. oscillans (De Vis), but reveals no additional information about inter- generic relationships. In the unique characters which differentiate it from P. oscillans, it is less like structurally ancestral forms such as Hypsiprymnodon than is P. oscillans, and could be a descendent of the lineage leading to P. oscillans. A specimen of Propleopus from the Pleistocene Wellington Caves is similar to P. chillagoensis, and suggests the possibility that the two species, or lineages leading to them, were contemporaneous during part of the Pleistocene, and probably represent end members of a late Tertiary radiation. Introduction Propleopus oscillans was described on the basis of a single dentary fragment by De Vis (1888). Since then a lower incisor and two additional dentary fragments have been re- ported (Woods, 1960 and Tedford, 1955), as well as a maxillary fragment (Bartholomai, 1972). All of this Pleistocene material has been referred to P. oscillans with little hesita- tion. Gill (1953, 1957) described a tooth from the Pliocene Grange Burn local fauna which Ride (1964) and Turnbull and Lundelius (1970) regard as similar to but smaller than P. oscil- lans. The present material, comprising a maxillary fragment and two isolated premolars repre- sents a second very distinct species. The exis- tence of at least two distinct forms of Propleo- pus in the Pleistocene indicates that there was probably a late Tertiary radiation of the genus. Registration numbers are those of the Queensland Museum (e.g. F6675, fossil col- lection; e.g. J6824, modern mammal collec- tion), the National Museum of Victoria (e.g. P15917), the American Museum of Natural History (e.g. AMNH 65279) and Museum of Palaeontology, University of California (e.g. UCMP 51697). Dental terminology follows that used else- where by Archer (1974, 1976). An alternative nomenclature for individual premolar teeth, as used elsewhere (e.g. by Bartholomai, 1972), is not used here, but not because it has been shown to be any less appropriate than the sys- tem used by Archer (1974) and some other authors (e.g. Mahoney and Ride, 1975). Rediagnosis of Propleopus Longman, 1924 Recognition of a second species broadens the concept of Propleopus as follows: Poto- roine macropodids differing from other poto- roine genera in unique extent of subhorizontal wear facet on I a ; also differ from Hypsypri- mnodon in having enamel on Ij covering only lower half of labial and lingual faces; posterior half of P 4 shorter crowned; M x trigonid wide; differing from Caloprymnus and Potorous in distribution of enamel of Ii, larger more finely serrated and arcuate plagiaulacoid premolars, in having a vestigial tooth immediately pos- terior to Ii, and in having lingual cingula (sometimes as rudiments) on upper molars; differing from Aepyprymnus in having more arcuate plagiaulacoid premolars, poorly- developed lingual cusp or crest develop- ment on P 4 , wider molars, poorly-developed transverse lophs on molars; vestigial tooth immediately posterior to l u obliquely set plagiaulacoid premolars, and lingual cingula (sometimes rudiments) on upper molars; also differing from Bettongia in having more arcu- 55 56 MICHAEL ARCHER, ALAN BARTHOLOMAI, and LARRY G. MARSHALL ate, mcsially swollen plagiaulacoid premolars with narrow posterior ends on permanent pre- molars, and a vestigial tooth immediately pos- terior to Ix. Propleopus chillagoensis sp. nov. (Plate 11, fig. 1) Holotype: PI 59 17, right maxillary fragment with erupting P 4 and erupting and broken M\ fragment of M 1 , and M 2 - 3 , donated by Mr Hugh C. Mainwaring in 1910. Referred Material: PI 59 18, isolated LP 4 ; P15919, isolated and broken RP 1 ; both from type locality. Type Locality: Fissure deposit in Chillagoe Formation, Chillagoe District, N.E. Queens- land. According to a letter (National Museum of Victoria) from Mr Hugh C. Mainwaring to Professor Baldwin Spencer, 21 July 1910, the specimens came from \ . . a bluff 800 ft. long, 300 ft. wide and 50 ft. above the plain of the surrounding country. This bluff is being worked [1908-11] as a quarry to supply the smelter with flux. In breaking down the stone a number of small caves were found. These were fitted with material resembling silicified clay in which the fossils were imbedded'. Mr William Morrow (pers. comm.) notes that the locality was 'Smelter's Junction, one mile from Chilla- goe Railway Station, along the Mungana line, on the left hand side on the way to Mungana.' Origin of Name: Reference to type locality. Diagnosis: Differs from the only other species in the genus, Propleopus oscillans, in having a markedly larger P 4 and P 4 , each with nine ver- tical ribs; anterior ribs on P 4 are rectilinear rather than curved; P 4 crown base more steeply inclined posteriorly; upper molars shorter, with more swollen bases; lingual cingulum around base of protocone on at least M 2 ~ 3 ; preproto- crista connects directly with preparacrista; pos- terior half of upper molar crowns markedly narrower than anterior half. Description: P 4 : Two-rooted tooth; occlusal outline oval, widest at anterior one-half; an- terior end gently convex, posterior end blunt. Occlusal cutting edge gently convex, ventrally with maximum convexity occurring towards posterior one-half of edge. Anterior edge of tooth has curved, labially convex, vertical crest confluent with occlusal cutting crest of tooth. Posterior to this anterior crest are eight pairs (lingual and labial) of prominent vertical ridges which reach the occlusal edge of tooth. Ninth pair fails to extend as far as occlusal edge. Longitudinal spacing between ridges de- creases posteriorly. Bases of second pair of ridges bifurcated. Bases of eighth and ninth buccal ridges confluent. Ninth ridges more massive than others and form shallow vertical fossettes between themselves and eighth ridges. Relative ridge lengths as follows 3 sub- equal to 4, 2 subequal to 5, 6, 7, 8, 9. Eighth ridge interrupted by gap on each side. Base of crown lacks cingula. Judging by position of erupting crown, P 4 probably oriented antero- labially, with respect to long axis of molar row. P 4 : Identical to P 4 except as follows: Pos- terior edge of crown keeled with non-serrated vertical continuation of occlusal cutting crest; widest part of crown closer to middle of tooth; ninth ridges less massive than corresponding ridge in P 4 ; base of second ridge not bifurcated; base of second and third lingual ridges con- fluent. M 1 : Crown represented by only posterior one- third. Roots indicate anterior one-half of tooth much wider than posterior one-half, and that lingual one-half of tooth longer than labial one-half, result of evidently very large protocone. Remnant of posthypocrista and postmetacrista linked in arc, middle of which functions as a posterior cingulum. Base of crown very swollen, such that transverse dia- meter of crown base below hypocene and meta- cone estimated at three times distance between those cusps. M 2 : Anterior one-half of crown wider than pos- terior one-half. Lingual and labial sides of crown subequal in length. Protocone, hypo- cone and paracone subequal in height, and taller than metacone. Preprotocrista and pre- paracrista connect as an anteriorly convex arc. Postimetacrista and posthypocrista joined as in M 1 . Prehypocrista and postprotocrista linked in barely obtuse angle. Postparacrista and pre- metacrista become irregular towards middle of crown, and are only just linked by low lon- gitudinal crest. Prominent, transverse lingual EXTINCT GIANT RAT-KANGAROO PROPLEOPUS 51 ridges descend flanks of paracone and meta- cone from longitudinal midline of crown. Poorly-defined transverse labial ridge descends flank of hypocone and meets lingual rib of metacone in longitudinal midvalley. Very poorly-defined anterolingually directed labial ridge descends protocone, but intersects the preprotocrista anterior to the protocone. La- biad to point of intersection of postparacrista and premetacrista are two small adjacent cusps along labial end of transverse midline. Prom- inent lingual anterolingually inclined ridge ascends flank of hypocone to base of proto- cone, adjacent to small basal lingual cuspule, which in turn is adjacent to lingual end of an- terolingual basal cingulum which terminates labially at point where M 1 abuts against M 2 . Very small cingular bulge occurs on postero- lingual base of hypocone. M 3 : Anterior one-half of tooth much wider than posterior one-half. Lingual one-half subequal in length to labial one-half. Proto- cone subequal in height to hypocone, aller than paracone, and much taller than re- duced metacone. Prehypocrista, postproto- crista, preprotocrista and preparacrista linked in M 2 . Posthypocrista does not contact ves- tigial postmetacrista, being separated by posterolabial depression. Poorly developed premetacrista and postparacrista only just fail to contact in transverse midvalley of tooth. Prominent lingual rib descends para- cone in posterolabial direction from longitu- dinal midline of crown. Much more poorly de- veloped linqual and labial ridges descend flanks of metacone and hypocone but just may not meet each other in longitudinal midvalley. Poorly-developed labial ridge descends proto- cone in same manner as in M 2 . Prominent lingual rib ascends flank of hypocone to base of protocone. Vertical lingual ridge, also des- cends from base of protocone to point halfway down its flank. Base of this ridge vaguely con- tacts oblique lingual hypocone ridge and an- terolingual basal cingulum on anterior flank of protocone. Anterolingual cingulum terminates labially just short of point at which M 2 abuts M 3 . Poorly-developed (but better than M 2 ) posterior cingulum descends from posterior flank of hypocone to posthypocrista near its labial end. M 3 less worn than M 2 and reveals fine crenulations over much of crown surface enclosed by principal cusps. Also evidence of accessory vertical crenulations parallel to la- bial ridges of protocone and hypocone, and connecting posthypocrista with longitudinal midvalley of crown. M 4 : Only anterior one-half of crown remains. From remnant of base of posterior one-half, evident that anterior one-half wider than pos- terior one-half. Protocone taller than paracone. Preprotocrista connected to preparacrista as in M 3 . Postparacrista not connected to pre- metacrista. Lingual ridge of paracone promin- ent and extends transversely to longitudinal midvalley to meet labial ridge of protocone. Some evidence for lingual protocone ridge as in M 3 . Tooth erupting so anterior cingulum not visible, if present. Edge of enamel near pos- terior base of protocone suggests lingual hy- pocone ridge present as in M 3 . Very poorly- developed vertical ridge occurs on anterolabial flank of paracone linking preparacrista with base of crown. As in M 3 , clear indications of accessory crenulations occur in crown valleys and on flanks of cusps. Meristic changes along tooth row : Pos- teriorly, teeth become relatively narrower and longer; metacone markedly reduces in size; paracone reduces in size but at a slower rate; crests linking metacone to other crests become smaller; transverse midvalley of crown be- comes more open at labial end; bases of pro- tocone and hypocone become less swollen; distance between hypocone and metacone diminishes. Tooth eruption sequence indicated suggests P 4 and M 4 erupt at approximately the same time. Discussion: The type specimen of Propleopus oscillans is a dentary while that of P. chillago- ensis is a maxillary fragment. A possibility therefore exists that both specimens represent P. oscillans. We have considered this possi- bility and discounted it for the following rea- sons. First, Bartholomai (1972) has described F6675 from the Darling Downs, a maxillary fragment with LP 4 -M 2 , and referred it to P. oscillans. It is specifically distinct from the 58 MICHAEL ARCHER, ALAN BARTHOLOMAI, and LARRY G. MARSHALL 0.96 1.10 1.25 1.50 B 1.47 0.97 0.81 0.69 Figure 1— Measurements of specimens of Propleo- pus chillagoensis sp. nov. A-C, P15917, holotype. D-E, P15918, LP 4 . F, P159I9, RP 4 . Measurements are in centimetres. Chillagoc maxillary fragment, for reasons noted in the diagnosis. In addition, although P 1 of the type of P, chillagoensis is larger than P ( of F6675, its molars are proportionately smaller. For this reason, the size differences in the two maxilla cannot be the result of allo- metry. Additional evidence for reference of F6675 to Propleopus oscillans comes from the num- ber of ridges on upper and lower adult pre- molars of living potoroines. In modern Aepy- prymnus rufescens, of twenty-three specimens examined with relatively unworn premolars, nine have the same number of ridges (seven or eight) on the upper and lower premolars, while the remainder differ by only one ridge. Varia- tion in ridge number is as follows: P 4 : one has seven ridges; thirteen have eight; and eight have nine; P.,: three have seven ridges; thirteen have eight; and three have nine. In Hypsiprimnodon moschatus, of all ten speci- mens with relatively unworn adult premolars, P 4 has six and P 4 seven ridges. Both upper and the single lower premolars of P. chillagoensis have nine ridges, while the P 4 of the holotype of P. oscillans and P 4 of the referred specimen both appear to have seven ridges. The con- stancy of ridge number in Hypsiprimnodon, the consistently different ridge number in P. chilla- goensis from the holotype of P. oscillans, and the apparently close relationship of Propleopus to Hypsiprimnodon (see below) which has a species constant number of ridges, convince us of the correctness of Bartholomai's (1972) re- ference of F6675 to P. oscillans. Added to the other morphological characters noted in the diagnosis, it also convinces us of the distinction between the two species of Propleopus. Further, but less convincing evidence for the association suggested by Bartholomai (1972), is the fact that both the holotype of P. oscillans and F6675 come from the Darling Downs of southeastern Queensland, while the specimens of P. chillagoensis come from far northeastern Queensland. These data, as well as the close approxima- tion of upper and lower molar sizes, support Bartholomai's (1972) reference of F6675 to Propleopus oscillans and convince us that the EXTINCT GIANT RAT-KANGAROO PROPLEOPUS 59 Chillagoe material with its nine vertical ridges, and relatively short and wide premolars can- not be referred to P. oscillans. ^ Most authors who have considered the affini- ties of Propleopus oscillans regard it to be closely related to Hypsiprymnodon, a conclu- sion with which we generally agree, although some characters more closely resemble Betton- gia than Hypsiprymnodon. P. chillagoensis re- veals no characters that more closely ally it with Hypsiprymnodon or Bettongia than with P. oscillans. The characters which differentiate it from P. oscillans are unique derived charac- ters and are of neutral value in assessing inter- generic relationships. Because P. oscillans re- tains more structurally pr vitimiecharac tains more structurally primitive character states (i.e. fewer ridges on P\ relatively un- swollen molars, poorly-developed lingual cin- gula, and unreduced metacones on the upper molars), and has no derived characters of its own which are not also present in P. chilla- goensis and hence diagnostic of the genus as a whole, it can be regarded as structurally, if not actually ancestral to P. chillagoensis. The precise age of the eastern Darling Downs specimens within the Pleistocene is in doubt. Similarly, the age of the Chillagoe speci- mens is in doubt but the presence of Sarco- philus, suggests that the Chillagoe deposit is no older than Pleistocene (Bartholomai and Marshall, 1973). Tedford (1967) mentions that the material described here as P. chilla- goensis is associated with Sarcophilus and small macropodines apparently related to liv- ing species. The Propleopus specimen (UCMP 45171) figured by Tedford (1967, fig. 5b) as P. oscil- lans from Pleistocene deposits in Wellington Caves, New South Wales, although only a frag- mentary dentary lacking P 4 , shows in some re- spects, the kinds of characters which might be expected in lower molars of P. chillagoen- sis. Compared with P. oscillans, the anterior half of Mi is relatively wider, the molars are somewhat shorter, and most importantly, the talonid of M 4 is narrower as noted by Ted- ford (1967) and its entoconid is more reduced. The entoconid is the occlusal counterpart of the metacone in the upper molars. Reduction of one is likely to be reflected by reduction of the other. Its taxonomic position is considered as doubtful until better material becomes avail- able. Tedford (1967) concludes that the late Pleis- tocene, Lake Menindee, New South Wales specimen (UCMP51697) of Propleopus is refer- able to P. oscillans, a conclusion with which we agree. A second tooth fragment from Men- indee (UCMP51698) was compared by Ted- ford (1967) with the Chillagoe material. It is likely that this specimen is also referable to P. oscillans. These occurrences suggest the possibility that the lineages resulting in Propleopus oscil- lans and P. chillagoensis may have been con- temporaneous during the Pleistocene and rep- resent parts of a late Tertiary radiation of the genus. Acknowledgements We thank Dr T. Rich (National Museum of Victoria) for making the Chillagoe specimens available to us. The late Professor R. A. Stir- ton provided the Queensland Museum with a cast (F3287) of the Wellington Caves speci- men. Mr A. Easton and Mr R. Burgess (Queensland Museum) produced the photo- graphs for the plate. Ms J. Utz and Ms R. Owens (Queensland Museum) typed drafts of the paper. Mr S. Van Dyke (Queensland Mu- seum) assisted in study of premolar ridge num- bers in modern potoroines. References Archer, M., 1974. The development of the cheek- teeth in Antechinus flavipes (Marsupialia, Dasyu- ridae). /. R. Soc. West. Aust. 51: 54-63. , 1976. Phascolarctid origins and the po- tential of the selenodont molar in the evolution of diprotodont marsupials. Mem. Qcl Mus. 17: 367-71. Bartholomal, A., 1972. Some upper cheek teeth in Propleopus oscillans (Dc Vis). Mem. Qd Mus. 16: 211-213. , in press. The fossil vertebrate fauna from Pleistocene deposits at Cement Mills, Gore, south-eastern Queensland. Mem. Qd Mus. 18. Bartholomai, A., and L. G. Marshall, 1973. The identity of the supposed dasyurid marsupial, Sarcophilus prior De Vis, 1883, with comments on other reported 'Pliocene' occurrences of Sarco- philus. Mem. Qd Mus. 16: 369-374. 60 MICHAEL ARCHER, ALAN BARTHOLOMAI, and LARRY G. MARSHALL De Vis, C. W., 1888. On an extinct genus of the marsupials allied to Hypsiprymnodon. Proc. Linn. Soc.N. S.W.I: 5-8. Gill, E., 1953. Australian Tertiary marsupials. Aust. J.Sci. 16: 106-108. , 1957. The stratigraphical occurrence and palaeoecology of some Australian Tertiary mar- supials. Mem. Nat. Mus. Vic. 21: 135-203. Mahoney, J. A. and W. D. L. Ride, 1975. Index to the genera and species of fossil Mammalia described from Australia and New Guinea be- tween 1838 and 1968. Spec. Pubis. West. Aust. Mus. 6: 1-250. Ride, W. D. L., 1964. A review of Australian fossil marsupials. /. R. Soc. West. Aust. 47: 97-131. Tedford, R. H., 1955. Report on the extinct mam- malian remains at Lake Menindee, New South Wales. Rec. S. Aust. Mus., 11: 299-305. , 1967. The fossil Macropodidae from Lake Menindee, New South Wales. Univ. Calij. Publ. Geol. Sci., 64: 1-156. Turnbull, W. D., and E. Lundelius, Jnr., 1970. The Hamilton fauna — a late Pliocene mamma- lian fauna from the Grange Burn, Victoria, Australia. Fieldiana (Geology) 19: 5-84. Woods, J. T., 1960. The genera Propleopus and Hypsiprymnodon and their position in the Macro- podidae. Mem. QdMus. 13: 199-212. Explanation of Plate PLATE 11 Propleopus chillagoensis sp. nov. A, stereopair, oc- clusal view, P15917, holotype, RP 4 -M 4 . B, labial view, P15917, holotype. C, stereopair, occlusal view, P15918, LP 4 . D, labial view, P15918, E, stereopair, occlusal view, P15919, RP 4 . F, labial view, P15919. White line is one cm in length. MEM. NAT. MUS. VICT. 39 PLATE 11 DETAILED COMPARISONS OF THE DENTITIONS OF EXTANT HEXANCHID SHARKS AND TERTIARY HEXANCHID TEETH FROM SOUTH AUSTRALIA AND VICTORIA, AUSTRALIA (SELACHII: HEXANCHIDAE) By Noel R. Kemp Tasmanian Museum, G.P.O. Box 1164M, Hobart, 7001, Australia Abstract In extant hexanchid sharks except for (usually) a bigger primary cusp, isolated teeth of a given size of the smaller species Hexanchus vitulus (Springer and Waller) may be confused with those of H. griseus (Bonnaterre). This specific size difference has significance in the fossil record. Heptranchias perlo (Bonn.) differs in its more slender and relatively larger primary cusp with basal denticles (not serrations as in Hexanchus) on its mesial margin and fewer crownlets increasing and then decreasing in size distally. Notorynchus cepedianus (Peron) differs mainly in its more robust primary cusp and crownlets which are fewer in number than in Hexanchus and which decrease evenly in size distally, like Hexanchus. Cali- fornian specimens of Notorynchus are included in the monotypic TV. cepedianus until the taxonomic significance of the variability of the upper medial teeth is established and defined. N. cepedianus is recorded from the fossil record for the first time. Heptranchias haswelli Ogil- by is regarded as a species inquirenda. Hexanchus agassizi Cappetta is expanded to include the Eocene Hexanchus teeth from South Australia. Heptranchias howellii (Reed) from the South Australian Eocene and Notorynchus primigcnius (Agassiz) from the Victorian Mio- cene are recorded and described from Australia for the first time. Introduction The sixgill and sevengill sharks of the family Hexanchidae, although a relatively small group, are well represented in the fossil record, es- pecially outside Australia, for example Wood- ward (1886), Jordan (1907), Leriche (1910, 1927, 1957) and are common in modern seas, for example Garman (1913), Bigelow and Schroeder (1948), Whitley (1968), Bass, D'- Aubrey and Kistnasamy (1975). Taxonomic- ally, however, they are not a stable group (see below). Less than a dozen fossil hexanchid teeth have been described from the Australian fossil record (Pledge, 1967; Kemp, 1970). Three teeth and several fragments from the Eocene of South Australia were assigned by Pledge (1967) to 'Notidanus' serratissimus Ag., W cf. serra- tissimus and W? serritissimus. The three fragments from the Miocene of Victoria were described by Kemp (1970) as Notorynchus cf. primigenius. In light of recent publications (Welton, 1974; Cappetta, 1976) and with the recovery of further specimens from the South Australian Eocene during the last decade, the present study shows that the three extant gen- era are represented in the South Australian and Victorian Tertiary material. Compagno (1973) lists two families in the suborder Hexanchoidei; Hexanchidae, with two genera, Hexanchus and Notorynchus and Heptranchidae with the monotypic genus Hep- tranchias. He based this separation of Hep- tranchias into its own family on a suggestion by Dr Shelton P. Applegate 'because of many differences in cranial and external morphology between it and the other hexanchoids {Hexan- chus and Notorynchus)' (Compagno, 1973:33). Recently Bass et al. (1975) suggest that more detailed anatomical studies of the six- and sevengill sharks may show that the smaller species of Hexanchus, H. vitulus, may have more in common with Heptranchias perlo than with the larger species, Hexanchus griseus. H. griseus, on the other hand, may have more in common with Notorynchus cepedianus. Until these suggestions are confirmed or repudiated by further detailed work the suprageneric clas- sification adopted here is that of Patterson (1967). Terminology The tooth types medials, laterals and pos- teriors are used in the sense of Applegate (1965a). The faces of a tooth are termed la- bial and lingual and the margins mesial and 61 62 NOEL R. KEMP entire ma basal primary cusp denticles \ T crown root crow nlets LABIAL LINGUAL serrations ingual thickening mesial + distal MESIAL LABIAL Figures 1-4 — Terminology. 1-3. Notary nchus cepedianus (Peron), lower right second lateral tooth (Figs. 1, 2, x2;Fig. 3, xl-5). 4. Hexanchus agassizi Cappetta, lower left lateral of about fourth, fifth or sixth row (xl-5). distal as defined by Hoojier (1954). In the cockscomb-like dentition of the hexanchids the larger primary cusp is followed by a number of crownlets (Applegate, 1965b). The mesial mar- gin of the primary cusp may be serrated, or, if the serrations are relatively large, they are termed basal denticles. A character of the family is the presence of one series of func- tional teeth in both jaws except for the upper medials which have two or three and the posteriors of both jaws which have two to four functional series. The description of dentition of Hexanchus griseus is given in de- tail as a datum for the diagnoses of the other species. Abbreviations The following abbreviations are used: AMS, Australian Museum, Sydney; AUGD, Univer- sity of Adelaide, Department of Geology; CM, Canterbury Museum, Christchurch, New Zea- land; MUGD, University of Melbourne, De- partment of Geology; NMV, National Museum of Victoria, Melbourne; NZNM, National Mu- seum of New Zealand, Wellington; RJFJ, pri- vate collection of Dr R. J. F. Jenkins, South Australia; SAMD, South Australian Depart- ment of Mines; SAM, South Australian Mu- seum, Adelaide; TFF, private collection of Mr T. F. Flannery, Beaumaris, Victoria; USNM, United States National Museum, Washington. Systemalics Order HEXANCHIFORMES Suborder HEXANCHOIDEI Family HEXANCHIDAE The Hexanchidae are characterized by six or seven gill-openings and heterodont denti- SOUTH-EASTERN AUSTRALIAN HEXANCHID SHARKS 63 tion, that is the teeth are of dissimilar shape both within and between the two jaws. Except for another two families all other sharks have only five gill openings. The Chlamydoselachidae, or frill shark, of which only one specimen has been recorded from Australian waters, in 1976 (J. R. Paxton, pers. comm.), has a terminal mouth compared with the subterminal mouth of all other sharks and six gill openings with the first opening con- tinuous across the throat. The remaining five openings are interrupted on the ventral sur- face. The dentition of Chlamydoselachus, the single genus of the family, is homeodont, that is all the teeth are of similar shape both within and between the two jaws. The Pristiophoridae, or saw sharks, has a sixgill genus, Pliotrema, which has yet to be found in Australian waters. It is easily sepa- rated from the sixgill hexanchid species by its two dorsal fins, rostral snout with barbells and marginal teeth, homeodont dentition and no anal fin. The hexanchids have only one dor- sal fin, no rostral snout and possess an anal fin. Main characters One dorsal fin, always posterior to pelvics; anal fin present; caudal fin with a definite lobe and well marked subterminal notch ; pre- caudal pit absent; six or seven gill-openings not connected ventrally, last gill opening in front of and extending below pectoral origin; spiracle present but small; nictitating mem- brane absent; upper labial furrow absent, lower labial furrow present, developed to varying de- grees. Dentition heterodont: upper jaw, teeth with slender primary cusp and a variable number of small crownlets; small medial tooth present or absent; lower jaw, teeth basically trapezoi- dal in outline and multicusped; small medial tooth present. Small undifferentiated posterior teeth present in both jaws. As all species of hexanchids possess a simi- lar pavement-like dentition formed by the posteriors in both jaws and as they are rarely recognized in the fossil record, teeth of this type are omitted in the following descriptions and diagnoses. Genera Rafinesque (1810) erected two genera, Hex- anchus and Heptranchias, based on six- and sevengill sharks respectively. Cuvier (1817, fide Bigelow and Schroeder, 1948) included both of these genera in his new genus Noti- danus. This genus also came to include an- other sevengill shark, Notorynchus, both before and after Ayres (1855) described it as a new genus. Subsequent studies showed that the three genera of Rafinesque and Ayres were valid and Cuvier's Notidanus has not been used for extant sharks for more than half a century. Palaeontologists have until recently used Notidanus as an encompassing genus, for example Leriche (1957), Casier (1966), Cap- petta et al. (1967) (fide Cappetta 1970), Pledge (1967), as it was widely held that sep- aration of the genera of hexanchids was only possible on the basis of the anatomy of their soft parts. A number of authors, however (Ap- plegate 1965b; Waldman 1970; Welton 1974; Cappetta 1975) have noted some of the sa- lient dental differences, mainly in the lower lateral teeth, between these three genera. An- tunes and Jonet (1969), in describing Miocene teeth from Portugal, include in the family Hex- anchidae only two genera, Hexanchus Rafines- que and Heptranchias Rafinesque. They in- clude Notorynchus Ayres in the latter genus but give no reason for this. As set out in the key below the sixgill genus is readily separated from the two sevengill genera which differ from each other in the shape and size of their snouts. The descriptions under each genus further differentiates them on their dental characteristics. KEY TO LIVING GENERA (After Bigelow and Schroeder, 1948). 1 A Six gill-openings Hexanchus IB Seven gill-openings 2 2A Head narrow; snout tapering; hori- zontal diameter of eye considerably greater than distance between nos- trils Heptranchias 2B Head broad; snout broadly rounded; horizontal diameter of eye consider- 64 NOEL R. KEMP ably smaller than distance between nostrils Notorynchus Genus HEXANCHUS Rafinesque, 1810 Hexanchus Rafinesque, 1810:14; type species Squalus griseus Bonnaterre, 1788. Generic Diagnosis Six gill-openings, not joining across the ventral surface; lower labial furrow well de- veloped. No central medial tooth in upper jaw; lower laterals with 5 to 6 crownlets in specimens of about 500 mm total length, increasing up to 10 to 12 in specimens of about 4000 mm total length. Characters otherwise those of family. Living Species Two species of the genus Hexanchus are recognized (Springer and Waller, 1969), H. griseus and H. vitulus. The larger species, H. griseus, which reaches sexual maturity at about 4500 mm has 5 lower lateral teeth on each side of the jaw. The smaller species, H. vitulus, which reaches sexual maturity at about 1400 mm to 1750 mm has 6. KEY TO LIVING SPECIES OF HEXANCHUS (After Springer and Waller, 1969) 1A Five rows of large trapezoidal teeth on each side of the lower jaw griseus Bonnaterre, 1788 IB Six rows of large trapezoidal teeth on each side of the lower jaw vitulus Springer and Waller, 1969 Bass et al. (1975) list a number of ap- parent differences between the two species. One of these differences, relating to the shape of the symphyseal region of the upper jaw, may be a post-mortem artifact caused either by drying of the removed jaws or by fixation of in situ jaws in spirit collections. In H. griseus the upper medial teeth are set ap- proximately in line with the general curvature of the jaw whereas in H. vitulus the upper me- diate bulge markedly forward from the line of the rest of the jaw (Bass et al., 1975:7, PI 3A, 3B). In three of the five jaws of H. griseus in the present study, AMS 119110-001, USNM 188048 and NMV A235 the upper medials bulge forward exactly as shown by the South African H. vitulus (Bass et al, 1975, PI. 3B). In one specimen of H. vitulus (USNM 1 12600) the upper medials do protrude but only slightly, much less so than in the Victorian H. griseus. These jaws are still in situ, the shark being pre- served in the National Museum of Victoria's spirit collection. Hexanchus griseus (Bonnaterre, 1788)* * The date of Bonnaterre's Squalus griseus is often quoted in post-1948 literature as 1780. This error seems to stem from Bigelow and Schroeder's (1948) incorrect date on page 80. The correct date is used on pp. 78 and 85. Bluntnose sixgill shark, Mud shark, Cow shark (Plate 12, figures 1-4) Squalus griseus Bonnaterre, 1788: 9. Notidanus griseus (Cuvier, 1817); Agassiz, 1835, PL E, figs. 2-4; 1838: 92; 1843: 218. Notidanus griseus Cuvier, 1817; Agassiz, 1870:397. Notidanus griseus (Linne-Gmelin, 1788); Le- richc, 1910: 225, Fig. 70. Hexanchus griseus (Bonnaterre, 1788); Cle- mens and Wilbv, 1946: 52, Fig. 12; Parrott, 1958: 83, text fig.; Whitlev, f968: 5; Wel- ton, 1974: 2, PI. IE; Bass et al, 1975: 8, Fig. 5, PI. 1,3 A. Hexanchus griseus (Bonnaterre, 1780); Bige- low and Schroder, 1948. 80 {part), Figs. 8, 9 (non Fig. 9A = H. vitulus, S. Springer, pres. comm.); Lynch 1963: 295, Figs. 1-5; Springer and Waller, 1969: 169, Figs. 2B, 5-7. Hexanchus griseum (Bonnaterre, 1788); Springer and Garrick, 1964, Table 1. ? Isurus oxyrhinchus Rafinesque, 1810; An- tunes and Jonet, 1969: 136 (part.), PL 7, fig. 22 (non Fig. 24). (non) Hexanchus cf. griseus (Bonnaterre, 1780); Antunes and Jonet, 1969: 130, PL 4, figs. 1-3. Extant Material Examined. 2187 mm female taken in 200 m off Port Fairey, Vict., Mar., 1963 (NMV A235, spirit collection); jaws of 4250 mm specimen taken in 450 m off Norah Head, N.S.W., June, 1976 (AMS 119110-001); photographs of jaws of 4330 mm female, Gulf SOUTH-EASTERN AUSTRALIAN HEXANCHID SHARKS 65 of Mexico (USNM 188048) (Springer and Wal- ler, 1969: 169); photographs of tooth sets (Applegate, 1965a) of one side of jaws of eight specimens ranging from 696 mm to 4346 mm from California, U.S.A. Teeth from specimens of about a metre or less in length show less variation. Mesial margin may be entire and distal margin may have only small crownlets, perhaps only 2 or 3, even in rows nearest jaw articulation. Dentition Dental formula of AMS 1191100-001 (PI. 12, figs. 1-4) P8L9M4L9P8. Numerical variation P8 L6 Ml L6 P9 is restricted to the upper laterals, from 7 to 9 rows on each side. The number of upper and lower medials and lower laterals is constant in the above material and in the figured dentitions e.g. Bigelow and Schroeder, 1948, Fig. 8E; Bass et al, 1975, PI. 1. Bigelow and Schroe- der (1948: 82, footnote) record a small Medi- terranian specimen in which there is no lower medial tooth. In all teeth the crown and its subdivisions — cusps, crownlets, basal denticles or serrae — are equally biconvex labio-lingually. The root is simple, not branching, approximately square or trapezoidal in outline and is wedge-shaped, thinning from the base of the crown to the basal margin. Upper Jaw Medials. Four rows with lanceolate crowns curved away from the symphysis, two rows on each side. Margins entire except in large speci- mens about 2-3 m or more in length which may have margins serrated or even with basal den- ticles. Root rhomboidal in outline and with a pronounced thickening labio-lingually. Laterals. From 7 to 9 rows on each side of the jaw. Teeth of first row about as high as broad with a slightly inclined, curved primary cusp and 1 or 2 crownlets distally. Basal half of mesial margin serrated. Teeth of last rows about twice as broad as high with a more in- clined and almost straight primary cusp only slightly bigger than the first crownlet; 3 to 6 more crownlets decreasing evenly in size dis- tally. Basal two-thirds of mesial margin of tooth finely serrated. Other laterals grading in size and shape between these end members. Lower Jaw Medial. Tooth about as high as broad with from 3 to 6 laterally directed secondary cusps and either with or without a central cusp. Cen- tral cusp when present may be bifurcated and symmetrical or single and asymmetrical. Root approximately square in outline. Laterals. Six rows on either side of the medial, lower laterals largest in a set of jaws; basically trapezoidal in shape and about twice as broad as high. Crown divided into a primary cusp only slightly bigger than the following crown- lets which decrease evenly in size distally. From 6 to 8 or 9 crownlets in specimens about 2 m or less in length. In larger specimens the number of crownlets increases from 9 to 10 in first rows through to 10 to 12 in last rows. In- clination of crownlets more or less constant throughout laterals. Mesial margins smooth in small specimens. Basal two-thirds finely ser- rated in specimens about 1-5 m in length and more coarsely serrated in larger specimens. Serrations coarsest midway along mesial mar- gin, serrations of first rows coarser than those of last. Root more or less rectangular in out- line. Discussion. In all hexanchids the upper lal teral rows towards the jaw articulation are similar in shape to their lower equivalents. Common to the three genera however are dif- ferences which may serve to differentiate these respective tooth types. In the uppers the prim- ary cusp may be relatively higher and broader than in the lowers. The mesial margin of the primary cusp is usually more than half the length of the upper tooth whereas in the lowers it is less than half and often near one-third, i.e. the primary cusp in the uppers is more cen- trally placed. There are always more crown- lets in the lowers than in the uppers — this may be less significant with isolated teeth. 66 NOEL R. KEMP Three Upper Miocene hexanchid teeth are referred to Hexanchus cf. griseus by Antunes and Jonet (1969). The crowns of the two up- per mediate (Antunes and Jonet, 1969, PL 4, figs. 1, 2) are too slender and too obliquely curved to be H. griseus. The lower lateral tooth (Antunes and Jonet, 1969, PL 4, fig. 3) has a large primary cusp with reasonably coarse serrations on the mesial margin fol- lowed by six well developed crownlets. The low number and relatively large size of the crownlets compared with the size of the tooth strongly suggests a notorynchid tooth, prob- ably Notorynchus primigenius. The tooth figured by Antunes and Jonet (1969, PI. 7, fig. 22) as an abnormal hunts oxyrinchus is very similar to an upper medial in an Austra- lian Hexanchus griseus (AMS 119110-001, PL 12, fig. 2) in its size, shape and obliquity of the crown and the presence of small denticles at the base of the mesial margin and is probably referable to that species. Distribution. Hexanchus griseus occurs in the Atlantic, Pacific and Indian Oceans and in the Mediterranean Sea inhabiting both tropi- cal and temperate water (Bigelow and Schroe- der, 1948). Bass et al. (1975) note that H. griseus and Hexanchus vitulus of equivalent size occupy different ranges. The young of H. griseus inhabit temperate regions while H. vitulus occupies the tropical areas. H. griseus is often taken on long lines at about 200 m and as deep as 1400 m; it is also known from shoal waters (Bigelow and Schroeder, 1948). Springer and Waller (1969) suggest that it lives on the continental slopes at depths of 350 m or more, occasionally mov- ing into shallower waters. Vaillant recorded a 4-5 m female being taken in shallow water on a bar off Archachan in the Bay of Biscay (fide Springer and Waller, 1969). Hexanchus vitulus Springer and Waller, 1969 Bigeye sixgill shark (Plate 12, figure 5) Hexanchus vitulus Springer and Waller 1969: 160, Figs. 1, 2A, 3, 4; Bass et al, 1975: 9, Fig. 6, PL 2, 3B. Extant Material Examined. Photographs of teeth in situ of two dried jaws: no data (USNM 110900); 1630 mm female taken at 275 m, Bahamas, 1948 (USNM 112600). Dentition Tooth counts (Springer and Waller, 1969; Bass et al., 1975) show a range of P8-9 L 7 M4 L7 pg . These data represent at least ten P3-5 L5 mi L5 ps j aws w hich suggests that the num- ber of upper laterals is constant (cf. H. griseus) and numerical variation is restricted to the pos- teriors of both jaws. Diagnosis. Teeth small, basically similar to those of H. griseus. Upper Jaw Medials. Crowns may be more curved than in H. griseus and margin between top of root and base of crown, distally, more strongly arcuate. Laterals. Similar to H . griseus but primary cusp proportionately bigger than distal crownlets and mesial margin more finely serrated. Lower Jaw Medial. Similar to medial of H. griseus includ- ing variability of central cusp. Laterals, Primary cusp usually proportionately bigger than distal crownlets compared with H. griseus. Discussion. An important feature demonstrated by the dentition of these two species of Hexan- chus is that of size difference. The largest tooth from a large, sexually mature H. vitulus rarely exceeds 22 to 25 mm whereas that from a large, sexually mature H. griseus could be from 45 to 50 mm in length. Isolated teeth of the same size and of the same jaw position could be difficult to refer to one species or the other. However, with a random sample of teeth such as in the fossil record, if the two species were present at the time of deposition one would ex- pect a size difference in the teeth to be ap- parent thus demonstrating the existence of the two species. The size difference could also be interpre- ted as simply being teeth from juvenile and adult specimens of H. griseus. Allometric SOUTH-EASTERN AUSTRALIAN HEXANCHID SHARKS 67 growth patterns would enable the differentia- tion of a tooth of a juvenile H. griseus from a tooth of an adult H t vitulus, for example there would be more crownlets on the H. vitulus tooth than on the similar-sized H. griseus specimen. Empirically though, in any one species of fossil teeth there is a tendency to size cluster- ing, especially in smaller samples. (In contrast, however, in the National Museum of Victoria's collection of sharks' teeth there are 3-4000 teeth of fcuru$ hastalis (Agassiz, 1838) and a size range of juvenile to adult teeth is ap- parent, but, the majority of specimens repre- sent large sharks; a collecting bias may favour this size distribution). There are many vari- ables involved to produce this result but one dominating factor could be that the teeth shed by living sharks are insignificant in the fossil record compared with those from a dead adult. If an adult dies, immediately there is the po- tential of hundreds of teeth and crowns being preserved. In Notorynchus cepedianus for ex- ample there would be about one hundred while in a species such as Odontaspis taurus Rafin- esque, 1810 — the grey nurse shark, in Austra- lia — it could be as high as three to four hun- dred. Distribution: H. vitulus has been recorded from the south-eastern Indian Ocean between Kenya and Natal (Bass et ah, 1975), from Florida — Gulf of Mexico- West Indies region — and from the Philippines (Springer and Waller, 1969). Miiller and Henle's (1841) records of H. griseus from the Atlantic and Mediterranean may well include H. vitulus as they state that their specimens had 5 or 6 rows of broad teeth in the lower jaws (fide Springer and Waller, 1969). H. vitulus is a tropical to subtropical de- mersal species being taken at depths between 90 and 600 m with one specimen being netted just off shore in Natal. It is rarely taken in temperate waters (Bass et ah, 1975). Hexanchus agassizi Cappetta, 1976 Plate 14, figures 4-11; Plate 15, figures 1-3 Hexanchus agassizi Cappetta, 1976: 553, PL 1, figs. 5-8. Notidanus serratissimus Agassiz: Woodward, 1886: 216, PI. 6, figs. 24, 25, 26 (non fig. 23); Woodward, 1899: 6, PI. 1, fig. 7 (non fig. 6); Casier, 1966: 44, PI. 1, figs. 10-12 (non figs. 1-9). 'Notidanus* serratissimus Agassiz: Pledge, 1967: 140, PI. l,fig. 1. 'Notidanus' cf. serratissimus Agassiz: Pledge, 1967: 140, PI. l,fig. 2. 'Notidanus' ? serratissimus Agassiz: Pledge, 1967: 140, fig. 3, two top specimens only (non bottom two specimens). Fossil Material Examined. Four teeth (SAMD V34; UAGD F17262; SAM P19552, P19643) Vand a dozen broken teeth and fragments (SAMD V60; SAM P19552). Occurrence. Knight Formation equivalents, E and WS, Naracoorte No. 5 Bore, 135-145 m, Naracoorte, South Australia; Blanche Point Marl at Blanche Point and Port Noarlunga, South Australia. Age. Knight Formation equivalents, Middle Eocene; Blanche Point Marl, Upper Eocene (Pledge, 1967). Diganosis. Teeth up to 23 mm long, very similar to those of the extant H. vitulus but with the primary cusp nearer in size to the dis- tal crownlets. The primary cusp of the uppers may be more attenuated and sinuous than in H. vitulus. Description. One tooth (PL 15, fig. 2 = Pledge, 1967, PL 1, fig. 1) shows clearly the main characters of H. agassizi: the crownlets which decrease evenly in size distally from the slightly larger primary cusp with its finely ser- rated mesial margin; the slight lingual con- cavity of the tooth when viewed incisally. The length of the specimen is twice that stated in Cappetta's (1976) diagnosis. As with Cap- petta (1976) upper teeth are poorly represented in the collection on hand. One almost complete tooth (PL 14, fig. 4) possesses the slender, sinuous primary cusp with a serrated mesial margin and the second distal crownlet, the first crownlet being lost. The presence of only 2 crownlets suggests that the tooth is from the first or second row of the laterals. The two incomplete crowns (PL 14, 68 NOEL R. KEMP figs. 5, 6) show that the difference in size be- tween the primary cusp and the first crownlet is less marked than in the equivalent teeth of the extant H. vitulus. A broken lower tooth possessing only the primary cusp, first crownlet and part of the root (PI. 14, figs. 10, 11) has one denticle on the base of the medial margin of the first crownlet. This feature has not been seen before in an hexanchid tooth and probably represents an individual variation. A well preserved tooth (PL 15, fig. 1 = Pledge, 1967, PL 1, fig. 2) differs slightly from other teeth o/ H. agassizi in having a relatively broad and more upright primary cusp. The length of the mesial margin of this cusp is not great enough for the tooth to be an upper lateral and this with the large number of crownlets — 10 — suggests a lower lateral. Further material may show such differences to be within the range of normal variation of the species. Discussion, The diagnosis by Cappetta (1976) for his H. agassizi from the London Clay states that the teeth do not exceed 1 3 mm in length and that they have from 7 to 10 crownlets including the principal cusp. A tooth from the London Clay figured by Wood- ward (1886, PL 6, fig. 25) under the name Notidanus serratissimus is 1 8 mm in length and has 1 1 crownlets. Two teeth from South Aus- tralia figured by Pledge (1967, PL 1, figs. 1, 2) as 'Notidanus' serratissimus and W\ cf. serratissimus, respectively, both exceed 20 mm in length and have 12 crownlets each. These three specimens are included in Cappetta's (1976) synonymy but the discrepancies in size and number of crownlets are not discussed. The two South Australian localities of Mid- dle and Upper Eocene age are stratigraphically younger than the London Clay which is Lower Eocene. An increase in tooth size of a taxon as the geological age of the specimens de- creases has been demonstrated before in sharks' teeth e.g. in Odontaspis macrota (Agas- siz) from the Lower Tertiaries of Europe and Angola, Darteville and Casier (1943) and Ca- sier (1946, 1966). The South Australian teeth are morphologically similar to those from the London Clay except for their greater size and an increase in the number of crownlets — up to 12 compared with 7 to 10 in the Lower Eo- cene teeth. This increase is to be expected as such allometric growth patterns are seen in ex- tant hexanchids (see H. griseus, above). Until specimens of either Lower Eocene age from Australia or Middle and Upper Eo- cene age from Europe, or elsewhere, become available the teeth from South Australia should be included in Cappetta's //. agassizi Addi- tional specimens may show the existence of two fossil species, which, like the dentition of the two extant Hexanchus species, are separ- able primarily on the basis of size. Distribution. H. agassizi is found in Middle and Upper Eocene beds from South Australia and has been recorded from the Eocene of Russia (Menner, 1928; Glyckman, 1964, fide Cappetta, 1976), the Lower Eocene of the London Basin and from North America (Cap- petta, 1976). Genus Heptranchias Rafinesque, 1810 Heptranchias Rafinesque, 1810; 13; type spe- cies Heptranchias cinereus Rafinesque equals Squalus perlo Bonnaterre, 1788 (fide Bigelow and Schroeder, 1948). Generic Diagnosis Seven gill openings; lower labial furrow poorly developed; snout narrow and tapering, length always more than 1*5 x distance be- tween nostrils; horizontal diameter of eye al- ways greater than distance between nostrils. No central medial tooth in upper jaw; medial tooth in lower jaw with central cusp, which may be asymmetrical; first rows of lower laterals with from 4 to 5 or 6 crownlets; prim- ary cusp markedly bigger than distal crownlets its mesial margin with from 1 to 4 basal den- ticles. Characters otherwise those of family. Living Species One species, Heptranchias perlo, of world- wide range. Garrick and Paul (1971) have shown the Australian nominal species H. dak- ini Whitley, 1931 to be conspecific with H. perlo. They also note that some New Zealand specimens of H. perlo have 6 lateral teeth on SOUTH-EASTERN AUSTRALIAN HEXANCHID SHARKS 69 each side of the lower jaw compared with the usual 5. Specimens of H. perlo from south western Indian Ocean represent an isolated population in that they have very different ver- tebral counts from those of Australian and Western North Atlantic specimens (Bass et al, 1975). Heptranchias perlo (Bonnaterrc, 1788) Sharpsnout sevengill shark; Perlon shark (Plate 13, figure 1) Squalus perlo Bonnaterre, 1788: 10. Notidanus cinereus Cuvier, 1829; Gunther, 1870: 398; Agassiz, 1843: 218. Heptranchias perlo (Bonnaterre, 1788); Mc- Culloch, 1911: 2, PL 1, fig. 1; McCulloch, 1919: 219, PL 16, fig. 4a; McCulloch, 1929: 3; Fowler, 1941: 9; Bigelow and Schroeder, 1948: 88, Figs. 10, 11; Parrott, 1958: 86, text fig.; Applegate and Uyeno, 1968: 197, PL IB; Garrick and Paul, 1971: 1, PL 1, 2; Welton, 1974: 6, Fig. 1C, PL 1; Bass era/., 1975: 11, Fig. 7, PL 4. Heptranchias dakini Whitley, 193 la: 3 10; Fowler, 1941: 5"; Munro, 1956: 2, Fig. 4; Scott, 1962: 19, text fig.; Whitley, 1968: 5; Scott et ah, 1974: 23, text fig. (non) Heptranchias cf. perlo (Rafinesque); An- tunes and Jonet, 1969: 131, PL 4, fig. 4. Extant Material Examined. 695 mm female, Victorian coast (AMS 110825) (possible holo- type of H. dakini Whitley, 1931, see Garrick and Paul, 1971); 720 mm female, off Barren- joey, N.S.W., Jan., 1955, J. E. Smith (AMS IB3277); 778 mm female, taken at 100 m, N.E. of Cape Solander, N.S.W., 1916, State Trawl- ing Department (AMS 113929); 845 mm male, taken at 120-140 m, 100 km S. of Cape Ever- ard, Victoria, 1910, 'Endeavour' (AMS 110794 (paratype of H. dakini Whitley, 1931, see Garrick and Paul, 1971); 720 mm male, taken at 620 m, 56 km S.E. of Newcastle, N.S.W., 33° ll'S, 152° 23'E, 29 Apr., 1971, F.R.V. 'Kapala', prawn trawl (AMS 115975- 014); 810 mm female, trawled off Botany Bay, June, 1943, G. P. Whitley (AMS IB1341); 890 mm female, taken at 177 m, 32 km, S.E. Port Stephens, N.S.W., Dec., 1975, 'Kaiyo Maru' (TM D1245); 887 mm female, same data as TM D1245 (TM D1247); 870 mm male, same data as AMS 110794 (NMV El 184). Dentition Dental formula of H. perlo TM D1245 pip L8 M4L 7 P9 - The number of upper and lower P9 O mi L5 Piomedials is constant while upper laterals may vary between 7 and 9 on each side. As noted above some New Zealand specimens have 6 lower laterals on each side. Diagnosis. Teeth small with the largest — the 4th or 5th row of lower laterals — not exceeding 25 mm in length, and showing a similar hetero- donty to Hexanchus griseus but differing sig- nificantly in detail. Upper Jaw Medials. Crown more finely attenuated, much flatter labio-Iingually, more strongly sinuous and curved away from the jaw midline than in H. griseus. Similar to H. vitulus especially the arcuate distal margin, but crown more inclined and tooth much broader. Laterals. Primary cusp more slender and rela- tively much larger than the distal crownlets compared with both extant species of Hexan- chus. One or 2 small crownlets in first row in- creasing to 2 to 4 in last row. Mesial margin smooth or with small basal denticles in first row. Subsequent rows with 1 or 2, increasing to 2 or 3, may be 4, basal denticles about equal in size to distal crownlets, i.e. differing mark- edly from the upper laterals of H. griseus and //. vitulus which have a finely serrate mesial margin and a greater number of relatively larger distal crownlets. Lower Jaw Medial. Tooth with only 2 or 3 cusps on me- sial and distal margins, much less inclined than in Jl. griseus and H. vitulus. Central cusp rela- tively larger and approximately symmetrical. Laterals. Primary cusp about twice as high as first crownlet and more finely attenuated than in H. griseus and H. vitulus and primary cusp and crownlets more upright than in those spe- cies. From 4 to 5 or 6 crownlets in first row to 6 to 8 in last row; crownlets increase in size distally then gradually decrease, the last 2 or 70 NOEL R. KEMP 3 in each tooth being markedly smaller than the third or fourth last. This is contrast to the even decrease in size from the first to last crownlet in Hexanchus spp. Mesial margin with from 2 to 4 basal denticles in first rows to 1 or 2 in last; basal denticles relatively large, in last rows largest may be almost as big as first crownlet. Discussion. Bigelow and Schroeder ( 1 948 ) note a difference in the number of crownlets in the lower laterals between a male and their female specimens. In the present study material this sexual dimorphism is not apparent, the number of crownlets showing a similar range between the two sexes. The largest tooth seen in the present study is a fifth lower lateral 20 mm in length from a 1340 mm female (NZNM 2180). In a slightly longer specimen, a 1365 mm female, also from New Zealand (CM 418) the fifth lower lateral is only 17 8 mm while the fourth is 19 mm (G. A. Tunnicliffe, pers. comm.). With one ex- ception this is the largest specimen of H. perlo seen recorded in the literature. Giinther (1870: 398) notes an 'Adult male; stuffed, 7 feet long. From the Antarctic Expedition' under the name Notidanus cinereus. This specimen is actually 6 ft 8 in (2030 mm) long, has no teeth, the mouth and gill slits are sewn up and 'it is not possible to identify to species for certain due to its shocking condition' (Mr A. Wheeler, pers. comm.). In light of this Giin- ther's (1870) record for a maximum size of H. perlo is regarded as dubious. Antunes and Jonet (1969) assign a Late Miocene upper right anterior (= medial) hex- anchid tooth to Heptranchias cf. perlo. The thickness of the tooth labio-lingually separates it immediately from Rafinesque's H. perlo as does the less sinuous crown and the less curved distal margin. Garrick and Paul (1971) have shown that the differences between the nominal H. dakini Whitley and H. perlo (Bonnaterre) are due to sexual dimorphism. The main differences are in the relative position of the dorsal, pectoral and anal fins. In females the anal fin origin is below about the middle of the base of the dorsal fin, in males it is below the end of the dorsal base. Also, the pelvic fin is situated further back in females resulting in a shorter pelvic-anal dis- tance than in males (Garrick and Paul, 1971). In the following Table 1 are compared selected dimensions of one male and two female H. perlo. The latter were taken off N.S.W. (TM D1245, TM D1247) while the male was taken off southern Victoria (NMV El 184) by the Endeavour in 1910 along with the holo- and paratype of Whitley's H. dakini. Although differing in detail these data de- monstrate the same trends of fin placement characteristic of each sex as shown by Gar- rick and Paul (1971, Table 1). The two H. perlo from off Port Stephens, N.S.W. gave the following vertebral counts: TMD1247 TMD1245 Precaudal 93 91 Caudal 56 54 Total 149 145 These counts fall within the range of other Australian and New Zealand specimens of H. perlo (Garrick and Paul, 1971) and agree closely with those of three specimens from the western North Atlantic (Springer and Garrick, 1964). They differ from the average counts given by Bass et al. (1975) of 75 precaudal and 53 caudal. Distribution. H. perlo has been recorded from the western and eastern North Atlantic, the Mediterranean, the Cape of Good Hope and Japan (Bigelow and Schroeder, 1948), from Australasia (Garrick and Paul, 1971), from the eastern South Atlantic and south- west Indian Ocean (Bass et al, 1975). It appears to be a shelf-edge species as it is usually taken at depths between 100-400 m; some specimens have been taken in shallow water — about 50 m — in New Zealand. Shal- low water records are probably due to confu- sion with the broadsnout sevengill genus, Notorynchus (Garrick and Paul, 1971). Bass et al. (1975) also record H. perlo from depths of 50-400 m but note that most were taken from the deeper part of this range. SOUTH-EASTERN AUSTRALIAN HEXANCHID SHARKS 71 TABLE 1 Sex and total Catalogue no. % dorsal base Predorsal Preanal Prepelvic Pelvic origin to length anterior to anal length as length as length as anal fin origin fin origin % tl. %tl. %tl. as % tl Female 887 TMD1247 82-8 50-2 55-5 411 14-4 Female 890 TM D1245 61-6 51-1 55-6 41-9 13-7 Mean of 72-2 50-7 55-6 41-7 14-1 females 930 50-3 56-9 40-8 16-1 Male 870 NMVE1184 Heptranchias howellii (Reed, 1946) Plate 15, figures 4-7 Notidanion howelli Reed, 1946: 1, figs. 1-3, Fig. 4. 'Notidanus* ? serratissimus Ag.; Pledge, 1967: 140 (part.), PL 1, fig. 3, bottom two speci- mens (non top two specimens). Heptranchias ezoensis Applegate and Uyeno, 1968: 195, PL 1A. Heptranchias Waldman, 1971: 166, PL 1, figs. 1,2. Heptranchias howelli (Reed); Welton, 1974: 1, Fig. 1A, B, PL 2A, B. Fossil Material Examined. Four incomplete crowns (RJFJ no. 121; SAM P19573; UAGD F17284). Occurrence. Blanche Point Marl, Blanche Point, South Australia. Age. Upper Eocene. Diagnosis. Based on lower laterals, the only tooth type of this species so far identified. Teeth similar to the extant H. perlo but larger, up to 25 mm in length (Waldman, 1971) and with mesial basal denticles, primary cusp and distal crownlets all relatively broader and less attenuated than in the extant species. The primary cusp of H. perlo is always relatively higher. H. howellii may have up to 5 basal denticles, H. perlo from 1 to 4. Description. Three incomplete crowns (PL 15, figs. 4, 5, 7) represent mesial portions of teeth, each having the primary cusp and two crownlets with the second crownlet being characteristically larger than the first. The re- maining specimen (PL 15, fig. 6) is the distal portion of a tooth with the crownlets increas- ing in size distally until the last two which de- crease, the most distal being very much smaller. Figure 5 — Locality map showing occurrences of fossil hexanchid teeth in S.E. Australia. 72 NOEL R. KEMP Discussion. A tooth of H. howellii figured by Welton (1974, PL 2B) although very close morphologically to an H. perlo lower lateral, e.g. row 2 (PL 13, fig. 1 ), still has a more ro- bust shape, a primary cusp not as high and cusp and crownlets inclined and straight rather than inclined and slightly curved as found in the extant species. The other tooth (Welton, 1974, PL 2 A) is nearer to Reed's (1946, Figs. 1-4) holotype of H. howellii and similar to the four incomplete crowns from South Australia. Reed (1946) did not compare her new spe- cies with H. perlo but Welton (1974) differen- tiates H. perlo from H. howellii on the basis of the former species having a higher primary cusp and no more than 3 basal denticles on the lower laterals. Lower laterals of both left and right rows of two jaws to hand (TM D1245; TM D1247, PL 13, fig. 1) have 4 basal den- ticles on the mesial margin. The main criteria to separate H. perlo from H. howellii are the higher primary cusp in the extant species, as Welton (1974) notes, but also the generally broader and less attenuated basal denticles, primary cusp and crownlets of H. howellii, Applegate and Uyeno (1968) did not com- pare their H. ezoensis with H. howellii The figured holotype of H. ezoensis is very similar to some figured specimens of H. howellii, e.g. PL 15, fig. 7; Welton (1974, PL 2B) except that it has only 2 basal denticles and the prim- ary cusp and crownlets are more inclined. Such differences however are within the normal range of variation seen between the first and last rows of lower laterals of H. perlo (PL 13, fig. 1) and is in accord with Applegate and Uyeno's own interpretation of the tooth being referable to the fifth lower lateral row. For these reasons H. ezoensis is included in the synonymy of H. howellii. The size of teeth of H. howellii is compar- able to those of H. perlo suggesting that the fossil shark too may have grown to a maxi- mum length of less than 2 m. The type speci- men is 13 mm in length (Reed, 1946), Welton (1974) records four teeth ranging from 6 mm to 17 mm and Waldman's (1971) two speci- mens are 19 mm and 25 mm long. The four South Australian fragments are all incomplete but by comparison with the above specimens — measuring from the mesial margin to the apex of the second crownlet — they fall within this size range. Distribution. Although teeth of H. howellii are apparently rare in the fossil record — only about a dozen having been described, includ- ing the four in the present study — they are rea- sonably widespread geographically. In North America they are recorded from the Eocene of New Jersey (Reed, 1946) of Oregon and Wash- ington (Welton, 1974) and British Columbia (Waldman, 1971; Welton, 1974). The tooth from Japan is from the Upper Oligocene (Ap- plegate and Uyeno, 1968) while the South Aus- tralian specimens are of Upper Eocene age. Genus Notorynchus Ayres, 1855 Notorynchus Ayres, 1855: 76; type species Notorynchus maculatus Ayres, 1855, Califor- nia. Generic Diagnosis Seven gill openings; spiracles small; lower labial furrows well developed; snout broadly rounded, length always less than 15 x distance between nostrils; horizontal diameter of eye 2 x or 3 x distance between nostrils; dorsal surface of body and paired fins with small dark splotches, white spots may or may not be present. Central tooth in both upper and lower jaws, except in some specimens, e.g. off California, which have a variable number of medials either on or either side of the upper symphysis (see below); lower medial without central cusp or, if present, small and strongly asymmetrical; lower laterals with 4 to 7 distal crownlets. Characters otherwise those of family. Living Species Possibly only one species Notorynchus cepe- dianus (Peron, 1807) of world-wide range. The taxonomic status of the group at the sub- generic level is particularly confused. N. macdonaldi Whitley (1931b) was later included by Whitley (1934) himself in the synonymy of N. cepedianus when he found than Peron's species had priority. Bass et ah (1975) note that the white spots on the dorsal SOUTH-EASTERN AUSTRALIAN HEXANCHID SHARKS 73 surface of JV. macdonaldi described by Whit- ley (1931) had not been reported on further specimens. In fact this coloration seems to be quite widespread as it has been recorded from a number of localities such as Bass Strait (Mac- donald and Barron, 1868); New Zealand (Phil- lips, 1924) and eastern Pacific (no detailed localities given) (Kato et at., 1967). The size of the spots is about that of the eye (Whitley, 1931B; Kato et aL, 1967: 6). On the two local specimens caught recently (TM D1291 and TM D1292) the white spots were ran- domly scattered over the dorsal surface and ranged from 7-30 mm in diameter. Macdonald and Barron (1868) suggested that the spots on their specimen might be the result of disease. An examination of the white spots on the two local specimens revealed no pathological dis- turbance (B. Munday, pers. comm.) The presence or absence of a central cusp on the lower medial tooth, the degree of serration of the mesial margin of the lower laterals and the presence or absence of a medial tooth on the symphysis of the upper jaw have been the main criteria used in differentiating the nom- inal species of Notorynchus by a number of authors, for example Giinther (1870), Garman (1913), Fowler (1941). The first two charac- ters are variable within the species (see below) but the presence or absence of a tooth on the symphysis of the upper jaw is a difference whose taxonomic status has yet to be estab- lished. Many authors such as Agassiz (1835), Miiller and Henle (1841), Macdonald and Barron (1864), Garman (1884), Lahille (1928), Whitley (1931), Sadowsky (1970) and Bass et al. (1975) either figure or describe in detail the presence of a central upper medial tooth with a single medial on each side. The issue is confused by some authors, for example Garman (1913), Fowler (1941), who include in the synonymies of their species of Notorynchus supposedly lacking an upper cen- tral medial, references which actually figure this tooth, for example Agassiz (1835), Miiller and Henle (1841). In addition Fowler (1941) includes a number of Australian references in his synonymy of Heptranchias cepedianus (Peron), the species supposedly lacking an up- per central medial. All Australian specimens so far described in the literature and specimens seen by the present writer possess this tooth. Fowler's single example, H. cepedianus, from southern Africa ('Cape Colony coast', Fowler, 1941: 7) was actually a skinned out specimen, thus presumably the jaws were not examined. Bass et al. (1975) note that all southern Afri- can specimens do possess a tooth on the upper symphysis. Tenore's (1810) description of his new species, Squalus platycephalus, from the Mediterranean, gives no clear indication of the number or placement of teeth in the symphyseal area of the upper jaw, 'Superior prominentia intermedia dentibus decern triplici serie dispositis . . .' (Tenore, 1810: 258). Two sets of jaws of the broadsnout seven- gill shark from off the Californian coast re- cently received at the Tasmanian Museum (see below) have a pair of medials on either side of the upper symphysis. The type of Notorynchus maculatus Ayres, 1855 is from California. However, in the description of the teeth of the upper jaw of the type it is not clear whether there is a tooth on the symphysis or not 'Those of the center are narrow, acute, without den- ticles at the base . . .' (Ayres, 1855: 76-77). Some Californian specimens of Notorynchus do have an upper central medial but a range of from 0-2 on each side of the symphysis is known (B. J. Welton, pers. comm.). In the des- cription of tooth types (see below) only the more common condition of three upper me- dials with one on the symphysis is included as the writer has not seen the complete range of variation present in the Californian speci- mens. Further detailed comparative studies may show the dental variation of the upper symphyseal area of Notorynchus from Cali- fornia and perhaps the Northern Hemisphere in general to be of specific value. If a second spe- cies is established and clearly defined only then can the synonymies of this genus be unravelled. Heptranchias haswelli Ogilby, 1897 was erected on the basis of a set of jaws then in Macleay Museum University of Sydney. Ma- terial in this Museum was eventually handed over to the Australian Museum but the type jaws are now lost (J. R. Paxton, L. Bushell, 74 NOEL R. KEMP pers. comm.). From the description (Ogilby, 1897) the jaws definitely were of hexanchid (see below) but its status must now remain as a species inquirenda. Notorynchus cepedianus (Peron, 1807) Broadsnout sevengill shark (Plate 13, figure 2) Squalus cepedianus Peron, 1807: 337. Squalus platycephalus Tenore, 1810: 241, 258. Notidanus indicus Agassiz, 1835, PL E, fig. 1; Agassiz, 1838: 92; Agassiz, 1843: 217; Giin- ther, 1870: 398; Hutton, 1873: 271; Day, 1878; 723, PI. 189, fig. 4; Johnston, 1882: 138; Johnston, 1890: 38. Heptanchus indicus Miiller and Henle, 1841: 82, PL 32. Notorynchus maculatus Ayres, 1855: 72; Gill, 1862: 495; Herald, 1968: 412. Notorynchus borealis Gill, 1864: 150. Heptranchus indicus McDonald and Barron, 1868: 371, PL 32, figs. 1-6; Castlenau, 1872: 217;Haswell, 1880: 96;HasweIl, 1884a: 88, PL 1, fig. 5; Haswell, 1884b: 381, PL 10, fig. 1,2; Ogilby, 1889: 179. Heptranchus griseus McDonald, 1873: 312. Notidanus (Heptanchus) indicus McCoy, 1880: 16, Fig. A, B, PL 43, fig. 2. Heptranchias pectorosus Garman, 1884: 56; Lahille, 1928: 299, Figs. 1, 2; Fowler, 1941: 7. Notorhynchus maculatus Jordan and Ever- mann, 1896: 17; Jordan and Evermann, 1900, PL 2, fig. 7; Welton, 1974: 3, Fig. ID. Heptranchias indicus; Waite, 1907: 6; McCul- loch, 1911:2. Notorhynchus indicus; Zeitz, 1908: 289. Heptranchias platycephalus (Tenore, 1810); Lahille, 1928: 300, 302. Notorynchus platycephalus Garman, 1913: 18. Heptranchias spilotus Lahille, 1913: 26, Figs. 1-3, PL 8, fig. 1. Notorhynchus pectorosus; McCulloch, 1919: 219, PL 6, fig. 3a; Waite, 1921: 10, Fig. 5; Waite, 1923: 24, three Figs; Phillips, 1924: 259, Fig. 1. Notorynchus griseus (McDonald, 1873); Mc- Culloch, 1929: 3. Notorynchus macdonaldi Whitley, 1931b: 138, PL 20, figs. 3-5; Phillips, 1935: 236, Fig. 1. Notorynchus cepedianus; Whitley, 1934: 181, 197; Whitley, 1940: 70, Figs. 4, 50, 51; Clemens and Wilby, 1946: 51, Fig. 11; Gra- ham, 1953: 62, one Fig.; Scott, 1962: 19, one Fig.; Whitley, 1968: 5; Scott et al., 1974: 23, three Figs; Bass et aL, 1975: 14, Fig. 8, PL 5. Heptranchias cepedianus; Fowler, 1 941 : 6; Parrott, 1958: 84, one Fig. Notorhynchus cepedianus; Munro, 1956: 2, Fig. 5. Notorhynchus pectorosus; Garman, 1913: 20; Sadowsky, 1970: 33, Fig. 1. (non) Heptranchious haswelli; Ogilby, 1897: 62. Extant Material Examined. 555 mm female, Altona, Vict. (NMV A106); 2391 mm female, mounted specimen (NMV); dried jaws, no data (NMV); 910 mm male, State Fisheries, N.S.W., 1920, no data (AMS 16813); head of specimen taken off Babel Is., N.S.W. at 130 m, 26 June, 1911 (AMS E2161); dried jaws, no data (MUGD); 1520 mm male, Storm Bay, S.E. Tasmania, Safcol (Tas.) Pty. Ltd., May, 1977 (TM D1291); 1648 mm male, same data as TM D1291 (TM D1292) flaws only of these two specimens retained); dried jaws, 1310 mm immature female, Berkeley Flats, San Fran- cisco Bay, California, U.S.A., B. J. Welton, June, 1976 (TM D1302); dried jaws, 2180 mm immature female, California, U.S.A., B. J. Welton, Jan., 1977 (TM D1303). Fossil Material Examined. One incomplete tooth (TMZ1991). Occurrence. Jemmy's Point road cutting, 5 km E of Lakes Entrance, Victoria. Age. Early Pliocene (Abele et al., 1976). Dentition Dental formula of dried jaws, MUGD (PL 13, fig. 2) :«m£fJf|«. The number of me- dials and laterals in both jaws is in most spe- cimens usually constant (except in Californian specimens, see above), but some variation does occur in the laterals, e.g. Bass et aL (1975) record one specimen with 7 upper laterals on one side. The 555 mm male from Victoria has SOUTH-EASTERN AUSTRALIAN HEXANCHID SHARKS 75 7 lower laterals on the right but the usual 6 on the left. As in Hexanchus the number of crownlets, especially in the lower laterals and the degree of denticulation of the mesial mar- gin of all teeth increases with age, and thus the size of the tooth. On the mesial margin of teeth of a given size the basal denticles of Notorynchus are larger and more developed than are the serrations of Hexanchus. The ba- sal denticles of Heptranchias are relatively larger again than those of Notorynchus. In both Notorynchus and Heptranchias the basal den- ticles extend only half to one-third along the mesial margin, in Hexanchus the serrations extend along about two-thirds. The basal den- ticles of Heptranchias are relatively larger again than those of Notorynchus. Diagnosis. Teeth small with the largest — usually the 2nd lower lateral row — probably not ex- ceeding 25 mm in length, and showing a similar heterodonty to Hexanchus griseus but differing significantly in detail. Upper Jaw Medials. Three rows, the central one of which is on the symphysis and is usually upright but may be slightly inclined to either the right or left. Serrations or basal denticles not usually present on margins of central medial. Basal denticles present on both margins of the two lateral medials but may be absent in juvenile specimens. Crown of medials of Heptranchias perlo much more slender and inclined than in Notorynchus cepedianus. Root more or less square or slightly tapering basally in outline when viewed labially. Laterals. Teeth in first rows a little higher than broad ranging through to a little broader than high in last rows. Teeth of Hexanchus are lower and broader in comparison. Primary cusp similar to H. griseus but with fewer crown- lets distally, ranging from only 1 in first row to 4 to 5 or 6 in the last row. Primary cusp and crownlets all relatively broader and larger than in Heptranchias perlo. Lower Jaw Medial. Tooth similar to that of Hexanchus griseus but usually lacking a central cusp. Medials of some specimens, not necessarily large or small, may be quite asymmetrical hav- ing one or two more cusps on one side than the other and a strongly oblique central cusp. Laterals. Teeth about constant breadth and only one-third to one-half broader than high from first to last row. In Hexanchus they are about twice as broad as high while in Heptran- chias perlo they range from a little broader than high in the first row to more than twice as broad as high in the last. Primary cusp with 4 to 6 or 7 basal denticles on mesial margin and 4 to 5 or 6 crownlets which decrease evenly in size distally. Both extant Hexanchus species have a greater number of crownlets, from 6 to 12, depending on tooth size, and a serrated mesial margin. Heptranchias perlo has about the same number or slightly more than in Notorynchus cepedianus but they increase in size distally in the former before decreasing, and then rapidly in the last 1 to 3 crownlets. The primary cusp of H. perlo is relatively higher and more attenuated than in the other three species of the family. Description. Table 2 gives the measurements as per cent of total length of two male topotypes of N. cepedianus. Peron's specimen was from Adventure Bay, a bay on Bruny Island which is in Storm Bay, SE Tasmania. No detailed measurements of Australasian material has previously been published to the author's knowledge. Both specimens were light grey above grading to off-white below with nu- merous dark grey to black splotches about 2-10 mm across on the dorsal surface and paired fins. Also on the dorsal surface of each shark were about a dozen white spots from 7-30 mm diameter, randomly distributed. Dental formula: TM D1291 g % ™ ^ g 1 TMD1292 p9 3 l6mil6p1o - The lower medial of TM D1291 is symmetrical with no central cusp while in TM D1292 the lower medial is asymmetrical with a very small oblique cen- tral cusp. Discussion. Ogilby (1897) placed his new species of hexanchid in the genus Heptranchias, H. haswelli, on the basis of a central cusp, in- clined to the right, on the lower medial tooth. He also described in the upper jaw: 76 NOEL R. KEMP Total length Snout tip to: Nostrils: Eye: Mouth : TABLE 2 Dimensions of Notorynchus cepedianus as per cent of total length outer nostrils mouth eye spiracle 1st gill opening 4th gill opening 7th gill opening pectoral origin pelvic origin dorsal origin anal origin upper caudal origin distance between inner ends length horizontal diameter breadth height lower labial furrow Gill openings, length of: Pectoral fin: Pelvic fin: I>orsal fin: Anal fin: Caudal fin: Interspace between bases of: Distance from origin to origin: 1st 4th 7th outer margin inner margin distal margin length of base origin to lateral lobe origin to median tip length of claspers length of base length of rear tip vertical height length of base length of rear tip vertical height length of upper margin length of lower margin base of notch to tip dorsal and caudal anal and caudal pectorals and pelvics pelvics and anal Male Male TM D1291 TMD1292 1520 mm 1648 mm 1*3 1-6 3-8 3-8 4-1 4*4 10-2 10-7 13 2 13-9 15-3 16 3 160 171 160 171 41*5 40-1 49-2 48-3 54-9 54-3 66-5 66-2 4-4 4-4 11 1-2 1*3 1-3 12-7 11-8 6-5 6*3 4-4 4-2 5-6 5-4 4-9 4-4 2-8 3-2 13-7 13 5 5-7 5-7 111 11-4 7-6 7-8 5-7 6-4 8-9 10-4 5 6 6-4 6 6 7-7 2-0 2-3 4 1 4-3 4-5 6 1-6 1*9 2-4 2-7 33-7 34-5 8-7 9-4 7-5 6-8 10-3 10-6 70 6-0 25-5 23-1 15-5 14-2 3 medial teeth, the outer 2 with basal denticles on both margins, the central tooth with entire margins; 8 rows of laterals with an increasing num- ber of basal denticles and crownlets from symphysis to articulation, the last 2 rows being lower and broader than the first rows; • 10 rows of posteriors on each side; in the lower jaw: SOUTH-EASTERN AUSTRALIAN HEXANCH1D SHARKS 77 • a medial tooth with an inclined strong central cusp and 4 lateral secondary cusps on each side; • lateral teeth, the number of which is not given, with 5 to 7 basal denticles on the mesial margin of the primary cusp, fol- lowed by 5 to 6 crownlets which de- crease regularly in size distally; •. 10 rows of posteriors on each side. Except for the inclined, strong, central cusp on the lower medial and 8 upper laterals on each side the description is that of Notorynchus cepedianus. Heptranchias perlo does and both Hexanchus species may have a strong central cusp on the lower medial tooth. An asymmet- rical lower medial in N. cepedianus may have a central inclined cusp but in comparison with Hexanchus and Heptranchias the central cusp could not be called 'strong' (Ogilby, 1897: 63). Eight upper laterals have not been recorded be- fore from N. cepedianus but 8 or 9 are com- mon in both Hexanchus griseus and Heptran- chias perlo. Because the type jaws are lost //. haswelli must then remain a species inquirenda. An abnormality in the form of bifid teeth (PL 14, fig. 3) is noteworthy in that in each case the two 'teeth' both show the basic mor- phology of N. cepedianus teeth. Peyer (1968: 42) tends to 'deny the deeper morphological significance of the presence or absence of ac- cessory cusps'. The anlage, in this case of the first lateral row of the lower jaw of this speci- men of N. cepedianus, did continue to produce teeth of a morphology characteristic of this species including the basal denticles and the crownlets which decrease evenly in size distally. The mesial 'tooth' has 4 basal denticles on the mesial margin of the primary cusp which is a little larger than the remaining 3 crownlets. The distal 'tooth' which is about one-third the width of the mesial portion, has only 1 basal denticle on the mesial margin of the primary cusp which is noticeably larger than the 3 crownlets. Gudger (1937) noted similar abnor- malities in other species of sharks, e.g. Galeo- cerdo cuvieri. The causative mechanism may be a foreign object such as a fish spine being imbedded in the anlage which is subsequently divided as each tooth advances. The largest definite record of Notorynchus cepedianus seen in the literature is that given by Phillips (1935) of a 2888 mm— '9 ft 5-5 in' (Phillips, 1935: 236) — specimen taken in Oriental Bay, New Zealand; the size of the teeth is not recorded. The 2nd lower lateral of the 2391 mm mounted specimen in the Na- tional Museum of Victoria is 22 mm in length. The dental sexual dimorphism in JV. cepedi- anus illustrated and noted by Macdonald and Barron (1868) and briefly reiterated by Phil- lips (1935) has not been seen in any other lit- erature nor in any of the present study material. Indeed, the hand-drawn illustration by Mac- donald (Macdonald and Barron, 1868, PI. 33, fig. 3a, 3d) of the teeth of a female N. cepedia- nus, with the basal denticles of the mesial mar- gins of the lower laterals nearly equal in size to the primary cusp and crownlets, is more akin to teeth of the Upper Cretaceous N. pectinatus (Agassiz, 1843: 221; Agassiz, 1844, PI. 36, fig. 3a; Applegate, 1965b, Figs. 1, 2) than to Pe- ron's extant species and must be regarded as a subjective interpretation of their specimen. The Early Pliocene tooth (TM Z1991) repre- senting a lower lateral from the 2nd or 4th row is, to the author's knowledge, the first fossil occurrence of this species. Although the basal denticles and the first two crownlets are miss- ing, the tooth, on the basis of the shape of the root and the relative proportions and curvature of the primary cusp and remaining crownlets, can be definitely assigned to the extant species. N. cepedianus and N. primigenius (see below) were contemporaneous species, but until further fossil specimens of the former species are recognized discussion of their relative geo- graphical distribution would be speculative only. Distribution. Notorynchus cepedianus is re- corded from the Pacific Ocean, California to British Columbia (Clemens and Wilby, 1946) and British Columbia to Chile, excluding the tropics (Kato et aL y 1967), from Japan and Taiwan (Matsubua, 1936 and Chen, 1963, re- spectively, fide Bass et a/., 1975), from New Zealand (Phillips, 1935) and from Australia, but only from southern waters, e.g. McDonald and Barron, 1868, McCulloch, 1919, Waite, 78 NOEL R. KEMP 1921, Scott et ah, 1974, no specimens having ever been recorded from northern waters. J. D. Ogilby (fide Whitley, 1931) states that N. cepe- dianus does not occur in Queensland waters; the Atlantic Ocean, from Argentina (Lahille, 1928) and Brazil, but not from other parts of the Atlantic (Sadowsky, 1970) and the Indian Ocean (Day, 1878). This report by Day, of a specimen taken off Madras appears to be the most tropical occurrence (about 14° N) of N. cepedianus. The broadsnout sevengill shark is commonly taken in shallow coastal waters in South Africa (Bass et at., 1975) and Australia. Phillips (1935) records a 3 m female being taken off a swimming beach in Wellington, New Zealand, but also notes that it is an open ocean shark sometimes entering harbours and inlets. Herald and Ripley (1951) report that while smaller specimens, less than 18 m, are com- mon in the shallower waters of San Francisco Bay, larger specimens live in deeper water out of the Bay. Notorynchus primigenius (Agassiz, 1835) (Plate 15, figures 8-10) Notidanus primigenius Agassiz, 1835, PL 27, figs. 6-8, 13-17 (non figs. 4, 5); Agassiz, 1843: 218; Woodward, 1886: 216, PI. 6, figs. 19-22; Davis, 1888: 33, PL 6, fig. 6; Leriche, 1905: 207, Fig. 62; Chapman, 1914: 271; Leriche, 1926: 388; Leriche, 1927: 8, PL 1, fig. 1; Leriche, 1957: 22, PL l,figs. 1-6 (synonymy). Notidanus marginalis Davis, 1888: 34, PL 6, fig. 8 (non fig. 7); Chapman, 1914: 268, 271, Fig. 130A; Chapman, 1918: 4, PL 6, fig. 8 (?PL 9, fig. 1). Hexanchus cf griseus (Bonnaterre); Antunes and Jonet, 1969: 130 (part.), PL 4, fig. 3 (non figs. 1, 2). Hexanchus primigenius (Agassiz) ; Cappetta, 1970: 16, PL 4, figs. 11-19. Fossil Material Examined. Four incomplete crowns (TMZ 1992; NMV P27410, P27411, an unnumbered fragment). Occurrence. Batesford Limestone, Batesford, Vict. Muddy Creek Marl, Clifton Bank, Mud- dy Creek, Hamilton, Vict. Age. Batesford Limestone, Lower Miocene; Muddy Creek Marl, Middle Miocene (Abclc etaL, 1976). Diagnosis. Teeth similar to N. cepedianus but larger in size, up to 30 mm (Leriche, 1910), and with the primary cusp and crownlets gene- rally straightcr and more erect and broader and more robust. Description. Two specimens (PL 15, figs. 8, 9) represent lower lateral teeth, a third speci- men (PL 15, fig. 10) possibly a lower lateral while a fourth, of only 2 crownlets, cannot be placed in the jaw. One incomplete crown (PL 15, f\g. 10) consists of a primary cusp with 2 basal denticles — a fracture surface suggests that there were more — and the first crownlet Both cusp and crownlet are relatively broad, not markedly attenuated and with the cusp about half as high again as the crownlet. The other incomplete crown (PL 15, fig. 8) consists of only the primary cusp with 4 basal denticles on the mesial margin. The largest basal den- ticle is nearly twice the size of the first. The distal portion of a tooth (PL 15, fig. 9) has 5 crownlets which evenly decrease in size distally. The root is flat labially with an angular longi- tudinal ridge on the lingual face about two- thirds up from the base; basal margin very thin labio-lingually. Discussion. The relatively large, broad pri- mary cusps and crownlets differentiates these fragments from those of both Hexanchus and Heptranchias. They can be further separated from Hexanchus by the relative size difference of the primary cusp and first crownlet; the presence of basal denticles on the mesial mar- gin and cusps and crownlets being more or less straight. In contrast, in Hexanchus, the size of the primary cusp is nearer that of the first crownlet; the mesial margin is serrated rather than denticulated and the cusps and crownlets are slightly curved. Additional features sep- arating the fragments from Heptranchias are the smallest basal denticle being only about half the size of the biggest and crownlets evenly decreasing in size distally. In contrast, in Hep- tranchias, the smallest basal denticle is only about one-fifth the size of the biggest and the SOUTH-EASTERN AUSTRALIAN HEXANCHID SHARKS 79 last one or two crownlets decrease rapidly in size from the third or fourth last. The shape and dimensions of the 2 crown- lets of the fourth specimen are conspecific with the other four fragments. All the fragments differ from the extant N. cepedianus in their more erect cusp and crownlets. On the basis of the broad, robust and only slightly oblique cusp and crownlets the specimens are assigned to Agassiz's N. primigenius. Chapman's (1918) emended description of the lectotype of Noti- danus marginalis Davis, 1888 is based on an incomplete lateral tooth of a notorynchid. In the figure of the lectotype the primary cusp, which is missing, has been drawn in as a dotted outline, but of a slightly smaller size than the first crownlet. It appears that this interpretation forms the basis for his differentiation (Chap- man, 1918: 5) 'N. marginalis differs essentially in the almost equal size of the two anterior cones, those of N. primigenius and N. serra- tissimus being graduated'. Such a character is not found in any hexanchid and as all the characters of the lectotype — the basal denticles showing a small size range and the 6 relatively broad, robust, slightly oblique crownlets de- creasing evenly in size distally — are typical of Notorynchus primigenius, Notidanus margin- alis is here included in that species. Distribution. Notorhynchus primigenius is a widespread fossil species ranging from the Middle Eocene through to the Pleistocene. In Europe it is recorded from the Middle Eocene, Lower Oligocene, Miocene and Pliocene of Belgium (Leriche, 1905, 1910, 1926), from the Miocene of France (Leriche, 1957, Cap- petta, 1970), from the Upper Miocene of Por- tugal as Hexanchus cf griseus (Antunes and Jonet, 1969) and from the Middle Eocene, Miocene and Pleistocene of England (Wood- ward, 1886). Davis (1888) and Chapman (1918) note its presence in the Miocene of New Zealand under the name Notidanus mar- ginalis. The occurrence of Notorynchus primigenius in the Lower Miocene Batesford Limestone and the Middle Miocene Muddy Creek Forma- tion is the first record of the species from the Australian Tertiary deposits. Addendum A jaw of a notorynchid shark (specimen with no number labelled "Cook Strait, Hexan- chus raberi, 9.5.1944") seen recently in the National Museum, Wellington, New Zealand has the dental formula; P10— L7— M4— L7— P14. P11_L6— Ml— L6— P10 This occurrence of a Notorynchus cepedianus with 4 medial teeth in the upper jaw, outside the Californian coast region, does suggest that such a difference may be due only to individual variation within the species. A fossil tooth from South Australia recently to hand and referable to Hexanthus agassizi Cappetta has been noted by Pledge (pers. coram.; Pledge, N.S., 1977. Metasqualodon harwoodi (Sanger, 1887. — A rediscription. Rec. S. Aust. Mus. Adelaide, 17 (17): 285- 297, 3 Figs). The tooth (SAM P10867) from the River Murray Cliffis near Wellington is a lower lateral, 20 mm long and 9-4 mm high with 10 crownlets which decrease evenly in size distally and a primary cusp with a den- ticulated mesial margin. Pledge (1977) shows it to be from the Ettrick Formation which is of Late Oligocene age. This then is the youngest occurrence of H. agassizi so far on record. Acknowledgements The author wishes to thank Dr O. P. Single- ton, University of Melbourne for suggesting the initial project which was the basis of a M.Sc. thesis under his supervision and from which the present study arose. The following people greatly assisted by allowing access to or loan of specimens in their care: Miss J. M. Dixon, National Museum of Victoria; Mr T. F. Flannery, Beaumaris, Victoria; Mr J. M. Lind- say, Department of Mines, South Australia; Dr J. R. Paxton, Australian Museum, Sydney; Mr N. S. Pledge, South Australian Museum, Adelaide and Dr T. H. Rich, National Mu- seum of Victoria. Appreciation is expressed to those who donated specimens, photographs or supplied information especially Dr J. A. Bass previously Oceanographic Research Institute, Durban, South Africa; Dr H. Cappetta, Lab- 80 NOEL R. KEMP oratoire de Paleontologie, Montpellier, France and Mr B. J. Welton previously University of California, Berkeley, U.S.A. and now Natural History Museum, Los Angeles, U.S.A. as well as Mrs L. Bushell, Macleay Museum, Univer- sity of Sydney, N.S.W.; Mr M. J. Cooper, Uni- versity of Melbourne; Mr T. A. Darragh, National Museum of Victoria; Professor J. A. F. Garrick, Victoria University of Wel- lington, New Zealand; Dr. D. F. Hoese, Aus- tralian Museum, Sydney; Mr J. M. Morland, National Museum of New Zealand, Wellington; Mr S. Springer, previously Bureau of Commer- cial Fisheries, U.S.A.; Dr V. G. Springer, United States National Museum, Washington; Mr G. A. Tunnicliffe, Canterbury Museum, Christchurch, New Zealand; Mr D. Ward, London, U.K.; Mr B. F. Webb, Marine Re- search Laboratories, Taroona, Tasmania and Mr A. Wheeler, British Museum (Natural His- tory), London. Radiographs of two sharks were kindly made by Dr A. J. Mcintosh, Hobart, Tasmania and Mr B. Munday, Department of Agriculture, Launceston, Tasmania carried out the pathological examination of the shark skin. Thanks are also due to Mr R. F. Durham, Black Rock, Victoria for his constant enthu- siasm and assistance in the initial stages of the photography. Acknowledgement is gratefully made to the Director and Trustees of the Tas- manian Museum for the use of the Museum's time and facilities. References Abele, C. et al., 1976. Tertiary. In Douglas, J. G. and Ferguson, J. A. (Eds.) Geology of Victoria: i-xi, 1-528. Geological Society of Australia, Special Publication No. 5. Melbourne. Agassiz, L. J. R„ 1833-1844. Research sur les pois- sons fossiles. 3: i-viii, 1-390, Atlas: 1-34, 83 pis. Neuchatel. Antunes, M. T. and Jonet, S., 1969. Requins de l'Helvetien superieur et du Tortonien de Lis- bonne. Rev. Fac. Cienc. Lisboa, (2, C) 16 (1): 119-280, 14 Figs., 20 pis. Applegate, S. P., 1965a. Tooth terminology and variation in sharks with special reference to the sand shark Carcharias taurus Rafinesque. Contr. Sci., Los Angeles Co. Mus. 86: 1-18, 5 Figs. , 1965b. 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Paris: i-vi, 1-215, 102 pis. Cappetta, H., 1970. Les selaciens du Miocene de la region de Montpellier. Mem Ext. Palaeoverte- brata, Montpellier: 1-139, 22 Figs.: 1-19, 27 pis. , 1975. Selaciens et Holocephale du Gar- gasien de la region de Gargas (Vaucluse). Geol. Mediterr., Aix-en- Provence, 2(3): 115-134, 10 Figs., 2 pis. , 1976. Selaciens nouveaux du London Clay de l'Essex (Ypresien du Bassin de Londres). Geobios, Lyon, 9(5): 551-575, 1 Fig., 4 pis. Casier, E., 1966. Faune ichthyologique du London Clay. Brit. Mus. (Nat. Hist.): 1-496, 82 Figs., 68 pis. Castelnau, F. L., 1872. Contribution to the ichthyo- logy of Australia. Proc. Zool. Acclim. Soc. Vict. 1: 29-248. Chapman, F., 1914. Australasian Fossils: 1-341, 150 Figs. Melbourne. , 1918. Descriptions and revisions of the Cretaceous and Tertiary fish-remains ol New Zealand. Palaeont. Bull., Wellington, 7: 1-45, 9 pis. Clements, W. A. and G. V. Wilby, 1946. Fishes of the Pacific Coast of Canada. Bull. Fish. Res. Bd. 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Fish Wildl. Serv. Wash. Ill: 1-47, 75 Figs. Kemp, N. R., 1970. Studies on the dentition of Aus- tralian Tertiary and Recent sharks. Unpublished M.Sc. thesis, University of Melbourne: 1-354, 63 pis. Lahille, F., 1913. Nota sobre los generos Heptran- chias y Hexanchus. An. Mus. nac. B. Aires. 18: 25-32. 3 Figs., 2 pis. , 1928. Nota solore unos peces elasmobran- quios. An. Mus. nac. B. Aires 34: 299-340, 27 Figs., 5 pis. Leriche, M., 1905. Les poissons eocene de la Bel- gique. Mem. Mus. r. Hist. nat. Belg., Bruxelles, 3(11): 49-228, 9 pis. , 1910. Les poissons oligocenes de la Bel- gique. Mem. Mus. r. Hist. nat. Belg., Bruxelles, 5: 229-363, 27 pis. -, 1926. Les poissons neogene de la Belgique. Mem. Mus. r. Hist. nat. Belg., Bruxelles, 32: 368-472, 14 pis. -, 1927. Les poissons de la Molasse Suisse. Abh. Schweiz. palaont. ges., Basel, 46: 1-56, 7 pis., 47: 57-120, 7 pis. -, 1957. Les poissons neogenes de la Bretagne de l'Anjou et de la Touraine. Mem. Soc. geol. Fr., Paris, (5) 81: 1-64, 4 pis. Lynch, D. D., 1963. First Australian record, Hexan- chus griseus (Bonnaterre, 1780). The Six-gilled Shark. Mem. Natn. Mus. Vict. 26: 259-260, 1 pi. McCoy, F., 1880. Prodromus Zool. Vict., Melbourne 5: 1-58, 10 pis. McCulloch, A. R., 1911. Report on the fishes ob- tained by the FJ.S. 'Endeavour* on the coasts of New South Wales, Victoria, South Australia and Tasmania. Zool. Res. Endeavour 1(1): 1-87, 26 pis. , 1919. Check-list of the fish and fish-like animals of New South Wales. Part i. Austr. Zool. 1(7): 217-229, 3 pis. , 1929. A check-list of the fishes recorded from Australia. Aust. Mus. Mem. 5(1): 1-144. MacDonald, J. D., 1873. A communication . . . containing descriptions of two sharks taken off Flinders Island, Bass's Straits, during the voy- age of H.M.S. 'Herald' in the South Seas. Proc. Zool Soc. (Lond.): 312. Macdonald, J. D. and C. Barron, 1868. Notes on Heptranchus indicus, chiefly as regards sexual characters. Proc. Zool. Soc. (Lond.): 371-373, lpl. Muller, J. and F. G. Henle, 1841. Systematische Beschreibung der Plagiostomen. Berlin: i-xxii, 1-200, 60 pis. Munro, I. S. R., 1956. Handbook of Australian fishes. Fisheries Newsletter, Australia, 1 : 1-8, 56 Figs. Ogilby, J. D., 1889. List of the Australian Palaeich- thyes, with notes on their synonymy and distribu- tion. Proc. Linn. Soc. N.S.W. 4(1): 178-186. , 1897. New genera and species of Austra- lian fishes. Proc. Linn. Soc. N.S.W. 22(1): 62-95. 82 NOEL R. KEMP Parrott, A. W., 1958. Big game fishes and sharks of New Zealand: 1-127. London. Patterson, C, 1967. Classes Selachii and Holo- cephali. In W. B. Harland el al. (Eds.) The Fossil Record. London (Geological Society) : 666-675. Peron, F., 1807. Voyage de Decouverts Aux Terres Australes. 1: 1-496. Paris. Peyer, B., 1968. Comparative Odontology. R. Zan- gerl, Trans, and Ed.: 1-347, 95 pis. Chicago and London. Phillips, W. J., 1924. A review of the elasmobranch fishes of New Zealand, N.Z. J. Sci. Tech. 6(5-6) : 257-269, 14 Figs. , 1935. Sharks of New Zealand; No. 4, N.Z. J. Sci. Tech. 16: 236-241, 3 Figs. Pledge, N. S., 1967. Fossil elasmobranch teeth of South Australia and their stratigraphic distribu- tion. Trans. R. Soc. S. Aust. 91: 135-160, 4 pis. RafinesQUE, C. S., 1810. Caratteri di alcuni nuovi generi e nuovi specie de anitnali (Presci) . * . defla Sicilia: 1-105, 20 pis. Palerme. Reed, M. D., 1946. A new species of fossil shark from New Jersey. Notul. Nat., Philadelphia, 172: 1-3, 1 pi. Sadowsky, V., 1970. First record of broad-snouted seven-gilled shark from Cananeia, coast of Bra- zil. Bohn Inst, oceanogr. S. Paulo 18(1): 33-35, 1 Fig. Scott, T. D., 1962. The marine and fresh water fishes of South Australia. Adelaide: 1-338. , C. J. M. Glover and R. V. Southcott, 1974. The marine and fresh water fishes of South Australia. Adelaide: 1-392. Springer, S. and R. A. Waller, 1969. Hexanchus vitulus, a new sixgill shark from the Bahamas. Bull. mar. Sci. 19(1): 159-174. Springer, V. G. and J. A. F. Garrick, 1964. A survey of vertebral numbers in sharks. Proc. U.S. natn. Mus., Washington, 116: 73-96. Tenore, M., 1810. Sopra una nuova specie di Squa- dro pescato nelle acque della riviera di Chiaja del littorale di Napoli. Mem. Acad. Pont. Naples, 1: 241-265. Waite, E. R., 1907. A basic list of the fishes of New Zealand. Rec. Cantab. Mus., Christchurch, 1: 3-39. , 1921. Catalogue of the fishes of South Australia. Rec. S. Aust. Mus., Adelaide, 2: 1-208, 332 Figs., 1 pi. , 1923. The Fishes of South Australia, Ade- laide: 1-243. Waldman, M., 1971. Hexanchid and orthacodontid shark teeth from the Lower Tertiary of Van- couver Island, British Columbia. Can. J. Earth Sci. 8(1): 166-170. Welton, B. J., 1974. Heptranchias howellii (Reed, 1946) (Selachii: Hexanchidae) in the Eocene of the United States and British Columbia. Paleo- bios, Berkeley, Calif., 17: 1-15, 2 Figs., 2 pis. Whitley, G. P., 1931a. New names for Australian fishes. Austr. Zool. 6: 310-334, 1 Fig., 3 pis. , 1931b. Studies in ichthyology, No. 5 Rec. Austr. Mus., Sydney, 18(4): 138-160, 2 Figs., 2 pis. , 1934. Notes on some Australian sharks. Mem. Qd. Mus., Brisbane, 10(4): 180-200, 4 Figs., 3 pis. 1940. The fishes of Australia. Part J. The sharks, rays, devil fish and other primitive fishes of Australia and New Zealand. Sydney (Royal Zoological Society): 1-280, 302 Figs. ( 1968. A check-list of the fishes recorded from the New Zealand region. Austr. Zool. 15(1): 1-102. Woodward, A. S., 1886. On the palaeontology of the Selachian genus Notidanus Cuvier. Geol. Mag., London (3) 3: 205-217, 253-259, 1 pi. Zeitz, A. A synopsis of the fish of South Australia. Part 1 Trans. R. Soc. S. Aust. 32: 288-293. Explanation of Plates All photographs by the author PLATE 12 Figures 1-4 — Hexanchus griseus (Bonnaterre). Teeth in situ of 4250 mm specimen from N.S.W. (AMS 119110-001). 1. Photomontage of labial view of re- placement series of right side of up- per and lower jaws including medial rows of left side of upper jaw and lower medial row. Functional series of upper jaw is seen inciso-lingually. 2. Incisal and labial views of functional series of medial rows of upper jaw. Note basal denticles on medial mar- gin especially of first right medial row (xl). 3. Labial view of replacement series of medial rows of upper jaw. Note large basal denticles on distal margins of first right (apex of crown broken) and second left medial teeth. First left row is almost obscured due to shrink- age of jaw in symphyseal region (xl). 4. Lower medial replacement row. Note first tooth with bifurcated central cusp while second tooth has no cen- tral cusp (xl). Figure 5 — Hexanchus vitulus (Springer and Waller). Labial view of teeth of 1550 mm male from Natal, South Africa; right side of upper and lower jaws including lower medial tooth. From Bass et al., 1975, with permission. SOUTH-EASTERN AUSTRALIAN HEXANCHID SHARKS 83 PLATE 13 Figure 1 — Heptranchias perlo (Bonnaterre). Labial view of teeth of 887 mm female from N.S.W.; right side of upper and lower jaws including lower medial tooth; pos- teriors, except for first upper, are not shown (TMD1247). Figure 2 — Notorynchus cepedianus (Peron). Labial view of teeth of right side of upper and lower jaws including upper central and lower medial teeth; posteriors, except for first upper, are not shown (NMV). 7. Lower lateral, labial face, lingual face (SAMP19552d). 8. Lower lateral, labial face, lingual face (SAM P19552e). 9. Lower lateral, labial face, lingual face. Blanche Point Marl (RJFJ no. 121a) (x2). 10-11. Lower lateral, Blanche Point Marl (RJFJ no. 121b). 10. Labial face, lingual face (x2). 11. Labial face; note basal den- ticle on first crownlet (xl2). PLATE 14 Figures 1-3 — Notorynchus cepedianus (Peron). 1 . Symphy seal area of upper jaw of specimen from California, U.S.A. showing medial and first lateral rows. Note two medial rows on either side of symphysis (TMD1303) (xl-5). 2 . Symphy seal area of upper j aw of specimen — topotype — from Storm Bay, S.E. Tasmania showing medial and first lateral rows. Note central medial row on the symphysis (TMD1291) (x2). 3. Abnormal, bifid tooth of first lateral row of lower jaws (NMV) (x2). Figures 4-11 — >Hexanchus agassizi (Cappetta). 4-8 From Naracoorte No. 5 Bore, 135- 145 m (x2). 4. Upper lateral, labial face, lingual face (SAMP19552a). 5. Upper lateral, labial face, lingual face (SAMP19552b). 6. Upper lateral, labial face, lingual face (SAMP19552c). PLATE 15 (all teeth x2) Figures 1-3 — Hexanchus agassizi Cappetta. 1. Lower lateral, labial face, lingual face, Naracoorte No. 5 Bore, 135-145 m (SAMD V34). 2. Lower lateral, labial face, lingual face, Blanche Point Marl (UAGD F17262). 3. Lower lateral, labial face, lingual face, London Clay, Isle of Sheppey, U.K.; for comparison (NRK). Figures 4-7 — Heptranchias howelli (Reed). Lower laterals, labial face, lingual face, Blanche Point Marl. 4. (SAMP19572). 5. (UAGD F172284a). 6. (UAGDF17284b). 7. (SAMP19573). Figures 8-10 — Notorynchus primigenius ( Agassiz) . Lower laterals, labial face, lingual face. 8. Muddy Creek Marl (TFF). 9. Batesford Limestone (NMVP27411). 10. Batesford Limestone (NMV P27410). MEM. NAT. MUS. VICT. 39 PLATE 12 MHM. NAT. MUS. VICT. 39 PLATE 13 OJ MEM. NAT. MUS. VICT. 39 PLATE 14 ? f T T # % 10 8 MIM. NAT. MWS. VICT. 39 PLAN IS ^ni t>7, -ffvy, » „ Wt * \ ■ 1* 'Mm t \ - '■ t '■■ mm 5 8 1 p i i ^jb pi* 10 RAEMEOTHERIUM YATKOLAI, gen. et sp. nov., A PRIMITIVE DIPROTODONTID FROM THE MEDIAL MIOCENE OF SOUTH AUSTRALIA By Thomas H. Rich, 1 Michael Archer, 2 and Richard H. Tedford* 1 National Museum of Victoria. 2 Queensland Museum, Brisbane 3 American Museum of Natural History, New York Abstract A new genus and species, Raemeotherium yatkolai, from the medial Miocene Namba Formation in South. Australia, is the most primitive member of the Diprotodontidae yet des- cribed. On its most anterior lower molar is a well-developed paracristid and protocristid. This, together with a strong cristid obliqua on all lower molars suggests derivation of diprolodontids from selenodont rather than quadritubercular ancestors. Introduction In September 1976, Mr Ian Stewart (Na- tional Museum of Victoria) discovered the di- protodontid dentary here described as Rae- meotherium yatkolai (SAM P19764). It was found at the south end of Lake Pinpa, South Australia, in the medial Miocene Namba For- mation. Only one or two centimetres away at the same level was a mandible of a species of pseudocheirine similar generically to that Ted- ford et al. (1976, p. 56) characterized as \ . , the most abundant mammal in the Pinpa Fauna . . .', (NMV P48616, see pi. 16, fig. 10). Three additional isolated teeth referred to Raemeotherium yatkolai were collected ten kilometres to the south on the west side of Lake Namba at a site in the Namba Formation designated as South Prospect B by Woodburne and Tedford (1975). A single isolated upper incisor of R. yatkolai was found at Ericmas Quarry, four kilometres to the north on the west side of the same lake, again in the Namba Formation. Callen and Tedford (1976, p. 135) and Ted- ford et al. (1977, pp. 56-57) discuss the basis for assigning a medial Miocene age to the Namba Formation. Abbreviations AM Australian Museum, Sydney. AMNH American Museum of Natural History, New York. BMNH British Museum (Natural History), Lon- don. NMV National Museum of Victoria, Melbourne. SAM South Australian Museum, Adelaide. Terminology and Measurements Mandibular terminology follows Stirton (1967), dental terminology is modified from Archer (1976) (see fig. 1), and enumeration of the teeth follows Archer (in press) (see fig. 1, 2). The system of tooth enumeration used here departs radically from all previously proposed. In it, the permanent cheek teeth of diproto- dontids are P3 ? M2, M3, M4, M5. In contrast, in the two systems previously employed by Twentieth Century authors, the same teeth are designated P3 or P4, Ml, M2, M3, M4. Measurements were made with a Wild microscope equipped with a mechanical stage, enabling an accuracy of 0- 1 millimetres. Systematics The following diagnoses are not intended to be exhaustive at each taxonomic level, but rather are restricted to those features which can be observed on the type and referred specimens of Raemeotherium yatkolai. Class MAMMALIA Linnaeus, 1758 Subclass THERIA Parker and Haswell, 1897 Infraclass METATHERIA Huxley, 1880 Superorder Marsupialia Illiger, 1811 Order Diprotodonta Owen, 1866 Family Diprotodontidae Gill, 1872 Diagnosis: Distinguished from all other Marsu- 85 86 THOMAS H. RICH, MICHAEL ARCHER, and RICHARD H. TEDFORD end — *C_^ hlch aepd pmcd med \ \ / ^rld ^paed hycK / pcd CO Figure 1 — Schematic diagram of diprotodontid lower molar showing terminology em- ployed. Abbreviations: aecd, anterior en- tocristid; co, cristid obliqua; end, ento- conid; hid, hypolophid; hyd, hypoconid; med, metaconid; pacd, paracristid; pcd, protocristid; pmcd, posterior metacristid; prd, protoconid; prld, protolophid; psd, protostylid. Modified from Archer (1976). Figure 2 — Schematic diagram of a diprotodontid mandible illustrating system of numbering teeth employed in this paper and method of measuring inclination of anterior bor- der of ascending ramus (angle a) and elevation of condyle (angle b). pialia by the presence of four simultaneously functional bilophodont molars lacking fore and midlinks and absence of a masseteric canal in the dentary. Subfamily Zygomaturinae Stirton, Woodburne, and Plane 1967. Diagnosis: Most members, including Raemeo- therium yatkolai, distinguished from all other diprotodontids by the presence of a distinct paracristid on M 2 . Raemeotherium gen. no v. Type species: Raemeotherium yatkolai sp. nov. Known distribution: Medial Miocene, Tarka- rooloo Basin, South Australia. Diagnosis: Distinguished from all other dipro- todontids by the presence of a distinct crest, the protocristid, directed buccally from the proto- conid on M 2 ; a well-developed anterior ento- cristid on M 2 ^; and the ascending ramus more recumbent than in other species. Etymology: RAEME, acronym for the Royal Australian Electrical and Mechanical En- gineers; Orjfjiov, Greek, beast. Raemeotherium yatkolai sp. nov. Plate 16, figures 1-5, 7, 8, 11) H oh type: SAM PI 9764, right dentary frag- ment with Ii, M 2 -5, and alveoli for P 3 . It lacks the P,i, condyle, and angular region and has been crushed in a mediolateral direction so that traces of many structures on the medial side such as the digastric fossa have been ob- literated. Type locality and stratigraphic position: Nam- ba Formation, southwest corner of Lake Pin- pa, grid zone 6, grid reference 317146 on the Curnamona topographic series SH 54-14, 1: 250,000 (1965, South Australia), 31° 8i'S., 140° 13'E. Diagnosis: That of the genus until other species are described. Etymology: Named in honour of the late Mr Daniel A. Yatkola, a student of mammalian palaeontology. Referred specimens: NMV P48537, isolated right M 4 or M 5 . NMV P48538, isolated right P. AMNH 102186, isolated left M 4 . Collected from the Namba Formation, South Prospect B, Lake Namba, grid zone 6, grid reference 3201 35, Curnamona topographic series SH 54-14, 1:250,000 (1965, South Australia), 31° 14'S., 140° 14'E. They were found in the white coloured unit described by Woodburne and Tedford (1975, p. 3) as a '. . . very fine sand . . .', developed as lenses within the \ . . thin-bedded black claystone . . .', at South Prospect B. MIOCENE DIPROTODONTID RAEMEOTHERIUM 87 AMNH 102183, isolated right P. Collected from the Namba Formation, Ericmas Quarry, Lake Namba, grid zone 6, grid reference 320140, Curnamona topographic series SH 54-14, 1:250,000 (1965, South Australia), 31° 12'S., 140° 14'E. It was found in a channel de- posit formed of white quartzose sand contain- ing green limonite-cemented clayballs. TABLE 1 Measurements (mm) of lower molars of Raemeotherium yatkolai Width Width Anterior Posterior Length Moiety Moiety SAM P19764 M» 10 7 5-9 6-7 M^ 110 65 7-4 M 4 111 71 7-1 M- 11-2 7-2 70 NMV P48537 M 4 or M 5 11 7-6 7-3 AMNH 102186 M 4 or M r> 11 7-7 7-8 TABLE 2 Measurements (mm) of the mandible of SAM PI 9764, type specimen of Raemeotherium yatkolai Length, diastema l!-P 3 : I5 a Depth, horizontal ramus at anterior end of P 3 : 19 Depth, horizontal ramus at middle of M„: 25 u Depth, horizontal ramus at middle of M 4 : 26 u Length, M 2 _ 5 : 44 2" a = approximately. Description Mandible: The mandible is remarkably delicate compared with other diprotodontids and in this respect resembles those of similar-sized macro- podids. The diastema between Ii and P :J is only about a third the length of the lower molar row. In outline, the symphysis is almost ellip- tical. Its major axis is about 25 mm in length and dips posteriorly at an angle of 35° with re- spect to the dorsal edge of the horizontal ra- mus. Its posterior edge is below the posterior root of Pa- The length of the minor axis is 13 mm. The surface of the symphysis is rough and there is no indication of its having been fused with the left mandible. The genial pit, for the insertion of the geniohyoideus muscle, dis- rupts the outline of the symphysis at its pos- terior end as a notch of smooth bone. The mental foramen is located 1*7 mm anterior of the anterior edge of P ;J , and 8 1 mm below the dorsal edge of the horizontal ramus. It is ellip- tical in outline with the major axis 4-2 mm in length and gently inclined anteriorly; the minor axis is 2 8 mm. Above and somewhat posterior to the mental foramen is a much smaller foramen. The ventral edge of the horizontal ramus below the molar row is gently convex downward. Posterior to that re- gion, the shape of the ventral border is un- certain owing to crushing. In the badly crushed posterior area of the mandible, part of the opening for the mandi- bular foramen is preserved 24 mm behind M 5 and about 3 mm below the level of the dorsal edge of the horizontal ramus. Distortion in this area is so great that the position in life of this foramen might have been several milli- metres away. Despite the shattered nature of the posterior part of the mandible, there is no doubt that a masseteric canal was not present. The anterior edge of the ascending ramus is posteriorly inclined. Angle a as defined in Figure 2 is 62°. Although the condyle is not preserved on this specimen, because the dorsal part of the posterior edge of the coronoid pro- cess is intact, angle b as defined in Figure 2 could not have been more than 34°. Lower dentition, /,; The thin layer of enamel on this tooth is confined to the buccal and ven- tral surfaces above the alveolar border. Along the dorsal border of the enamel, a well-de- veloped wear facet extends from the tip to about midway along the posterior edge of the enamel. An appression fossette is developed on the ventromedial surface of the enamel where this tooth contacted the left Ii. As on the molars behind, the enamel surface when unworn is wrinkled with gentle, irregular sinu- ous folds. The posterior end of the root is open and located beneath the middle of P 3 . P 3 : This tooth is not preserved but its two al- veoli are present in the type specimen. On the basis of the alveoli, this tooth, although much smaller, was similar to the P 3 of Kolopsis torus 88 THOMAS H. RICH, MICHAEL ARCHER. :mt] RICHARD H. TFDFORD in the ratio of its length to that of M,, and in the markedly greater posterior than anterior width- M s ; Unlike M 3 - r „ a distinct trigonid is de- veloped on the anterior moiety of the Mo. A well-developed paracristid extends from the protoconid anterior to the forward margin of the tooth. A paraconid is not present at the anterior end of the paracristid, but a cingulum begins at that point and extends posterolingu- ally along the margin of the tooth, terminating at the anterior end of the swelling at the base of the metaconid. Part of the mctaconid is missing but enough remains to confirm that the protolophid extends from the protoconid to the medial and slightly posteriorly positioned metaconid. A short ridge or cristid, the proto- cristid is directed posterobuccally from the tip of the protoconid. At the buccal end of the protocristid, there is a slight swelling to indi- cate the possible presence of a distinct cusp, the protostylid. The cristid obliqua is directed anterolingu- ally to abut against the posterior side of the trigonid at a point directly below and behind the protoconid. The hypoconid and entoconid are further apart from one another than the protoconid and metaconid. The hypolophid, connecting the hypoconid and entoconid, is convex posteriorly. A short anterior ento- cristid extends anterobuccally from the entoconid. Although present in other dipro- todontids such as Kolopsk torus, this an- terior entocristid is best developed in Racmco- therlum yatkolaL Along the posterior mar- gin of the tooth is a well-developed, slightly crenulated postcingulum. Near the centre of the cingulum is a small cuspule. An extremely weak vertical ridge or prominent crenulation extends up the posterior slope of the hypolo- phid from this cuspule, and merges with the hypolophid wear facette. A definite wear facet extends the length of this ridge. The small cus- pule is likewise truncated by a wear facet. None of the nearby crenulations display any signific- ant degree of wear. Of the four principal cusps, the protoconid is the tallest and the other three (metaconid, hypoconid, and entoconid) are subequal in height. In occlusal view, the tri- gonid and transverse valley are subequal in width and distinctly narrower than the talonid. M s - S ; The anterior moieties of these teeth are quite different from that of M.. In contrast, the posterior moieties of all molars are similar. A well-developed precingulum extends from the anterobuccal side of the protoconid base to the anterior side of the metaconid base. Un- like the M 2i the paracristid forms only a poorly- defined vertical crest from the tip of the pro- toconid to its base, and because of this the anterior moiety is not distinctly triangular in outline when viewed from above. Between the protoconid and metaconid is a well-developed protolophid that is convex posteriorly in occlu- sal view. Projecting a short distance posterolin- gually from the tip of the metaconid is a small posterior metacristid. This structure is some- what weaker on M rj than on M :{ -.». If it was present on M 2 , damage to that area of the tooth has removed all trace of the structure. Because of the similarity in the posterior moiety of all lower molars, only differences from the condition of M-> will be noted. The anterior entocristid becomes less-prominent posteriorly until on M a it has all but disap- peared. The postcingulum is wider and extends further towards the lingual and buccal margins of the tooth than on M L .. Cuspules are de- veloped posterior and slightly buccal to the entoconid on the postcingulum. Only on M 3 however, is there a homologue of the ridge con- necting the postcingulum with the crest of the hypolophid such as occurs on Ms, and even here it is not as well-developed as it is on M 2 . In order of decreasing height, the four principal cusps are the protoconid, metaconid, hypoconid, and entoconid. TABLE 3 Measurements (mm) of I 3 s of Raemeotherium yatkolai Length Width NMV P48538 4-6 4-6 AMNH 102183 5 4-4 Upper dentition, P: In occlusal view, this tooth has the outline of an isosceles right triangle, the corners of which are rounded, and the hypo- tenuse of which forms the posterolingual side. On the buccal side of the tooth is a broad, dor- MIOCENE DIPROTODONTID RAEMEOTHERIUM 89 soventral groove and on the anterior side, a well-developed appression fossette resulting from contact with I 2 . This tooth is quite similar in morphology to the homologous ones in Ngapakaldia tedjordi, Neohelos tirarensis, and Plaisodon centralis. Compared with measurements of Ngapakal- dia tedjordi (Stirton 1967), Kolopsis torus (Woodburne 1967), and Plaisiodon centralis (Woodburne 1967), the Fs described here are about the proper size to correspond with the molars of Raemeotherium yatkolaL Because no other diprotodontids of this size are known from the Namba Formation and two referred molars of R. yatkolai were found in the same site and level as one of the two incisors (NMV P48538 at South Prospect B), we are confident that reference here to R t yatkolai is justified. Discussion Raemeotherium yatkolai is unique within the Diprotodontidae and the most primitive member in that the paracristid and protocristid are well-developed on M 2 . The well-developed paracristid on M 2 is a feature of tribosphenic marsupials as well as some diprotodonts such as the selenodont possums. Archer (1976) sug- gested the possibility that lophodont marsupials (hence diprotodontids) may have been des- cended from selenodont or subselenodont pos- sums. The well-developed paracristid on M 2 , and the well-developed cristid obliqua on M 2 . 5 in Raemeotherium yatkolai at least suggest that diprotodontids did not descend from bunodont possums, but do not prohibit derivation from pre-selenodont tribosphenic ancestors, such as perameloids. On the Mo of selenodont diprotodonts buccal to the protoconid there is a variably-developed cusp that represents a protostylid; e.g. Phasco- larctos cinerus (NMV C2660, see pi. 16, fig. 9), and Pseudocheirus peregrinus (NMV C13937, see p. 16, fig. 6). On the M 2 of Rae- meotherium yatkolai, the protocristid appears to have terminated in a protostylid (see pi. 16, fig. 7). Apparently the protostylid appeared as a neomorph on the M 2 of selenodont diproto- donts and their descendants secondarily re- duced and then lost it independently in dif- ferent lineages. Evidence for the presence of this cusp as the primitive condition in groups descendant from the selenodont diprotodonts is to be found not only in R. yatkolai in the case of the diprotodontids, but in an undes- cribed Miocene macropod as well. Primitive diprotodontid features found in Raemeotherium yatkolai that are shared with some other members of the Diprotodontidae include the posteriorly inclined ascending ra- mus, the low position of the condyle on the dentary, all molars nearly the same size rather than markedly increasing in length and width posteriorly, and the well-developed anterior entocristid on M 2 - n . Inclination of the ascend- ing ramus is a quite variable feature within a given diprotodontid species. However, the re- cumbent angle {a in fig. 2) of R. yatkolai, 62°, is so much less than in any other diprotodontid that the differences can be confidently regarded as significant. Kolopsis torus with a minimum recumbent angle of 70° is the most similar diprotodontid, others fall in the range 80°- 100°. The ratio of M 2 to M 4 length typifies the difference between Raemeotherium yatkolai and other diprotodontids in molar lengths. In R. yatkolai, this ratio is 096, practically unity. The closest other diprotodontids are Ngapakal- dia tedjordi where the ratio ranges from 78 to 0-89 in three specimens measured by Stirton (1967), 0-84 to 86 in three specimens of Palorchestes painei measured by Woodburne (1967), and 85 in one specimen (AM F44471) of Kolopsis rotundus. All other known diprotodontids in which this feature can be measured, have a ratio of less than 85. In the same manner, the degree of primitiveness of R. yatkolai in each of the remaining features listed in the preceding paragraph cannot be exceeded in another diprotodontid. Some slight doubt remains as to whether Raemeotherium yatkolai is properly assigned to the Diprotodontidae because, as far as now known, the species has no derived or advanced character states which preclude its inclusion as a primitive member of the Macropodoidea. Although the presence of a masseteric canal, a derived condition within the Diprotodonta, is 90 THOMAS H. RICH, MICHAEL ARCHER, and RICHARD H. TEDFORD a feature unique to macropodoids, the oldest member of the superfamily may well have lacked one. On the basis of parsimony, the last common ancestor of the macropodoids and their immediate sister-group, whatever that may have been, presumably lacked a mas- seteric canal, this lack being the primitive con- dition. This means that at the time of their separation, unless the masseteric canal ap- peared as a part of that particular speciation event, there would have been an ancestral macropodoid which lacked that structure. R. yatkolai could conceivably have been a des- cendant of such a primitive macropodoid or near-macropodoid. Hence allocation to the Diprotodontidae must be regarded as tentative although highly probable, pending discovery of better preserved specimens of R. yatkolai. Assignment of Raemeotherium yatkolai to the Zygomaturinae within the Diprotodontidae must also be tentative because it is based on a primitive character state. By all the credible phylogenetic hypotheses that have been pro- posed for the Marsupialia, the presence of a strong paracristid on the Mo is primitive within the diprotodontids. Hence, it is to be expected in the common ancestor of the family and could therefore be the condition in primitive, as yet unknown members of the other subfamilies. Discovery of a single P 3 of R. yatkolai would probably settle the matter because all the un- doubted Zygomaturinae share a unique de- rived morphology of that tooth. Only one diprotodontid specimen is sig- nificantly older than Raemeotherium yatkolai. This is a form tentatively regarded as a palor- chestine by Tedford et al. (1975) who reported its presence in the late Oligocene or older Geilston Bay local fauna collected near Ho- bart, Tasmania. Unfortunately, this specimen (BMNH 40157) is not directly comparable with any known material of R. yatkolai, being a maxilla with part of M 2 and M ; ^ r '. However, the lengths of the respective tooth rows indicate that these two diprotodontids are about the same size and the smallest members of the family. They likewise show molars of similar length without conspicuous backward increase in size typical of later diprotodontids. Literature Cited ARCHER, M., 1976. Phascolarctid origins and the po- tential of the selenodont molar in the evolution of cliprotodont marsupials. Mem. Qd Mas. 17 (3): 367-371. (in press). The nature of the molar-pre- molar boundary in marsupials and a reinterpre- tation of the homology of marsupial cheekteeth. Mem. Qd Mus. 18. Cauhn, R. A. and R. H. Tedford, 1976. New late Cainozoic rock units and depositional environ- ments. Lake Frome area, South Australia. Trans. R.Soc.S.Aust. 100(3): 125-168. Stirton, R. A., 1967. The Diprotodontidae from the Ngapakaldi fauna, South Australia. Pp. 1-44 in Stirton, R. A., M. O. Woodburne, and M. D. Plane, Tertiary Diprotodontidae from Australia and New Guinea. Butt. Bur. Miner. Resour. Geo!. Geopliys, At/si. 87. Tedford, R. H., M. R. Banks, N. R. Kemp, I. Mc- Doucall, F. L. Sutherland, 1975. Recognition of the oldest known fossil marsupials from Aus- tralia. Nature 255 (5504): 141-142. Tedford, R. H., M. Archer, A. Bartholomai, M. Plane, N. S. Pledge. T. Rich, P. Rich and R. T. Wells, 1977. The discovery of Miocene verte- brates, Lake Frome area. South Australia. Bur. Miner. Resour. Jour. Aust. Geol. Geophvs. 2 : 53-57. Woodburne, M. O., 1967. Three new diprotodontids from the Tertiary of the Northern Territory. Pp. 53-103 in Stirton, R. A., M. O. Woodburne, and M. D. Plane, Tertiary Diprotodontidae from Australia and New Guinea. BulL Bur. Miner. Resour. Geol. Geophys. Aust. 87. Woodburne, M. O. and R. H. Tedford, 1975. The first Tertiary monotreme from Australia. Am. Mus. Novit. 2588: 1-11. Acknowledgements National Geographic Society grant no. 1562 to Rich made possible the 1976 expedition during which the type and two of the referred specimens were discovered. Essential ground- work for this expedition was laid in 1974 by Rich in collaboration with a contingent of the Third Royal Australian Electrical and Mech- anical Engineers Group of the Army under the command of Capt. Norman Moxey. Rich's interest in the general area east of Lake Frome stemmed from having been to the region as a member of Tedford's 1971 party supported by National Science Foundation Grant GB 18273X. Tedford was further supported by the MIOCENE DIPROTODONTID RAEMEOTHERWM 91 same agency and grant in 1973 when he re- visited the area and discovered the first referred specimens. Messrs. Roy Linke and Buddy Napier, managers of Frome Downes Station, and their families were most hospitable and provided facilities necessary for the success of our field programmes. Mr. Frank Coffa took the photographs and Mr. Alex Jelowyi did the lettering for the plate, Miss Jeanette Cook drew fig. 1 and Mr Harley Veitch, fig. 2. Explanation of Plate 16 Raemeotherium yatkolai. SAM PI 9764, type, right mandible, Lake Pinpa, South Australia, fig. 4, lateral view, xl; fig. 5, medial view, xl; fig. 7, occlusal view of M _ 5 , x2; fig. 11, occlusal view xl. NMV P48537, isolated right M, or M n , South Prospect B, Lake Namba, South Australia, fig. 1, occlusal view, x2. AMNH 102186, isolated left M 4 or M 5 , South Prospect B, Lake Namba, South Australia, fig. 2, occlusal view, x2. NMV P48538, isolated right I\ South Prospect B, Lake Namba, South Australia, fig. 3, occlusal view; fig. 8, lateral view, all x2. Pseudocheirine, NMV P48616, right mandible, found one or two centimetres from type specimen of Rae- meotherium yatkolai, Lake Pinpa, South Australia, fig. 10, occlusal view, x4. Pseudocheirus peregrinus, NMV C13937, left man- dible, juvenile, fig. 6, occlusal view of M-,__ ;! , x4. Phascolarctos cinereus, NMV C2660, left mandible, juvenile, fig. 9, occlusal view of P 8 -tW 8 , x2. Abbreviations: med, metaconid; prd, protoconid; psd, protostylid. MKM. NAT, MUS. VICT. 19 PLATE 16 2b ^ SYNONYMY OF THE FOSSIL WOMBAT VOMBATUS PLIOCENUS (McCOY) WITH THE LIVING SPECIES VOMBATUS HIRSUTUS (PERRY) By H. E. Wilkinson Abstract The type specimens of the fossil wombat species Vombatus pliocenus (McCoy) are re- deSCribed, and evidence presented for synonymy of this species with the living Common Wombat, Vombatus hirsutus (Perry). It is demonstrated that McCoy's description of this species was based principally on a specimen from Lake Bullenmerri of Ilolocene age. It is concluded that acceptance of the validity of this species has been strongly influenced by the presumed Pliocene age of the type specimen from Dunolly. The sediments from which this fossil came may be as old as Late Miocene, but it is suggested that the fossil could have been an intrusive burial. Introduction The discovery of a fossil wombat jaw at Dunolly in 1856 caused considerable interest, and some newspaper publicity, but it was not until 1861 that the fossil was recorded in the scientific literature by Professor Frederick Mc- Coy. He said (p. 168) 'I have likewise recog- nised the bones of the Wombat (Phascolomys) in the solid, hard, stony, ferruginous, auriferous drift called "cement" by the gold diggers, at a great depth in the sinkings at Dunolly, the material being so hard that the jaws could only be cleared by a stonemasons chisel; this deter- mination enables me to say that the age of the gold drift of Victoria, like that of Russia, is as Sir Roderick Murchison showed for the latter country, that of the "Mammiliferous crag" of England'. McCoy named the fossil Phascolomys plio- cenus in an essay for the Intercolonial Exhi- bition of 1866, the first printing of which ap- peared in a French translation in 1866, but it was not until 1874 that a full description and illustrations appeared in the first decade of his Prodromus of the Palaeontology of Vic- toria. The occurrence of this fossil wombat was quoted by authors such as Barkly (1869) and Smyth (1874, 1876) as the chief evidence for the Pliocene age of the gold drifts. It was referred to by Howitt (1913) and Walcott (1920) mentioned the occurrence in a paper on the age of the mammal bearing gold drifts of Australia. It was listed in the fossil catalogue prepared by Etheridge (1878) and illustrated in A ustralasian Fossils (Chapman, 1914). Tate (1951) accepted the species as valid when he reviewed the family Votnbatidae, and placed it in the genus Vombatus, a generic determina- tion accepted by Stephenson (1967) in a later review. Gill (1972) presented evidence for a probable Upper Pliocene age for the Dunolly fossil wombat, and briefly reviewed the cir- cumstances of its discovery, and the literature concerning it. The validity of the species has never been called into question, at least as far as the litera- ture is concerned, and the name has been ap- plied to several other fossil wombats of varying age from Victoria. The significance of the type specimens from Lake Bullenmerri has not previously been recognized. As part of a review of the family Vombatidae, the author has made a careful examination of the type specimens. Comparisons have been made with living and fossil wombats of comparable size to the living species. All the specimens referrred to are in the collection of the National Museum of Victoria, and are numbered with the prefix P. Provenance and age of the Type Specimens P7422, DUNOLLY, VICTORIA. The earliest known reference to the Dunolly fossil wombat is in The Courier of Mines and Bendigo Daily Mail of 30 September 1856, No. 253, p. 2, col. 6, in which a Mr Ferguson of the firm of Dennis and Ferguson at Dunolly re- ported the discovery of a jaw bone in cement at 30 feet. (Cement was the miners' term for silicified or ferruginized gold bearing conglo- 93 94 H. E. WILKINSON meratcs). In the same journal on 8 October 1856, No. 260, p. 4, col. 1, there is a report of Warden J. A. Panton receiving the fossil at Bendigo. via l Mr. Dixon of the Shamrock' (Ho- tel), and mentioning 50 ft. as the depth. Pan- ton thought it to be a kangaroo. Flett (1956, p. 33) records that the fossil was dug up from the hard cement of Gooseberry Hill, and dis- played for a time in the window of Mantel] and Wills, gold brokers at Dunolly, but does not cite the source of this information. Panton forwarded the specimen to Mel- bourne, but the details of the acquisition are not known. The Mining Commission Letter- book at the National Museum contains a copy of a letter from the clerk G. Ulrich to Panton (30 June 1857, p. 34) advising that the Palae- ontologist of the Geological Survey (F. McCoy) had determined the fossil as Phascolomys, and requesting Panton to provide details of its 'exact position, depth and circumstances 1 . This information may have been duly provided by Panton, but there is no evidence that it was. McCoy's earliest reference (1861) gives the locality as being l at a great depth in the sink- ings at Dunolly\ The depth is not stated in this, or any other of McCoy's references (1862. 1866, 1868, 1874), nor is a precise locality given. There is now no original label with the specimen, although Walcott (1920) refers to one. A letter written by A. R. C. Schvyn, Government Geologist, to the Editor of The Age, published on 28 February 1861, in- cludes a quotation from this label Thascolo- mys. From the "cement" of the gold drifts of Dunolly, 30 feet deep. Pliocene tertiary. Pre- sented by Mr Panton'. (This letter was written to refute a claim by R. Brough Smyth that kangaroo fossils found in Canard's claim at Yandoit in 1861 were the first which would enable the age of the auriferous deposits to be determined). Gill (1972) stated that the fossil came from a depth of 50 feet (15 metres) at the bottom of a shaft on Slaughteryard Hill, Dunolly. This locality was given by W. H. Ferguson in an un- published MS on Dunolly. Gill was aware that Flett (1956) had quoted Gooseberry Hill as the locality, but thought that this was an alter- native name for Slaughteryard Hill. The posi- tion is further complicated by the fact that the shaft he was shown is actually on Spiller's Hill, as is made clear by comparing the sketch plan provided for him by Mr John Flett of Dunolly, yrith the Geological Parish Plan of Dunolly. Mr James Flett has been unable to relocate the source of his Gooseberry Hill reference, and no other contemporary references to the site have been located, despite an intensive search (T. A. Darragh, pers. comm.). The question of exact locality is largely aca- demic, since all three hills are essentially simi- lar in their geological formation. They are part of a line of hill top cappings of Upper Tertian' gravel deposits, which extend from Golds- borough to Betley, paralleling the present day Burnt Creek. Gooseberry Hill is at the western end, followed by Spiller's Hill, and Graveyard Hill with Slaughteryard Hill at the eastern end. The gravels overlie tightly folded Lower Ordo- vician bedrock, into which the ancestral Burnt Creek was incised. Lithification of these old river gravels has left them more resistant to erosion than the surrounding bedrock, and they now stand in low relief above the valley of Burnt Creek. Gill (1964) presented evidence for an Upper Pliocene or Lower Pleistocene age for the Dunolly wombat. This was based on its alleged presence in post-Timboon Terrain stream sedi- ments, which had accumulated following the main movements of the Plio-Pleistocene Kos- ciusko Uplift. The gravels at Dunolly are equivalent to the White Hills Gravels at Bendigo (Wilkinson, 1 977) and like them consist essentially of quartz gravel, which contained rich concentra- tions of alluvial gold. (The fossil was found during the Burnt Creek rushes of the mid 1850s). Similar gravels at Axedale, about 20 km east of Bendigo, underlie a basalt flow re- cently dated at 6 95 ± 7 m.y. (Department of Minerals and Energy sample VAD 56, Am- del). This puts a minimum age of Late Miocene on these gravels. P.G. Macumber (pers. comm.) considers that the White Hills Gravels and their correlatives are equivalent to the Calivil Formation of the Deep Leads and Murray I OMBATUS PUOCENUS l )5 Basin. Palacontological evidence is consistent with a Late Miocene age for this formation also. The Tertiary age of the sediments is not in doubt, but there is the possibility that the fossil may be intrusive. Living wombats burrow ex- tensively, and burrow lengths of up to 20 m are known. Quite a few fossil wombats are known to be intrusive, and to represent in- dividuals trapped in burrow collapses. An ex- ample is a fossil wombat skeleton recovered from a burrow in marine fossiliferous marl at Modewarre near Geelong in 1884 (P1917), In this case the marl is Upper Miocene in age, but the fossil is unlikely to be much older than Late Pleistocene. Although McCoy identified it as pliocenus, it is in fact Vombatus hirsutus. The gravels at Dunolly are only partially lithiiied at depth, and would have represented no great obstacle to a burrowing wombat. If it were absolutely certain that the fossil came from the stated depth, this explanation might be argued against on the grounds that no liv- ing wombat has been known to burrow so deep. However, the circumstances surrounding the discovery of the fossil are almost totally un- known, and there is conflicting evidence as to locality and depth. The fossil certainly has an appearance of some antiquity, because of the encrusting matrix, but this can be matched on fossil wombats which are known to be geo- logically much younger than the gravels at Dunolly. For example, there is a partly matrix encrusted mandible from the ?Upper Pliocene- Pleistocene Shepparton Formation at Bung Bong near Maryborough, Victoria (P7420) which shows some strong similarities to the Dunolly specimen in the degree of mineraliza- tion and lithification of matrix. This specimen was collected in 1875, and was also determined by McCoy as pliocenus. It too is rcfcrrable to Vombatus hirsutus. P7441, P7442, lake bullenmerri, NEAR CAMPERDOWN, VICTORIA. The specimens from Lake Bullenmerri arc very lightly mineralized, and free of carbonate encrustation. They arc typical of fossils which have been collected from the shores of Lake Bullenmerri since the early days of settlement. The Lake Bullenmerri fauna consists of ex- tant species, although this does include Sarco- phifiis ursinus ( Tasmanian Devil) now extinct on the mainland. Gill (1953) pointed out that all the Western District post-tuff faunas he had examined contained extant species only, in- cluding Lakes Bullenmerri, Gnotuk and Keil- ambete. Lake Bullenmerri formed in the Holo- cene period, and the fossils arc not likely io be more than a few thousand years old, and may be much younger. The latter possibility is supported by observation of the rapidity with which bones of domestic animals on the shores of Lake Bullenmerri have become min- eralized in the 130 years or so of European occupation. The fossil bones are being washed up on the present beach level, presumably from erosion of an earlier terrace below present water level. No information is recorded of the circum- stances involved in the collection of the speci- mens. McCoy (1874) staled that they were 'From the shores of Lake Bullenmerri near Camperdown\ The drawing used in the Pvo- dromus was completed in 1862, and it appears probable that the specimens were collected by Selwyn when he visited Lake Bullenmerri (T. A. Darragh, pers. comm.). Selection of Lectotype and Lectoparatypes (iill (1953) listed the type specimens of plio- cenus as syntypes. Mahoncy and Ride (1975) list the Dunolly specimen as the holotypc, al- though McCoy did not designate a type speci- men in his 1866 description, nor was there any illustration. It is known that Plates 3-4 of his 1874 description were completed by Ludwig Becker in 1858, and Plate 5 was completed by A. Bartholomew in 1862. Il is clear there- fore that McCoy had all three specimens to hand when he published the first brief descrip- tion in 1866. The one character which he of- fered as a diagnosis of his new species cannot be measured on the Dunolly specimen, although a rough estimate can be made. 1 1 seems rather unlikely that McCoy based his new species on the Dunolly specimen alone, even though it is the only one referred to by locality. 96 II I WILKINSON It is therefore proposed to designate P7441 from Lake Bullcnmerri as the Lectotype. This is the only one of the three specimens used by McCoy on which his key diagnostic character is adequately preserved (see below), and the evidence that he used this specimen in 1866 is very strong. P7442 from Lake Bullenmerri, and P7422 from Dunolly are designated as Lecto- paratypes. The name pliocenus is therefore fixed on the best preserved specimen, which shows the one diagnostic feature given by McCoy in 1866, and all the additional features given in 1874. Systematic Description Order Marsupialia Family Vombatidae Iredale and Troughton 1934 Genus Vombatus Geoffrey 1 803 Type species Vombatus ursimis (Shaw) 1800 Vombatus pliocenus (McCoy) 1866 (Plate 1, figures 1,4. Plate 2, figures 1-2) Phascolomys pliocenus McCoy, 1866. Notes sur le zoologie ei la palaeontologie de Vic- toria: 21-22. Phascolomys pliocenus McCov, 1874. Prod. Pal. Vict. Dec. 1: 21-22, Pis. '3-5. Vombatus pliocenus (McCoy) Tate, 1951. Am. Mus. Novitates No. 1525: 5. 1. Lectotype: P7441. PI. 17, figs. 3-4. Incom- plete mandible with incisors, diastemal re- gion and left ramus with p4-m4 preserved. Locality: From shore of Lake Bullenmerri, Camperdown, Victoria. Collected prior to 1862. Age: Holocene. 2. Lectoparatype: P7442. PI. 17, figs. 1-2. In- complete mandible with incisors and right molar row preserved, minus m4. Locality: From shore of Lake Bullenmerri, Camperdown, Victoria. Collected prior to 1862. Age: Holocene. 3. Material: Lectoparatype: P7422. Plate 18, figs. 1-2. Almost complete mandible with both molar rows preserved, but the incisors obscured by matrix. Locality: From a depth of either 30 or 50 feet (9 1 or 15 2 metres) in an alluvial shaft on either Gooseberry or Slaughter- yard Hill, near Burnt Creek, Dunolly, Vic- toria. Collected in 1856, and presented to National Museum by J. A. Panton. (See discussion above). Age: The containing sediments are prob- ably Late Miocene in age, but the fossil may be intrusive and therefore younger. Descriptions and Comparisons The lectotype specimen from Lake Bullen- merri (P7441) is a ligtly mineralized incom- plete mandible, free of encrusting matrix. The right ramus is well preserved, with a complete molar row. The diastemal region has lost some of its bone, but both incisors are preserved. The premolar and first molar of the left ramus are also preserved, but the remainder has bro- ken away. The teeth are near perfect in preser- vation. The lectoparatype from Lake Bullenmerri (P7442) is an incomplete mandible, with most of the left ramus missing. It is less complete on the right side, and m4 is missing. It is better preserved in the diastemal region than P7441. The lectoparatype specimen from Dunolly (P7422) is an almost complete, partly matrix encrusted, heavily mineralized mandible. The left ramus has been freed of matrix, but the outer side of the right ramus, and the incisor region are still encrusted with a calcareous, ferruginous, silty matrix. Both molar rows are present, but all the occlusal surfaces are hol- lowed out. The left ascending ramus is pre- served, but the right is missing. The fossil was originally very nearly complete, but appears to have been damaged during the attempt to clear it of matrix. All three mandibles have in common fea- tures which clearly show that they belong to the genus Vombatus. These include the V- shaped inter-lobe valleys of the molars, the obliquely set, antero-posteriorly elongated pre- molars, and the position of the anterior root of the coronoid opposite the anterior lobe of m4. McCoy recognized that pliocenus was com- VOMBATUS PUOCENUS 97 parable to the living Thascolomys platyrhinus' (= Vombatus hirsutus) in size, position of sym- physis and 'the whole length of the dental series from edge of incisor to hind alveolus of last molar'. He claimed that his new species could be \ . , easily distinguished by the great size of the molar series, these teeth being not only larger transversely, occupying a longer fore and aft space, but extending so very much nearer to the edge of the incisors as to afford an easy mode of discriminating the species'. He laid great stress on the comparatively short diastema between the incisor and premolar, although this region is obscured by matrix in the Dun- oily specimen. He also differentiated pliocenus on the basis of 'the lower outline of the man- dible being rounded with a more uniform curve'; diastema being narrower between the molars; 'outlet of dental canal 1 (mental fora- men) being closer to the 'anterior molar' (pre- molar) and the incisors being more nearly equal in vertical and transverse diameters. Comparison by McCoy with other fossil wombats was confined to Vombatus mitchellii (Owen) from which McCoy stated that plioce- nus differed in having much larger molars, and in the symphysis extending further back. It has been shown elsewhere (Wilkinson, 1 973; in prep.) that Owen's concept of mitchellii in- cluded specimens of Lasiorhinus krefftii. The differences in position of the symphysis and size of molars referred to by McCoy are in fact features of generic distinction between Vom- batus and Lasiorhinus. Examination of the characters regarded by McCoy as diagnostic of pliocenus, with comments on their validity (a) "Great size of the molar series 9 The length of the alveoli of p4 to m4 in the type specimens is 55-9 mm in P7422, 562 mm in P7441 and 56-1 mm in P7442. The author has measured a specimen of V . hirsutus with an alveolar length from p4 to m4 of 57 -2 mm, but even this is exceeded by the largest specimens in the sample examined by Merrilees (1967, Figs. 1 and 2), which reach about 58 mm. The pliocenus types all fall well within the range of variation in V. hirsutus in this feature. (b) Ratio of length of molar row to 'whole length of dental series' McCoy stated that the ratio of the length of p4 to m4 to the 'whole length of the dental series from hind edge of last molar alveolus to front edge of incisor 'was 55:100 in pliocenus but only 45; 100 in platyrhinus (= hirsutus). In effect he was saying that the living species had a relatively shorter molar row and longer dia- stema than the fossil. This is the feature on which he laid greatest stress, and claimed that it gave \ . . an easy mode of discriminating the species'. This was tested by measuring the same characters on specimens of Vombatus hir- sutus and expressing the result as a percentage, for comparison with the percentages obtained from the types. In P7422, the incisors are dam- aged, and matrix obscured, and the ratio can only be estimated. The length from the tip of il to posterior alveolus of m4 is estimated to have been about 100 mm, and the percentage of the order of 54 to 56%. In P7441 the cor- responding figures are 101-9 mm and 55 5% and in P7442 they are 103 mm and 54-5%. The range in a sample of specimens of V. hir- sutus examined by the author was 95-8 to 112-7 mm and the percentages range from 51-7% to 54-7%. Although the percentage figure for P7441 is slightly higher, overlap be- tween pliocenus and hirsutus in this feature is demonstrated, and McCoy's claim of an easy distinction is therefore invalid. No hirsutus specimen with a percentage as low as 45% has been seen by the author. (c) Diastema narrower between the molars McCoy measured the distance between the posterior lobes of the 'second molar' or ml, to support his contention that 'the diastema is narrower between the molars'. In P7422 this is 24-5 mm; in P7441 it is estimated to have been 20-5 mm, and in P7442 it is 19-4 mm. In modern V. hirsutus the range is from <18 mm to >28 mm. Once again, the pliocenus types fall within the range of variation of the living species. 98 II. E. WILKINSON (cl) Shorter diastema McCoy claimed that the diastema was 'much shorter in the fossif. This was tested by mea- suring the length of the diastema between p4 and II. In P7422 this is estimated to have been about 27 mm, in P744I it is 30 mm and in P7442 it is 29 mm. In a small sample of modern V. hirsutus, the diastema varied from 26-3 mm, to 32 6 mm. The pliocenus types therefore fall completely within this range. (e) 'Lower outline of the mandible rounded with a more uniform Curve* This is a subjective judgement, difficult to quantify. However it follows that specimens with a relatively short diastema will tend to look more uniformly rounded, whereas long diastema mandibles tend to have the lower outline more elongated and angular. It is a variable feature of very dubious value for specilic distinction. (f) Anterior outlet of 'dental canal (— men- tal foramen) closer to 'anterior molar (~ premolar). This feature is not visible on P7422, but in P7441 the mental foramen is 5 3 mm from the premolar, and in P7442 the distance is 4 mm. This is a very variable feature, which varies from as little as 4 mm to in excess of 8 mm in living V. hirsutus. The pliocenus types P7441, P7442 are within the range of varia- tion in this feature also. (g) Greater compression of incisors Examination of a large range of specimens makes it clear that incisor cross section is rather variable in V. hirsutus. The difference between P744i and P7442 in this regard is typical of the degree of variation. Incisor cross section cannot be determined on the Dunolly specimen. P7422. Height expressed as a percentage of width is 74*4% in P7441, and 872% in P7442. In modern V. hirsutus] the range observed is from 70% to almost 100%. The figures calculated for the types of pliocenus fall within this range. The type specimens of pliocenus fall within the range of variation of Vomhatus hirsutus in all the features regarded bv McCov as diagnos- tic of his new species. He was probably in- fluenced by his belief that all three specimens belonged to the extinct fauna, and the number of comparative specimens of the living species available to him was no doubt very limited. He could hardly have been aware of the degree of variability in Vomhatus hirsutus. Merrilees (1967) stated that 'Variability is a striking characteristic of modern wombats', and gave details of the abnormalities encountered in 35 out of 52 individuals in the sample examined by him from Dingo Dell — Tairbank\ A.C.T. This was a breeding population, in which variation would be expected to be lower than in the species as a whole. Other fossil species of Vomhatus are in doubt for the same reason. Merrilees (1967) has already shown that Vomhatus parvus (Owen) is a juvenile V. hirsutus, and it is al- most certain that the same is true of V. thomp- soni (Owen). Owen's fossil species V. mit- chellii is based partly on specimens of Lasior- hinus krefftii (Wilkinson, 1973). The status of these species is currently under review (Wilkin- son, in prep.), and preliminary studies suggest that Vomhatus mitchellii may also be synony- mous with hirsutus, although possibly sub- speciiically distinct. The only other fossil spe- cies of Vomhatus is V. hacketti described by Glauert (1910) from Western Australia. It is readily distinguished from the pliocenus types and V. hirsutus by its narrower molars with more rounded lobes. Given the degree of variability in modern V. hirsutus and the doubtful status of most Of the fossil species of Vomhatus, the conclu- sion seems inescapable that Vomhatus plioce- nus (McCoy) is a junior synonym of Vomhatus hirsutus (Perry). Conclusion All the diagnostic features given by McCoy in defining pliocenus have been carefully ex- amined, and found to be invalid. No other character is evident on any of the types which would warrant their specific separation from hirsutus. The lectotype and lectoparatype from Lake Bullenmerri are of Holocene age, and there can be no doubt that they represent the living species. The lectoparatype specimen VOMBATUS PLIOCENUS 99 from Dunolly cannot be differentiated from hirsufm either, although acceptance of Vom- batus pliocenus McCoy as a valid species ap- pears to have been strongly influenced by its presumed Pliocene age. It has been shown above that this is an assumption which is not supportable by direct evidence, and the possi- bility of the fossil being intrusive, and there- fore younger, must be considered. If it is con- temporary with the sediments, then this has im- portant implications for studies of wombat spe- ciation. It could be argued for example that Vombatus evolved in the late Miocene, and that the mandible has remained virtually un- changed since then, although other elements of the skeleton might show significant differences. Until better material of known age and proven- ance becomes available, this cannot be re- solved, and it is better to regard the Dunolly specimen in the same light as the Lake Bullen- merri specimens. That is, Vombatus pliocenus (McCoy) is a junior synonym of the rather variable modern species Vombatus hirsutus (Perry). Acknowledgements The author wishes to thank the following people for their help in preparing this paper. Mr T. A. Darragh, Deputy Director, National Museum of Victoria, has been particularly helpful in tracking down early references to the Dunolly wombat, including unpublished material in the Museum Archives. Dr Duncan Merrilees, Western Australian Museum, and the author made a joint study of problems in- volving Vombatus pliocenus, Vombatus mit- chellii, and Lasiorhinus krefftii in 1968. Grate- ful acknowledgement is made to his contribu- tion to this paper in the form of valued discus- sions and follow up correspondence, although the author accepts full responsibility for the views expressed. Benefit has also been gained from discussions with E. D. Gill, and P. G. Macumber on the age of the sediments. Thanks are due to the Director of the Na- tional Museum, Mr J. McNally, and the Mu- seum staff for assistance with specimens (T. A. Darragh, T. Rich, I. Stewart, J. M. Dixon, A. J. Coventry). The photographs were taken by Frank Coffa of the National Museum. Thanks are also due to Dr D. Spencer-Jones, Director, Geological Survey Division, who al- lowed time towards completion of this pro- ject, commenced while I was on the Museum staff, and granted permission for its publica- tion. References Barkly, H., 1 865. Anniversary Address. Trans. Row Soc. 17(7.6: XIX-XXIV. Chapman, F., 1914. 'Australasian Fossils', Melbourne (p. 290. Fig. 142). Ethebjcge, R.. 1 878. A catalogue Of Australian Fos- sils, Cambridge (p. 190). FLETT, .(., 1956. Dunollw Melbourne, p. 33. Gill, F. D., 1953a. Geological Evidence in Western Victoria relative to the Antiquity of the Austra- lian Aborigines. Mem. Nam. Mas. Vict. 18: 25-92. , 1953b. Catalogue of Quaternary Types and Figured Specimens in the National Museum, Melbourne. Ibid. 18: 157-168. -, 1964. Rocks contiguous with the basaltic cuirass of Western Victoria. Proc. Row Soc. Vict. 77(2): 331-355. . 1972. The Dunolly fossil wombat. Vic- torian Nat. 89(3): 64-66. Glauert, L., 1910. The Mammoth Cave. The wom- bat Phascolomys hackctti sp. no v. Rec. W. Aust. Mas. 1(1): 15-29. Howrrr, A. M.. 1913. The Maryborough Goldficld. Mem. Civol. Surv. Vict. 11:7. N&VKONEY, J. A. and W. D. L. Ride, 1975. index to the genera and species of fossil mammalia de- scribed from Australia and New Guinea between 1838 and 196S. W. Aust. Mus. Spec. Publn. No 6. McCoy, F.. 1861. Note on the ancient and recent natural history of Victoria, (in) Catalogue of the Victorian Exhibition, Melbourne, p. 168. , 1862. Note on the ancient and recent na- tural history of Victoria. Ann. Mag. Nat. Hist. 9 (3rd Ser.): 137-150. -, 1866. Notes sur la zoologic ei la palaeon- tologie tic Victoria, Melbourne, 35 pp. , 1867. On the recent zoology and palaeon- tology of Victoria (Intercolonial Exhibition Fs- say No. 7). Pp 307-330. Melbourne. 1868. On the species of wombats. Trans. Proc. Roy. Soc. Vict. 8(2): 266-270. ■ — ~ . 1 874. Phascolomys pliocenus ( McCoy ) . Proilromus of the Palaeontology of Victoria De- cade 1: 21-22, pis. 3-5. Mehrilees, D., 1967. Cranial and mandibular charac- ters of modern mainland wombats (Marsupialia Vombalidae) from a palaeontological Viewpoint, and their bearing on the fossils called Phascoia- 100 H. H. WILKINSON mys parvus by Owen (1872) Rec. S. Aust. Mus. 15(3): 399-418. Owen, R. (in) T. L. Mitchell, 1838. Three expedi- tions into the interior of eastern Australia, with descriptions of the recently explored region of Australia Felix, and of the present colony of New South Wales. London, Vol. 2: 362-363, pi. 30, figs. 4-7. OWEN, R. 1872. On the fossil mammals of Austra- lia — Hart VI. Genus Phascolomys, GeofTr. Phil. Trans R. Soc. 162: 173-196, pis. 17-23. Smyth, R. B., 1874. Geo!. Surv. Vict. Rept. Prog. 1: 36. Smyth, R. B., 1876. Ibid. 3: 74. Stephenson, N. G., 1967. Phylogenetic trends and speciation among wombats. Aust. J. Zool. 15: 873-880., Tate, G. H. H., 1951. The Wombats (Marsupialia, Phascolomyidae) Am. Mus. Novitates No. 1525, 18 pp. Walcott, R. H., 1920. Evidence of the age of some Australian gold drifts, with special refer- ence to those carrying mammalian remains. Rec. Geol. Surv. N.SW. 9(2): 66-97 (p. 69). Wilkinson, H. E. (in) Merrilees, D., 1973. Fossili- ferous deposits at Lake Tandou, New South Wales, Australia. Mem. Natn. Mus. Vict. 34: 177-182 (see pp. 180-181). Wilkinson, H. E., 1977. Geology of the Bendigo Area. Geol. Surv. Rept. Willi. (in prep.) Re-examination of the fossil wombat species Vomhatus mitchellii (Owen) and Lasiorhinus krefftii (Owen), and clarification of their taxonomy. Explanation of Plates PLATE 17 Vomhatus pliocenus (McCoy) Figure 1 — Labial view of lectoparatype, left man- dible P7442 from Lake Bullenmerri, Vic- toria; Holocene. Figure 2 — Occlusal view of same specimen. Figure 3 — Labial view of lectotype, left mandible P7441 from Lake Bullenmerri, Victoria; Holocene. Figure 4 — Occlusal view of same specimen. PLATE 18 Vomhatus pliocenus (McCoy) Figure 1 — Labial view of lectoparatype mandible P7422 from Dunolly, Victoria; ? Late Miocene. Figure 2 — Occlusal view of same specimen. (All specimens | natural size) MH.M. NAT. MUS. VICT. 39 PLATE 17 ■>. :-( fl ' 1 M ^■■"" J. * qB « V'-"* * t f i . *• ■•*■ £ W ■. ; ***** ■ ' y-jmi'- MI M. NAT. iVIUS. VICT. 39 PLATE IX 1 l_ COMMENTS ON THE SPECIES CONCEPT IN SOME AUSTRALIAN ANISOPS SPINOLA (HEMIPTERA: NOTONECTIDAE) By I. Lansbury Hope Department of Entomology, University Museum, Oxford Introduction Twenty-five species of Anisops are recorded from Australia and Tasmania, 16 are endemic. A few species are taxonomically confusing as they occur in more than one form. Anisops is compared with Bttenoa Kirkaldy which replaces Anisops in the New World. The species concept is discussed, especially of those with an extra-Australian distribution which tend to be superficially distinct at the periphery of their range. Figures and short descriptions are given of little known species from New Guinea which may eventually be found in Australia. Finally a new species is described from Lihir, Bis- marck Archipelago. Species Concept Alpha taxonomy To understand the problems posed by vari- ous species of Anisops, it is necessary to con- sider the species concept in the genus. With few exceptions, only the males can be named with reasonable certainty. Attempts by Sweeney (1965) and Lansbury (1969) to devise reasonably easy keys to the Australian Anisops have not been completely successful. The occasional failure of my 1969 key is caused by trying to key out forms of species which were not available to me when the key was prepared. The key couplet re- ferring to the comparative width of the head to the pronotum has proved unreliable when dealing with species which vary in the degree of development of the flight musculature. There are over one hundred described and seemingly valid species of Anisops, some of them with a widespread distribution. The characters used to divide up the genus into manageable proportions vary greatly in their reliability. Structural peculiarities of the head and rarely the pronotum easily distinguish a small number from most of the genus. Aus- tralian species with reliable k spof characters are stall Kirkaldy; calcaratus Hale; nasuta Fie- ber; semita Brooks; and tahitiensis Lundblad. Of the remainder, thienemanni Lundblad; no- ditlata and canaliculate Brooks are easily recognized by abnormally shaped rostral prongs. The characters used by Brooks (1951) to separate species fall into two categories (1) morphometries of the head capsule and the pronotum; (2) presence of grooves, ridges or prolongations of the head capsule especially the facial tubercle. Secondary sexual charac- ters with combinations of the primary features are reworked to enable species to be keyed out. The measurements of the head and prono- tum are not easy to make as the structures are nearly all convex. The chaetotaxy of the male front leg, number of pegs in the stridulatory row and distribution of spines and setae on the front tibia and tarsus are used. The ros- tral prongs which are projections borne either side of the third rostral segment are also im- portant. Wing polymorphism The terminology used to indicate if an Ani- sops can fly or not is rather confusing. No problem arises over the term macropterous, but brachypterous can be misleading. Ail Aus- tralian Anisops have fully developed fore wings and hind or metathoracie wings. Despite the presence of wings, most species produce forms with the wing musculature undeveloped, in its place parenchymatous tissue. Young (1962) uses the term 'normal' for individuals capable of llight and 'flightless' for those which although possessing full sized wings, have not developed (light muscle. The term 4 brachyelytrous' has been sug- gested, although possibly quite suitable, some confusion may arise as this term is also applied to the Staphylinidae and Dermaptcra with their characteristic short elytra, a condition not oc- 101 io: r. LANSBURY earring in Anisops. To avoid confusion, the terminology of Young (1962) is followed in these notes. The normal forms with fully developed flight musculature are easily recognized by the pig- mented scutellum, the clavus and corium also coloured, most commonly black, brown, red or yellow, usually a subtle combination of two or three colours. The metanotum is black or dark brown. Flightless forms (always with fully developed metathoracic wings in Austra- lian species) have hyaline scutellums, the dor- sal pigmentation showing through, likewise the clavus and corium. The metanotum is not pig- mented, but occasionally dark brown at the wing bases. The following descriptions are taken from Young (1962) and refer to assimi- lis and wakefieldi White, both endemic to New Zealand. The macropterous form of both sexes have the lateral margins of the pronotum di- vergent. The males of the brachypterous forms have more-or-less parallel sided pronotums and females with slightly divergent pronotums. A check of Brooks (1951) and Lansbury (1964) shows that they often described two colour forms of a species but did not comment on the shape of the pronotum. A further complication has been found in a number of species which are isolated from the rest because the head is as wide or wider than the pronotal humeral width, these are: evansi Brooks (Tasmania) robusta Hutchinson (E. Africa) lencothea Esa- ki (Samoa and elsewhere) doris Kirkaldy (Australia) tasmaniaensis Brooks (Tasmania) philippinensis Brooks (Philippines and else- where) and assimilis. Young's (1962) descrip- tion of assimilis differs from Brooks, the head irrespective of the form is always narrower than the pronotal humeral width, it seems to be a general rule in Anisops that the lateral margins of the pronotum of flightless males are more-or-less parallel sided, therefore the taxo- nomic value of the head-pronotal width is of limited use in those species which have more voluminous eyes. Series of evansi from Tas- mania and the mainland show both forms to be present, where females can safely be associated with males, the head is always narrower than the humeral width. In B. lirtinocastris Hungerford and other species, both sexes of the flightless forms have the head as wide or wider than the pronotal humeral width, normal forms of both sexes have the head narrower than the humeral width. Truxal (1953) describing the wing poly- morphic forms of limnocastris found that the scutellum of the normal form was longer than the median length of the pronotum, that of the flightless form clearly shorter. These differen- ces have not been found so far in Anisops. Colour It is not known how much the colour of Ani- sops species can vary. Young (1962) found a direct correlation between colour and flight musculature. There is some evidence that in some circumstances Anisops exhibit a pheno- menon rather better known in the Corixidae (see deanei). Specimens from turbid water with a peaty or dark substrate frequently have a much larger area of the hemelytra with dark brown transverse markings, the pronotum is often more heavily pigmented. The same spe- cies of corixid from a sandy or gravel substrate are much paler. The significance of colour in Anisops where it impinges on food gathering and its role in enabling Anisops to merge with the background are poorly documented. The differences noted by Young (1962) refer al- most exclusively to the colour of the scutellum. The pale flightless Anisops with little dorsal pigmentation may be difficult for a predator to locate visually, the normal form with a pig- mented scutellum and intensified colouring of the hemelytra may be easier for a predator to see. Both forms would be equally at risk viewed from any position sub-surface when ascending to the surface to renew their gas supply or at risk from predators which use other senses than sight to locate their prey. Bayly, Ebsworth and Wan (1975) present data from which it can be inferred that within closed systems such as exist on Fraser Island, Qld., that fish and water-bugs are mutually exclusive. They pos- tulate that either the fish eliminate the bugs by feeding on them, or the fish out-compete the bugs for the available food supply. Young's (1962) data on assimilis shows that both forms were found in almost equal numbers. The num- AUSTRALIAN AN ISO PS 103 ber of habitats where only the normal form were found was almost the same as the flight- less form. The total number of habitats with both forms present being just over double the number of those where only one form was present. Anisops wakefieldi differed, the nor- mal form was ten times more common than the flightless and four times as many habitats were populated by the normal form compared with the flightless form. Both species were found together in 'swarms', assimilis having a greater tolerance of habitat types. In the more stable weedy habitats with little open water wakefieldi was the dominant species. Genitalia The genitalia of Anisops are of very limited value at the species level. Brooks (1951) figured the left and right parameres but made no re- ference to them. Young (1962) did not figure or comment on the New Zealand species. Trux- al (1953) figured the parameres of seven Buenoa species which closely resemble Ani- sops in general plan. The female genitalia of Anisops are of less taxonomic value than the male, the same seems to apply to Buenoa al- though hungerjordi Truxal differs from the others in the shape and distribution of large setae on the first gonapophysis, hungerfordi oviposits in rock crevices rather than plant tissue as do most other species of Buenoa and, as far as is known, all Anisops. The figures of the male genitalia (Figs. 43-58) are slightly misleading as they tend to show differences which do not exist, this is because some diffi- culty was experienced in placing the parameres on identical planes on the slides leading to some distortion. The apex of the left paramere is rather variable, the tip sometimes being curved inwardly, thus when viewed from the side, the tip is not visible. The right paramere is a simple flat plate. Isolating mechanisms As it seems that the genitalia and the dis- tinctions contrived by the taxonomists using the morphometries of the head and pronotum are of limited value, the problem arises of what in certain circumstances constitutes a species in Anisops. Since Buenoa are so very much H alike Anisops, one of the main differences be- ing that the male Buenoa has two segmented front tarsi, whereas Anisops males have one- segmented tarsus. It is felt that the behaviour of Buenoa and Anisops are probably rather simi- lar. In laboratory experiments it has been found that B. limnocastris males will pair with macrotibialis Hungerford females, the Fj gen- eration are sterile, males morphologically in- termediate, their acoustical behaviour similar to that of their male parents. Stimson Wilcox (n.d.) (1975). Truxal (1953) descriptions of the stridulatory apparatus of Buenoa species, es- pecially the number of stridulatory pegs shows that within a species, the variation found is less than in Anisops. Young (1962) found that in assimilis males, the shape and number of stridulatory pegs was fairly constant; the size and arrangement in wakefieldi could be vari- able (28 and 15 pegs respectively). The short peg row of wakefieldi, rostral prong and ex- panded facial tubercle would produce a dif- ferent sound than the longer peg row of assimi- lis drawn across a larger rostral prong. The expanded facial tubercle of wakefieldi would greatly hinder their attempts to pair with assi- milis females. Presumably in New Zealand where Anisops has been geographically isolated for a long period and the number of species small, stable populations would develop. Sound production in Buenoa is assisted by the body which in whole or part acts as a frequency gen- erator. Experimental work with Buenoa shows that acoustical behaviour is important in repro- ductive isolation amongst coexistent species, Stimson Wilcox (n.d.), (1975). No information is available on Anisops. Acoustical behaviour as an isolating mechanism in mixed populations is well known in other groups. Ragge (1965) describing British Acrididae (Orthoptera) likely to be found in the same general habitat states that Chorthippus brunneus (Thunberg) has 50-90 stridulatory pegs on each hind femora; C. parallelus (Zetterstedt) 70-130 and Myr- meleotettix maculatus (Thunberg) 1 30-1 80 pegs etc. Ragge's diagrams of the songs of these and other species shows that irrespective of the number of pegs, the songs of the males of each species remain consistent in volume 104 I. LANSBURY and duration. It is reasonable to assume from this that Anisops males only utilize sufficient pegs to produce a consistent species signal. This factor also tends to reduce the taxonomic value of minor variations in the number, size and arrangement of the stridulatory comb. This variation seems to be a feature of a num- ber of nondescript species with an extensive geographical distribution. Assuming that part of any population of an Anisops species living in ponds, dams and ephemeral pools contains a percentage of in- dividuals with fully developed musculature, it is essential that should conditions deteriorate sufficiently to induce flight, the migrant popu- lation should produce and recognize acoustic signals from the same species in the habitat emigrated to, otherwise discrete populations would form in the same habitat. Exceptional habitats such as Lake Tengano, Rennell Is- land only seem to produce flightless forms see 'capitata'. Anisops elstoni, deanei and philippinensis Brooks Under elstoni material from various locali- ties in different countries is compared. With one or two exceptions nothing is known about the type of habitat, population density, species diversity or basic chemical and physical pro- perties of the habitats where elstoni was col- lected. The genetic aspects remain totally un- known. As more material becomes available it has become increasingly difficult to confine the species to its original concept which, lacking secondary sexual characters, reproductive or- gans indistinguishable from other species and where the morphometries are complicated by structural changes caused by wing polymor- phism. The discontinuous distribution of el- stoni and other species suggest that the species concept is not fully understood or the term 'super species' might be appropriate. In Tasmania which has five species of Ani- sops, unpublished data shows that where two or more species are found in the same habitat, one at least is easily recognized by its larger size and/or secondary sexual characters leav- ing the remaining species to be easily recog- nized by their lack of secondary features. Con- ditions in Tasmania tend to be rather different from those on the mainland with its greater di- versity of species. The carrying capacity of habitats, especially species diversity rather than the total population of water-bugs etc. is not too well known. In Tasmania, out of 175 samples, only one habitat had eight species of water-bug present (Blackmans Lagoon). Three habitats with six species and six with five. The great majority of samples composed of two-three species. These figures may reflect vagaries of sampling rather than an accurate reflection of these habitats. There are 24 species of water-bugs in Tas- mania, 28 if the Saldidae, Ochteridae and Ge- lastocoridae are included. Fraser Is., Qld. has been studied in some de- tail (Bayly et al.> 1975). Four species is the maximum recorded from Boomerang South Lake, 13 lakes sampled. Bensink and Bur- ton (1975) on the littoral fauna of Blue and Brown Lakes on Stradbroke Is., Qld. found five and nine species respectively. Brown Lake being sampled more often than Blue Lake. Some of the Anisops listed in Bayly et al. (1975) and Bensink and Burton (1975) are discussed in the taxonomic section of these notes. The problem arises of deciding whether the material from Tasmania is conspecific with those from mainland localities in Australia and further north in New Guinea etc. The data can be interpreted in one of two ways, elstoni may be a complex of races which because of geographical isolation are in the process of evolving into distinct species or it may be a widespread 'plastic' species which has spread from south east Asia into Austra- lasia, each population still linked by common genetic factors. Anisops deanei is confined to Australia and a closely allied form or species is rather com- mon in Tasmania, much more so than elstoni. Non-Tasmanian deanei varies rather more than elstoni. In Tasmania they are both some- times found in the same habitat, but do not seem to be so on the mainland. Where two or more species occur in the same habitat, acoustic behaviour, phased life AUSTRALIAN AN1S0PS 105 cycles and a subtle niche preference not so far revealed by sampling seem to enable Ani- sops species to coexist and avoid competition for the available food supply. Habitat and distribution Lack of data makes it difficult to be pre- cise about the habitat preferences of Austra- lian Anisops species. Sweeney (1965) describes elstoni as rare and confined to the south east, the most westerly record being Narrandera, N.S.W. Most Australian Anisops are found along the eastern coastal zone, a few are con- fined to the more arid interior. Some species typified by thienemanni, stali and Hyperion Kirkaldy are found across the continent. The coastal zone from Brisbane northwards has a number of species with prominent secondary sexual characters and not related to the New Guinea fauna. Species of the more temperate areas are less well endowed with such features. Anisops philippinensis appears to be a relative newcomer to the Queensland fauna rather like tahitiensis Sweeney (1965). Taxonomy Anisops elstoni Brooks, 1951 Figures 1-15, 43-50 and 59-61 This species was described from Myponga, South Australia; The Dorrigo, 3,00(y, New South Wales; Brisbane, Queensland and Szech- uan, Suifu, China. Brooks compared elstoni with exigera Horvath (New Guinea). His figures of the male front leg are misleading as he omits to include the large spine on the inner proximal margin of the front tarsus (Figs. 1, 4, 7, 10 and 13) and he overlooked it in his description. Fortunately, he mentioned it in the key to species. He does mention the pro- notal depression which is helpful in isolating elstoni from related species (Figs. 3, 6, 9, 11 and 14). The material studied has been found to vary from various localities in Tasmania, Australia, New Guinea, Vietnam and possibly Rennell Island. A male from Charters Towers Qld., Coll. Sedlacek (Bishop Museum) has a bright red scutellum. Eyes orange-brown and pronotum straw yellow. The hemelytra are hyaline ex- cept for the inner claval margin adjacent to the | Figures 1-3 — Anisops elstoni Brooks '? 13 ■ 15 9 14 Figures 10-12— .4. elstoni 26 . :_,v _ /. SSs^Cwi Figures 25-29— A. philippinensis <£ Fraser Island, Qd. } Boomerang South: 25, front fe- mur. 26, tibia. 27, tarsus. 28, head and pronotum from above. 29, head from side. typical deanei. In Brooks (1951) this form keys out to couplet 53 (philippinensis and win- di Brooks) the latter is distinguished by having the synthlipsis almost the same width as the anterior width of the vertex. In Lansbury (1969) it keys out as either deanei or nabilla Lansbury. The rostral prong of nabilla is smal- ler, rather slender, apically acuminate and the front femur is broader distally than in deanei. Bensink and Burton (1975) refer to a species 'near philippinensis' Det. Lansbury from Blue and Brown Lakes on North Stradbroke Island. This is the brachypterous form of deanei and forms an interesting parallel with those des- cribed by Young (1962) for New Zealand spe- cies. Bensink and Burton (1975) found as on Fraser Island (Bayly et aL, 1975) that no fish were present in the lake with the more diverse water-bug fauna (nine species) Brown Lake; :.- - I i^i^ L :-y,0 30 Figures 30-32 — A. deanei Brooks ^ is f Figures 33-36— ,4. rigoensis Brooks 6*, 33-35, paratype. 36, Genjam, N.G.: 33, front leg. 34, head from side. 35 and 36, head and pronotum from above. Figures 37-39 — A. biroi Brooks 7. Pereon with obsolete dor- solateral grooves, obsolete dorsal pits on pe- reonites 4-6. Pleon about as long as pereonite 7, pleonites distinct. Antenna 1 flagellum of 3-4 articles, first the longest, shorter than last two articles of ped- uncle. Antenna 2 flagellum rudimentary, of 4-5 short setose articles. LEPTANTHURA AND RELATED GENERA 137 Mandible with an acute incisor, palp not reaching to end of incisor; palp article 2 three times length of first, bearing 1 (up to 3 on large specimens) setae distally; palp article 3 small with 2 stout, barbed spines terminally. Maxilla a slender serrated spine. Maxilliped basis not distinct from head, bearing 2 (up to 4) ventral setae distally; maxillipedal palp of 3 articles (but not clearly distinct), article 1 about | length of basis, bearing 1 dorsal and 2-3 ventral setae distally, articles 2 and 3 minute, together with 4-7 setae. Pereopod 1 stout; palm axial and with a broadly-based thumb proximally. Article 5 of pereopod 1 with 2-3 spines; palm of article 6 with 7-9 spines laterally, a simple stout seta after the second spine, and 9-10 simple setae mesially. Pereopods 2, 3 similar to first but less stout, more elongate and palms with smal- ler spines. Pereopods 4-7 progressively a little more elongate; article 5 small, triangular and bearing 2-3 spines; article 6 with setae and 3-4 spines posteriorly; dactyl about as long as palm of article 6. Uropodal endopod triangular, setose, about § length of peduncle; exopod divided by a deep notch into an acute ventral lobe and a larger, erect dorsal lobe with setose margins. Telson little shorter than uropod, dorsally con- cave, its lateral margins curved to an evenly rounded apex; statocyst opening by a proximal pore; 2 long erect setae and several smaller setae dorsally, 2 small setae at apex, sometimes in a minute notch. Male: Differs from description given above in more elongate form; multi-articulate, setose flagellum of antenna 1 with about 20 articles reaching back to end of pereonite 1 ; more elon- gate pereopods; palm of pereopods with more simple setae laterally and highly setose me- sially; appendix masculinis with a tapering fla- gellum extending beyond the inner ramus of pleopod 2. In sub-males the multi-articulate flagellum of antenna 1 lacks setae and the appendix mas- culinis is missing or lacks its flagellum. Material examined: 19 males, 71 females, 531 ju- veniles; 5-20 mm. Holotype: AM P3317, juvenile, 13 1 mm. Type locality: TASMANIA, Hobart, 'between tide marks'. Other material: QUEENSLAND. Middle Banks, Moreton Bay, QUBS stn 5, December 1972 (1 specimen). N.S.W. Burwood Beach, 3 5 km S. of Hunter R., 14 m, HDWBS samples: AM P24027 (1 specimen). E. of Malabar, Sydney, 31-192 m, AMSBS stations: stn Al, AM P24350 (9); stn D2, AM P22791 (1); stn 2E, AM P22792 (4); stn 4E, AM P22793 (1); stn III, AM P22782 (6), AM P22783 (7); stn V, AM P22785 (3), AM P22786 (1); stn 31, AM P22788 (1); stn 44, AM P22787 (5). 2 km E. of Long Bay, Sydney, 66 m, AMSBS stn IV, AM P22784 (3). E. of North Head, Sydney, 20 m, AMSBS samples collected in association with the sponge Polymastea craficia: AM P22813 (1), AM P24362 (1). AMSBS station 22, AM P22789 (1). Near Kurnell, Botany Bay, AM P8965 (1). Near Sow and Pigs Reef, Port Jackson, AM P8969 (2). VICTORIA. Port Phillip Bay, 5-22 m, NMV J475, PPBES stations: stn 906 (3 specimens); stn 907 (11); stn 913 (32); stn 918 (29); stn 921 (1); stn 922 (7); stn 925 (1); stn 927 (2); stn 928 (3); stn 932 (2); stn 945 (4); stn 954 (2); stn 955 (1); stn 959 (2); stn 961 (1); stn 968 (1); stn 977 (1); stn 978 (1); stn 983 (1); stn 984 (67); stn 985 (10); stn 1224 (2); stn 1226(1). Western Port, 2-19 m, 1964 CPBS stations, NMV J476: stn Al (3); stn A4 (2); stn B2 (4); stn B3 (4); stn B4 (1); stn CI (3); stn C2 (1); stn C4 (2). 1965 CPBS stations, NMV J477: stn 01S (21); stn 02N (1); stn 03S (3); stn 10E (2); stn UN (49); stn US (24); stn 12N (1); stn 200 (8); stn 21N (10); stn 21S (10); stn 22N (6); stn 22S (16); stn 23N (6); stn 26N (1); stn 26S (6); stn 300 (8); stn 31N (6); stn 31S (6); stn 31E (1); stn 32N (15); stn 32S (1); stn 33N (14); stn 34N (4); stn 35N (1); stn 400 (1); stn 41N (4); stn 41S (5); stn 42S (2); stn 51N (21); stn 51S (9); stn 52N 138 GARY C. B. POOR!'. (7); stn 600 (13); stn 61N (6). 1966-70 CPBS stations, NMV J478: stn 300 (4); stn 31N (17); stn 31S (12); stn 32N (34); stn32S(13). Distribution: Southern Queensland, New South Wales, Victoria and Tasmania; intertidal region to 192 m, on fine to coarse sandy sediments, often with shell. Specimens from N.S.W. were taken in association with the sponge Poly mas- tea craficia. Remarks: Leptanthura diemenensis is the most common paranthurid in benthic collections from the south-eastern Australian coast. It is most easily distinguished by the notched exo- pod on the uropod and the broad, rounded, dished telson. Specimens may be confused with Bullowanthura pambula with which it co- occurs. L. diemenensis is most closely related to L. laevigata (Stimpson) from South Africa (Barnard, 1925) the only other species re- ported to have an apical notch in the uropodal exopod. The species differ in the form of pereopods 1. Leptanthura nunana sp. nov. Figures 4, 5 Description: Head as wide as long, about i length of pereonite 1; rostrum triangular, al- most as long as lateral lobes; eyes absent. Pe- reonite lengths as follows: 1 = 2 — 3 < 4 = 5 = 6 > 7. Pereon with obsolete dorsolat- eral grooves, obsolete dorsal pits on pereonites 4-6. Pleon almost twice as long as pereonite 7, pleonites distinct. Antenna 1 flagellum of 4 articles, first article very short and surrounding second laterally, flagellum shorter than last article of peduncle. Antenna 2 flagellum rudimentary, of 5-7 short, setose articles, as long as elon- gate last article of peduncle. Mandible with an acute incisor, palp not reaching to end of incisor; palp article 2 three times length of first, without setae; palp article 3 small with 2 stout, barbed spines ter- minally. Maxilla a slender, serrated spine. Maxilliped basis not distinct from head, bear- ing 1 ventral seta distally; maxillipedal palp of 3 almost-fused articles, article 1 about i length of basis, bearing 1 dorsal and 2 ventral setae distally, articles 2 and 3 minute, to- gether with 5 setae. Pereopod 1 stout; palm oblique and with a triangular thumb proximally. Article 5 of pereopod 1 with 1 spine; article 6 with 12-14 spines laterally, a simple stout seta after the third spine, and simple setae mesially. Pereo- pods 2, 3 similar to first but much less stout, more elongate and palms with smaller and fewer spines. Pereopods 4-7 subequal; article 5 small and triangular, bearing 1 spine; article 6 with 3-4 spines posteriorly; dactyl a little shorter than palm of article 6. Uropodal endopod elongate, extending well beyond telson, setose, about 1 length of ped- uncle; exopod lanceolate and with an acute apex, with setose margins. Telson little longer than uropod peduncle, dorsally convex, lateral margins parallel and apex evenly rounded; statocyst opening to a proximal dorsal pore; several setae dorsally and apically. Male: Differs from description given above in more elongate form; multi-articulate, setose flagellum of antenna 1 with about 9 articles reaching back little beyond end of head; more elongate pereopods; palm of pereopod 1 setose mesially; appendix masculinis with a curved end extending well beyond the outer ramus of pleopod 2. Material examined: 1 male, 1 female, 7 juveniles; 6-10 mm. Holotvpe: NMV J479, juvenile, 9-7 mm. Type locality: VICTORIA, Altona Bay, Port Phillip Bay, PPBES stn 901, sandy sediment, 8 m, 7 June 1971. Paratypes: VICTORIA. Port Phillip Bay, PPBES stations: stn 933, AM P25373, 25374 (2 specimens); stn 954, NMV J480, 481 (2); stn 955, NMV J482 (1); stn 983, NMV J483 (1). Other material: VICTORIA. Port Phillip Bay, PPBES stations: stn 914, NMV J484 (1 specimen); stn 934, NMVJ485(1). Distribution: Port Phillip Bay, Victoria, clayey- sand to sandy sediments, 8-22 m. LEPTANTHURA AND RELATED GENERA 139 Figure 1 — Leptanthura diemenensis. Female (NMV J 475, PPBES stn 983): a, lateral (limbs omitted); b, head; c, pereonite 6, pleon and tail fan; d, e, antennae 1, 2. 140 GARY C. B. POORE Figure 2 Leptanthura diemenensis. Female (NMV J475, PPBES stn 983) : a-d, e, pereopods LEPTANTHURA AND RELATED GENERA 141 SMI /, Figure 3—Leptanthura diemenensis. Female (NMV J475, PPBES stn 983): a, mandible; b, maxilliped. Male (NMV J475, PPBES stn 961): c, d, pereopods 1, 2 (mesial views); e, pleopod 2. Remarks: Leptanthura nunana is distinguished form of the telson and pereopods. by the long narrow uropodal endopods and the The Australian aboriginal word 'nunana' lanceolate exopods. The latter character is means 'little' and suggests the small size of this shared with L. agulhasensis Kensley from species. South Africa but the two species differ in the 142 GARY C. B. POORE Figure 4—Leptanthura nunana. Holotype (NMV J479): a, head; b, pleon and tail fair c, flagellum of antenna 2; d, mandible; e, maxilliped. LEPTANTHURA AND RELATED GENERA 143 Figure 5 — Leptanthura nunana. Holotype (NMV J479): a-d, e, pereopods 1-4, 7. Male (NMV J480, PPBES stn 954): f, pleopod 2. 144 GARY C. B. POORE Leptanthura kapala sp. nov. Figures 6, 7 Decription: Head a little wider than long, about fi as long as perconite 1; rostrum broadly tri- angular, i length of lateral lobes; eyes absent. Pereonite lengths as follows: 1 — 2 = 3 < 4 = 5 = 6 > 7. Pereon with obsolete dorso- lateral grooves, obsolete dorsal pits on pereo- nites 4-6. Pleon about as long as pereonite 7, pleonites distinct. Antenna 1 flagellum of 3-4 articles, shorter than last two articles of peduncle. Antenna 2 flagellum rudimentary, of 3-4 short setose ar- ticles, about as long as last article of peduncle. Mandible with an acute incisor, palp not reaching to end of incisor; palp article 2 al- most 3 times as long as first, without setae; palp article 3 small and with 2 stout barbed spines terminally. Maxilla a slender, serrated spine. Maxilliped basis not distinct from head, bearing 1-2 ventral setae distally; maxillipedal palp of 3 articles (but not clearly distinct); ar- ticle 1 about J length of basis, bearing 1 dorsal and 2 ventral setae distally; articles 2 and 3 minute, together with 4 setae. Pereopod 1 stout; palm oblique and with a broadly-based thumb proximally. Article 5 of pereopod 1 with 1 spine; palm of article 6 with 1 1-14 spines laterally, a simple stout seta after the third spine, and several simple setae me- sially. Pereopod 2 smaller than first and with- out a thumb; pereopod 3 much smaller than first, without a thumb but the palm posteriorly lobed. Pereopods 4-7 progressively a little more elongate; article 5 small and triangular, bearing 1 posterior spine and setae; article 6 with 2-3 posterior spines; dactyl longer than palm of article 6. Uropodal endopod triangular, setose, about i length of peduncle; exopod oval, its dorsal lobe overlying the telson, with setose margins. Telson reaching halfway along endopod, dor- sally flat, its lateral margins tapering to an acute apex; statocyst opening to a proximal dorsal pore; 2 long, erect setae dorsally. Male: Differs from description given above in moderately short, setose flagellum of an- tenna 1 with 8 articles reaching back to end of head; more elongate pereopods; palm of pereopod 1 with many setae mcsially; appendix masculinis with a curved tip reaching to the end of the outer ramus of plcopod 2. Materia! examined: 4 males, 4 females, 48 juven- iles; 7-1 1 mm. Holotype: AM P25375, juvenile, 9 8 mm. Type locality: N.S.W., E. of Wollongong, 34 27'S., 151° 27'E., F.R.V. 'Kapala' station K76-23-02, 1200 m, 13 December 1976. Paratypes: N.S.W. Type locality, AM P25376 (12 specimens). Other material: N.S.W. Type locality, AM P25055 (43 specimens). Distribution: New South Wales, bathyal zone, 1200 m. Remarks: Leptanthura kapala is the only an- thuridean so far described from the bathyal zone around Australia's coast. The species is recognized by the acute telson and broad uro- podal exopod. L, agulhasensis Kensley from South Africa and L. tenuis (Sars) from the North Atlantic share the acute telson but the uropods and pereopods of these species differ. The species is named for the N.S.W. State Fisheries research vessel, F.R.V. 'Kapala', which collected these specimens and has made some of the first biological collections from deep waters in Australia. Bullowanthura new genus Diagnosis: Paranthuridae without eyes. Pereon with feeble dorsolateral grooves, otherwise smooth; pereonites 4-6 with small dorsal pits. Pleonites distinct. Telson thin, concave dor- sally, not indurated; a single statocyst opening by a dorsal pore proximally. Uropod with en- dopod barely exceeding telson, exopod broad and erect. Antenna 1 flagellum rudimentary, of 3-4 articles. Antenna 2 flagellum rudimentary, of 4-5 articles. Mandible with a palp of a single article bearing one terminal seta. Maxil- liped elongate, articles 1 and 2 fused to head; endite obsolete; palp about J length of basis, of 2 articles, the last minute. Pereopod 1 stout, subchelate, palm with a proximal thumb, ob- lique. Pereopods 2, 3 much less well developed than first, barely subchelate. Pereopods 4-7 LEPTANTHURA AND RELATED GENERA 145 Figure 6 — Leptanthura kapala. Holotype (AM P25375): a, head; b, tail fan; c, mandibu- lar palp (terminal articles); d, maxilliped. Juvenile (AM P25376): e, mandibular palp. with article 5 triangular and lacking a free an- terior margin. Pleopod 1 operculiform, only moderately indurated. Adult male characterized by more elongate pereopods than juveniles or females and by a multi-articulate, setose flagel- lum on antenna 1. Females with oostegites on pereonites 2-5. Type species: Bullowanthura pambula new spe- cies. Remarks: Bullowanthura is close to Leptan- thura, differing in having a mandibular palp of a single article. Leptanthura possesses a 3- articulate palp of characteristic form. Both genera are blind, have distinct pleonites, have similar telson and uropods, similar antennae, maxillipeds and pereopods 4-7. Barnard (1925) recognized the constant form of the mandibular palp in Leptanthura but noted variations in other genera. He did not find 'it always possible to use these differences as generic characters'. Nevertheless, three an- thurid genera, Pendanthura, Ptilanthura and Xenanthura, and three paranthurid genera, Cruregens, Colanthura and possibly Cruran- thura, lack a mandibular palp so it seems the character may have some value. Bullo- wanthura and the following new genus, Via- I U) (.aky C. ii. poori: x s x *, Figure 1 Leptanthura kapala, I lolotype P25375): a-d, e, pereopoda 1-4, 7. (AM kanthura t are the only paranthurids with a palp of a single article. Bullowanthura and Ulakanthura differ in ihc degree of development of pereopods 2 ami 3. In Bullowanthura pereopods 2 and 3 arc poorly developed and approach walking legs in form. In Ulakanthura these two limbs are larger than the first pereopod. Pereopods 4-7 are also different in the two genera. The genus is monolypie. The prefix 'bullowa" used in formation of the generic name is an Australian aboriginal word meaning 'flower' and reflects the mean- ing of the root 'anthos' used in most generic names in the family. Bullowanthura pambula sp. now Figures 8, 9 Description: Head as wide as long, about i as long as perconite 1; rostrum broadly tri- angular, 3 length of lateral lobes; eyes absent. Pereonite lengths as follows: 1 -: 2 — 3 < 4 = 5 = 6 > 7. Pereon with obsolete dorso- lateral grooves, obsolete dorsal pits on pcrcon- ites 4-6. Pleon little longer than perconite 7, pleonites distinct Antenna I ilagellum of 3-4 articles, first the longest, little longer than last article of ped- LEPTANTHURA AND RELATED GENERA [47 ■uncle. Antenna 2 flagellum rudimentary, of 4-5 short setose articles, Mandible with an acute incisor, palp a single narrow article with 1 terminal seta. Maxilla a slender, serrated spine bearing a minutely hooked keel on its distal inner margin. Maxil- liped basis not distinct from head, bearing 2-3 ventral setae distally; maxiliipedal palp of 2 articles, first about \ length of basis and bear- ing 1 dorsal seta and 4 ventral setae, second minute and with 3 terminal setae. Pereopod 1 stout; palm oblique and with a strong proximal thumb. Article 5 of pereopod 1 with 2-4 spines; palm of article 6 with 7-13 spines laterally and in the same row a long simple seta near the base of the thumb and another after the last spine; palm with 8-12 simple setae mesially. Pereopods 2, 3 unlike the first, article 6 barely thickened, palms lack- ing a thumb and bearing 6 spines. Pereopods 4-7 subequal; article 5 small and triangular, bearing 2 spines; article 6 with 2-4 posterior spines; dactyl about as long as article 6. Uropodal endopod elongate-triangular, set- ose, little shorter than peduncle; exopod divided into two parts by a very shallow ter- minal notch, a smaller ventral lobe with a rounded apex and a larger semicircular dorsal lobe. Telson reaching more than halfway along endopod, dorsally concave, lateral margins curved to a broadly rounded apex; statocyst opening to a proximal dorsal pore; 2 pairs of setae terminally and a few dorsally. Male: Differs from the above description in multi-articulate, setose flagellum of antenna 1 with about 10-12 articles reaching back to end of head; more elongate pereopods; palm of pereopod 1 with a longer and narrower thumb, bearing 11-14 spines laterally and many setae mesially; appendix masculinis a simple rod ex- tending well beyond the outer ramus of pleo- pod2. Material examined: 6 males, 3 females, 17 ju- veniles; 3-11 mm. Holotype: AM P25378, juvenile, 10 3 mm. Type locality: N.S.W., 1-6 km E. of Malabar, Sydney, AMSBS stn III, 33° 58'S., 151° 17'E., dredged from 66 m, 31 July 1973. Paratypes: N.S.W. E of Malabar, Sydney, 66-69 m, AMSBS stations: stn III, AM P25379 (6 speci- mens), AM P22814 (2), AM P25380 (3); stn V, AM P25382 (2); sin 4C, AM P22790 (1). E. of Long Bay, Sydney, 66 m, AMSBS sta- tion IV, AM P25381 (3). VICTORIA. Port Phillip Bay, 8-25 m, PPBES stations: stn 901, NMV J486-J488 (3); stn 906, NMV J489 (1); stn 969, NMV J554 (1); stn 982, NMV J555-J557 (3). Distribution: New South Wales and Victoria, 8-69 m, on sandy sediments. Remarks: The species has been previously con- fused with Leptanthura dicmenensis with which it co-occurs. Bullowanthura pamlnda may be separated on its smaller size and more pro- nounced thumb on pereopod 1 and, more de- finitely, on the uni-articulate mandibular palp. The series from Port Phillip Bay differed slightly from those off Sydney in features which could only be attributed to their slightly smal- ler size. Jn general, smaller specimens have fewer spines on the cutting edges of articles 6 of the pereopods, the number on pereopod 1 , for example, showing quite a wide range. The Australian aboriginal word 'pambula' means 'two waters' and indicates the distribu- tion of the species. Ulakanthura new genus Diagnosis: Paranthuridae without eyes. Pc- rcon with feeble dorsolateral grooves, other- wise smooth; perconites 4-6 with small dorsal pits. Pleonites distinct. Telson thin, concave to moderately convex dorsally, not indurated; a single statocyst opening by a dorsal pore proxi- mally. Uropod with endopod exceeding tel- son by about half its length, exopod usually broad and erect. Antenna 1 flagellum rudi- mentary, of 3-4 articles. Antenna 2 flagellum rudimentary, of 4-5 articles. Mandible with a palp of a single article without setae. Maxilli- ped elongate, articles 1 and 2 fused to head; endite obsolete; palp about one-third of length of basis, of 2 or 3 articles, the last minute. Pereopod 1 stout, subchelate, palm with a proximal thumb, oblique-transverse. Pereopods 2 and 3 stout, second at least more well de- 148 GARY C. B. POORE Figure S—Bullowanthura pambula. Holotype (AM P25378): a, head; b, pleon and tail fan; c, mandible; d, maxilliped. Male (AM P25382, AMSBS stn V): e, pleopod 2. LEPTANTHURA AND RELATED GENERA 149 Figure 9—Bullowanthura pambula. Holotype (AM P25378): a-d, e, pereopods 1-4, 7. Male (NMV J486, PPBES stn 901): f, pereopod 1 (mesial setae omitted). Male (AM P25382, AMSBS stn V): g, pereopod 1 (mesial setae omitted). [50 GARY C. B. POORE 3 Telson broadest terminally, apical lobes squarish; article 4 of pereopod 4 with posterior lobe extending dis- tally almost to end of article 6 U. cooma — Telson widest at midpoint, apical lobes rounded; article 4 of pereopod 4 with posterior lobe extending distally to end of article 5 U. crassicornis 4 Telson tapering to two acute terminal lobes; uropodal exopod lanceolate (i as wide as long) U. namoo — Telson ending in two broadly semicir- cular lobes; uropodal exopod ovate (§■ as wide as long) 5 5 Telson widest at proximal one third; appendix masculinis of male shorter than outer ramus of pleopod 2 , ... U* lata — Telson parallel-sided; appendix mas- culinis of male longer than outer ra- mus of pleopod 2 U. wanda Ulakanthura crassicornis (Haswell) veloped than first; articles 2 and 6 especially broad. Pereopods 4-7 with article 5 lobed pos- teriorly and extending along posterior margin of more distal articles; article 5 triangular and lacking a free anterior margin; articles 5 and 6 with strong posterior spines and article 7 with terminal articulating spines. Pleopod 1 oper- culiform, only moderately indurated. Adult male characterized by multi-articulate setose flagellum on antenna 1 and more elongate limbs. Females with oostcgites on pereonites 2-5. Type species: Paranthura crassicornis Haswell, 1881. Remarks: Ulakanthura is very close to Lep- tanthura and the preceding new genus, Bui- lowanthura. All three genera are blind, have distinct pleonites and have similar telsons, uro- pods, antennae and maxillipeds. Leptanthura is distinguished by the 3-articulate mandibular palp of constant form. The other two genera share a mandibular palp of a single article, a feature found in few other anthurideans. The Figures 10, 11 terminal seta on the palp of B. pamhula is not p aran thura (?) crassicornis Haswell, 1881: 478, found in species of Ulakanthura described here. pI 18 ^ f]g 5._Haswell, 1882: 305. Ulakanthura is notable for the relative size Leptanthura crassicornis, — Barnard, 1925: of pereopods 1-3, the first being the smallest. 151. Nierstrasz, 1941: 232 (part). This relationship is unknown in other paran- no t Paranthura crassicornis. — Haswell, 1884: thurids. The posterior lobe on article 4 and the 1011-2, pi. 53, figs. 8, 9 (—Paranthura terminal articulating spines which replace the sp> ) # unguis on article 7 of pereopods 4-7 are also not Calathura, sp. — Stebbing, 1905: 8 (from not known in other paranthurid genera. Sri Lanka). The genus is found only from eastern Aus- not Leptanthura crassicornis (Haswell). Nier- tralia where at present six species are known. strasz, 1 941 : 242 (part from Sri Lanka). The prefix 'ulaka' used in formation of the Description: Head about as long as greatest generic name is an Australian aboriginal word w idth, tapering anteriorly, almost as long as meaning 'flower' and reflects the meaning of pe reonite 1; rostrum broadly triangular, about the root 'anthos 1 used in most generic names j_ as ] ong as lateral lobes; eyes absent. Pereon- in the family. ite lengths as follows 1<2 = 3<4 = 5 = 6 > 7. Pereon with dorsolateral grooves, ,r ^^^nnn. dorsal pits and paired rows of setae on dorsum KEY TO SPECIES OF ULAKANTHURA ^ p J QnltQS / 6 . p leon as long as pere onite 1 Uropodal exopod apically cleft 2 ? ^ pleon i tes distinct. — Uropodal exopod not apically cleft 4 Antenna 1 flagellum of 3-4 articles, shorter 2 Maxillipedal basis with many (>7) than last article of peduncle. Antenna 2 flagel- ventral and lateral setae £/. colac lum rudimentary, of 4 setose articles. — Maxillipedal basis with few «4) ven- Mandible with an acute incisor, palp a tral and lateral setae 3 single tapering article, not reaching to midpoint LEPTANTHURA AND RELATED GENERA 151 of mandible. Maxilla a slender, serrated spine bearing a minutely-hooked keel on its distal tinner margin. Maxilliped basis not distinct ifrom head, bearing 1 ventral seta distally; imaxillipedal palp of 2 barely-distinct articles, ;about 4 length of basis and bearing 8 ventral- distal setae and 1 dorsal-proximal seta. Pereopod 1 stout; palm oblique-transverse and with a broad square thumb proximally. Article 5 of pereopod 1 with 3-4 spines; article 6 with 12-15 spines laterally along the cutting edge and a stout seta in the same row after the fourth spine. Pereopods 2, 3 stouter than first, articles 2 and 6 broader than those of first. Article 5 of pereopods 2, 3 without spines; palm of article 6 with a blunt proximal thumb and a row of about 16 spines laterally. Pereo- pods 4-7 dissimilar to pereopods 1-3, subequal; article 4 of pereopods 4-7 setose, posteriorly lobed, the lobe not reaching end of article 5 on pereopod 4 and about -J way along article 5 on pereopod 7; article 5 triangular, bearing 3-4 strong posterior spines; article 6 with 4-5 strong posterior spines; article 7 proximally curved, not tapering, with 1 short and 1 long terminal spine. Uropodal endopod triangular, setose, about | as long as peduncle; exopod divided by a clear notch into a ventral lobe with an acute apex and a larger erect dorsal lobe, with setose margins. Telson widest at midpoint and reach- ing halfway along endopod, dorsally flat-con- cave, divided by a deep wide notch into two semi-circular lobes; statocyst opening to a dor- sal pore at the base of the telson; two pairs of setae in the terminal notch. Male: Differs from the above description in possessing an elongate, setose flagellum on an- tenna 1 with about 14 articles reach back to end of pereonite 1; axial setose palm of pereo- pod 1; longer dactyls on pereopods 4-7; ap- pendix masculinis on pleopod 2. The only adult male is the 'holotype' which is not in good condition, the remaining males examined lacked setae on the antenna 1 flagellum. Material examined: 5 males, 1 female, 5 juveniles; 10-13 mm, Holotype: AM P3316, male, 12.5 mm. L Type locality: N.S.W., Port Jackson. Other material: N.S.W. Belmont Beach, 16 km S. of Hunter R., 22 m, HDWBS samples AM P24021-24025 (6 specimens). Burwood Beach, 3 5 km S. of Hunter R., 22-28 m, HDWBS samples: AM P24028 (2), AM P24029 (1). E. of Malabar, Sydney, 31 m, AMSBS stn A1:AMP25383(1). Distribution: Central New South Wales, 22-3 1 m, coarse sandy sediments. Remarks: Three paranthurids are catalogued together in the Old Collection of the Austra- lian Museum as P3316 under the name Paran- thura crassicornis. I have deduced that one of these, a male, is the specimen described by Haswell in 1881 and its large setose antenna 1 suggested the specific name to Haswell. In 1884 Haswell ascribed two more specimens to P. crassicornis noting differences in the an- tennae. The two other specimens stored as P3316 are probably those referred to by Has- well in 1884 but are not the same species as the original. Haswell (1881) remarked that pereopod 1 was larger than the following ones, a feature common in most anthurids. This is not so in U. crassicornis but is not obvious at first glance. Ulakanthura crassicornis is one of three spe- cies in this genus in which the uropodal exo- pod is cleft. The others are U. cooma from which it differs in form of the pereopods and telson, and U. colac from which it differs in the same characters and in the number of setae on the maxilliped basis. U. crassicornis is sympatric only with U. cooma. Stebbing (1905) suggested that two speci- mens collected in Ceylon (Sri Lanka) may be- long to Paranthura (?) crassicornis Haswell. Nevertheless, he listed them under the name Calathura, sp. and remarked on their dark eyes. This character clearly excludes them from Ulakanthura crassicornis. 152 GARY C. B. POORE Figure 10 — Ulakanthura crassicornis. Juvenile (AM P24021, HDWBS sample): a, head; b, tail fan; c, mandible; d, maxilliped. Ulakanthura colac sp. nov. Figures 12, 13 Description: Head about as long as greatest width, tapering only anteriorly, about i length of pereonite 1; rostrum broadly triangular; eyes absent. Pereonite lengths as follows: 1 < 2 = 3<4 = 5 = 6>7. Pereon with obsolete dorsolateral grooves, dorsal pits and paired rows of few setae on dorsum of pereonites 4-6. Pleon little shorter than pereonite 7, pleonites distinct. Antenna 1 flagellum of 3-4 articles, little LEPTANTHURA AND RELATED GENERA 153 Figure 1 1 — Ulakanthura crassicornis. Juvenile (AM P24021, HDWBS sample): a, b, c, d, pereopods 1, 2, 4, 7. longer than last article of peduncle. Antenna 2 flagellum of 4-5" setose articles. Mandible with an acute incisor, palp a single article, about ^ length of incisor. Maxilla a slender, serrated spine bearing a minutely- hooked keel on its distal inner margin. Maxil- liped basis not distinct from head, bearing 1 distal ventral seta and 7-14 setae laterally; maxillipedal palp with a distinct, minute, ter- minal article; palp articles together about -J length of basis and with 1 dorsal seta and 6 ventral-distal setae. Pereopod 1 stout; palm axial-oblique and with a broad thumb proximally. Article 5 of pereopod 1 with 3-4 spines; article 6 with 7-10 154 GARY C. B. POORE spines laterally along cutting edge, a stout seta after the third or fourth and after the last spine. Percopod 2 stouter than pereopod 1, article 2 especially broader than that of first, palm oblique. Article 5 of pereopods 2, 3 with 2-5 spines; palm of article 6 with a proximal thumb and row of 6-9 spines laterally. Pereo- pods 4-7 dissimilar to pereopods 1-3, sub- equal. Article 4 of pereopods 4-7 setose, lobed posteriorly, the lobe reaching only a little be- yond the distal end of article 5 on percopod 4 and not reaching this point on pereopod 7; article 5 triangular, with 3-6 strong posterior spines; article 6 with 4-8 stout posterior spines; article 7 barely tapering, with 1 long and 1 short terminal spine. Uropodal endopod setose, triangular, about twice as long as wide, $ as long as peduncle; exopod broad and with a shallow apical notch, folded over telson. Telson reaching halfway along endopod, dorsally concave, more or less parallel-sided, tapering distally to two rounded- triangular lobes separated by a strong apical notch; statocyst opening by a small proximal dorsal pore; one pair of simple setae and one pair of 'brush-setae' in the terminal notch and setae on dorsal surface of the telson. Male: Differs from the above deescription in more elongate palm on pereopod 1 ; setose fla- gellum of antenna 1 with 12 articles reaching back to middle of pereonite 1. Appendix mas- culinis a simple rod reaching to the end of the outer ramus of pleopod 2. Materia! examined: 7 males, 3 females, 49 ju- veniles; 4-12 mm. Holotype: QM W7400, juvenile, 7 8 mm. Type locality: QUEENSLAND. Middle Banks, Moreton Bay, clean sand, 9-10 m, collected by S. Cook and S. Newlands for University of Queensland benthic studies, March 1973. Paratypes: QUEENSLAND: Middle Banks, Moreton Bay, QUBS samples: QM W7344 (4 speci- mens), QM W7345 (6), QM W7346 (1). Other material: N.S.W. Burwood Beach, 3 5 km S. of Hunter R., 14-22 m, HDWBS samples: AM P24026 (1 specimen), AM P25384 (1). Botany Bay, SPCC stations: stn 18, AM P25360 (1); stn 65, AM P25366 (1); stn 83, AM P25367 (1 ); stn 85, AM P25368 (1). VICTORIA: Port Phillip Bay, 5-10 m, PPBES stations: stn 960, NMV J490-494 (44); stn 967,NMVJ495(1). Distribution: Southern Queensland, New South Wales and Victoria, well-sorted sandy sedi- ments, 5-22 m. Remarks: Ulakanthura colac is a widespread species found in all shallow water coastal sur- veys from eastern and southeastern Australia. The species is recognized by the combination of cleft uropodal exopod and setose maxilliped basis (see Remarks for U. crassicornis and U. cooma). The epithet 'colac' is an aboriginal word for 'sand* and reflects the sandy habitat of this and other species in the genus. Ulakanthura cooma sp. nov. Figures 14, 15 Description: Head about as long as greatest width, tapering anteriorly, about I as long as pereonite 1 ; rostrum broadly triangular; eyes absent. Pereonite lengths as follows: 1 < 2 = 3<4 — 5 — 6>7. Pereon with dorso- lateral grooves, dorsal pits and paired rows of setae on dorsum of pereonites 4-6. Pleon little shorter than pereonite 7, pleonites distinct. Antenna 1 flagellum of 4 articles, about as long as last article of peduncle. Antenna 2 flagellum of 4 setose articles. Mandible with an acute incisor, palp a single article, about I as long as incisor. Maxilla a slender, serrated spine bearing a minutely- hooked keel on its distal inner margin. Maxilli- ped basis not distinct from head, bearing 1 dis- tal and 1-2 subdistal setae; maxillipedal palp with a barely-distinct second article and a dis- tinct terminal article; palp articles together with 1 dorsal seta and 7 ventral-distal setae. Pereopod 1 stout; palm axial and with a broad thumb proximally. Article 5 of pereo- pod 1 with 5 spines; article 6 with 10-11 spines laterally along cutting edge, a stout seta after the fourth or fifth spine and another after the last spine. Pereopods 2, 3 stouter than pereo- LEPTANTHURA AND RELATED GENERA 155 Figure 12 — Ulakanthura colac. Holotype (QM W7400): a. tail fan; b, mandible; c, maxilliped. Male (QM W7344, QUBS sample) : d, pleopod 2. pod 1, articles 2 and 6 broader than those of first, palm oblique. Article 5 of pereopods 2, 3 with 1-3 spines; palm of article 6 with a proximal thumb and a row of 12-14 spines lat- erally. Pereopods 4-7 dissimilar to pereopods 1-3, subequal. Article 4 of pereopods 4-7 set- ose, strongly lobed posteriorly, the lobe reach- ing almost to the distal end of article 6 on pereopod 4 and halfway along article 6 on pereopod 7; article 5 triangular, with 6-8 strong posterior spines; article 6 with 8-10 stout posterior spines; article 7 barely tapering, with 1 long and 1 short terminal spine. Uropodal endopod setose, linear-triangular, about 3 times as long as wide, almost as long as peduncle; exopod broad and apically notched, folded over telson. Telson reaching little beyond the end of the peduncle, dorsally concave, widest distally, two almost truncate terminal lobes separated by a strong apical notch; statocyst opening by a proximal dorsal pore; 1 pair of simple setae and 1 pair of 'brush-setae' in terminal notch, telson setose dorsally. Male: Differs from above description in more elongate pi eon; more elongate and more set- ose palm on pereopod 1; setose flagellum on antenna 1 with 12 articles reaching back to middle of pereonite 1. Appendix masculinis a simple rod exceeding the inner ramus of pleo- pod 2 by about i its length. 156 GARY C. B. POORE Figure 13 — Ulakanthura colac. Holotype (QM W7400) : a-d, e, pereopods 1-4, 7. Material examined: 5 males, 13 females, 60 ju- veniles; 7-18 mm. Holotype: QM W7347, juvenile, 14-2 mm. Type locality: QUEENSLAND. Middle Banks, Moreton Bay, clean sand, 9-10 m, collected by S. Cook and S. Newlands for the University of Queensland benthic studies, September 1973. Paratypes: QUEENSLAND. Middle Banks, Moreton Bay, QUBS samples: QM W6132 (14 specimens), QMW6133(19). Other material: QUEENSLAND. Middle Banks, Moreton Bay, QUBS samples: QM W6125 (1 specimen), QM W6130 (9), QM W6131 (16), NMV J496(4),AMP25385(14). LEPTANTHURA AND RELATED GENERA 157 Distribution: Southern Queensland, clean sand, 9-10 m. Remarks: Ulakanthura cooma is a species of restricted distribution distinguished from others in the genus with a cleft uropodal exo- pod by the splayed, almost truncate telson and the particularly long lobes on articles 4 of pereopods 4-7. The aboriginal word 'cooma' means a 'sand bank' and indicates the environment from which this species has been taken. Ulakanthura lara sp. nov. Figures 16-18 Description: Head about as long as greatest width, strongly tapering anteriorly, about I length of pereonite 1; rostrum broadly triangu- lar, about i length of lateral lobes; eyes absent. Pereonite lengths as follows: 1 < 2 = 3 < 4 = 5 — 6 > 7. Pereon with dorsolateral grooves, dorsal pits and paired rows of setae on dorsum of pereonites 4-6. Pleon as long as pereonite 7, pleonites distinct. Antenna 1 flagellum of 4 articles, about as long as last article of peduncle. Antenna 2 flagellum of 4 setose articles. Mandible with an acute incisor, palp a single article about i as long as incisor. Maxilla a slender, serrated spine bearing a minutely- hooked keel on its distal inner margin. Maxilli- ped basis not distinct from head, bearing 6-14 lateral setae (most on larger specimens), 2 ventral setae and 1 dorsal seta distally; maxilli- pedal palp with a barely-distinct second article, palp about i length of basis, bearing 7 ven- tral-distal setae and 1 dorsal seta. Pereopod 1 stout; palm axial-oblique and with a short, broad thumb proximally. Article 5 of pereopod 1 with 4 ventral spines; article 6 with 10 spines laterally along the cutting edge, a stout seta after the fourth spine and another after the last spine. Pereopods 2, 3 stouter than pereopod 1, articles 2 and 6 broader than those of pereopod 1, palm oblique. Article 5 of pereopods 2, 3 with 1 posterior spine; palm of article 6 with a squarish proximal thumb and a row of about 13 spines laterally. Pereopods 4-7 dissimiliar to pereopods 1-3, subequal; posterior spines; article 6 with 6 stout pos- terior spines; article 7 proximally curved, not tapering, with 1 short and 1 long terminal spine. Uropodal endopod triangular, setose, about f length of peduncle; exopod broadly lanceo- late, without an apical notch. Telson reaching halfway along endopod, dorsally flat-concave, broadest at proximal third and tapering to two short, rounded lobes separated by an apical notch; statocyst opening to a dorsal pore at the base of the telson; 1 pair of simple setae and 1 pair of 'brush-setae' in the terminal notch and many setae dorsally on the telson. Male: Differs from the above description in more elongate pleon; more setose palm of pereopod 1 ; setose flagellum of antenna 1 with 8 articles reaching back to end of head. Appen- dix masculinis a simple rod reaching almost to end of outer ramus of pleopod 2. Materia! examined: 5 males, 22 females, 91 ju- veniles; 8-16 mm. Holotype: NMV J497, juvenile, 160 mm. Type locality: VICTORIA. 3 5 km NE of Point King, South Channel, Port Phillip Bay, PPBES stn 980, sandy sediment, 31 m, 12 October 1971. Paratypes: VICTORIA: Port Phillip Bay, 2-31 m, PPBES stations: stn 967, NMV J498 (3 specimens); stn 973, AM P25386 (2); stn 974, NMV J499, J500 (4); stn 975, AM P25387 (1); stn 980, AM P25388 (1); stn 981, AM P25389 (6); stn 984, NMC J501-J503 (12); stn 986, NMV J504, J505 (10). Western Port, 9-16 m, 1965 CPBS stations: stn 24N, NMV J447 (3); stn 31E, AM P25377 (1); stn 40E, NMV J448 (2). Other material: VICTORIA: Port Phillip Bay, 4-31 m, PPBES stations, NMV J508: stn 921 (5 specimens); stn 967 (7); stn 973 (10); stn 974 (4); stn 975 (4); stn 979 (2); stn 980 (1); stn 981 (9); stn 984 (4); stn 985 (1); stn 986 (9); stn 1244 (1); stn 1264 (1). Western Port, 10-14 m, 1964 CPBS stations, NMV J509: stn A6 (2); stn B2 (1); stn B6 (4); stnC5(l). 158 GARY C. B. POORE Figure 14— Ulakanthura cooma. Holotype (QM W7347): a, tail fan; b, c, d, e, pereopods 1, 2, 4, 7. LEPTANTHURA AND RELATED GENERA 159 Figure 15 — Ulakanthura cooma. Holotype (QM W7347): a, mandible; b, maxilliped. Male (QM W6135, MBBS sample): c, pleopod 2. article 4 setose, strongly lobed posteriorly, the lobe reaching to the distal corner of article 5 on pereopod 4 and not quite to this point on pereopod 7; article 5 triangular, bearing 5 stout 1965 CPBS stations, NMV J510: stn 24S (1); stn 35N (1); stn 36N (1); stn 41N (1); stn 42N (1). 1966-70 CPBS station, NMV J511: stn 32N (1). Distribution: Victoria, well-sorted coarse sandy sediments often with shell, 2-31 m. Remarks: Ulakanthura lara is the only species dealt with in this paper endemic to Victoria. The species is very close to U. wanda from Moreton Bay but differs in the more oval form of the telson and the length of the appendix masculinis of the male. The epithet 'lara' is an aboriginal place name from near Port Phillip Bay. Ulakanthura namoo sp. nov. Figures 19, 20 Description: Head a little longer than greatest width, tapering slightly anteriorly, about i length of pereonite 1 ; rostrum triangular; eyes absent. Pereonite lengths as follows: 1 < 2 = 3 <4 = 5 = 6>7. Pereon with obsolete dorsolateral grooves, dorsal pits and paired rows of setae on dorsum of pereonites 4-6. Pleon about as long as pereonite 7, pleonites distinct. Antenna 1 flagellum of 5 articles, about as long as last 3-4 articles of peduncle. Antenna 2 flagellum of 3 setose articles. Mandible with an acute incisor, palp a single article about i as long as incisor. Maxilla a slender, serrated spine bearing a minutely- hooked keel on its distal inner margin. Maxil- liped basis fused on to head, bearing 2 ventral setae and 5-8 lateral setae; maxillipedal palp with a barely distinct terminal article; palp articles together with 1 dorsal seta and 7 ven- tral-distal setae. Pereopod 1 stout; palm axial-oblique and with a broad thumb proximally. Article 5 of 160 GARY C. B. POORE Figure 16— Ulakanthura lara. Holotype (NMV J497): a, lateral (limbs omitted); b, head; c, tail fan. LEPTANTHURA AND RELATED GENERA 161 Figure 17 — Ulakanthura lara. Holotype (NMV J497): a, mandible; b, maxilla; c, maxilli- ped. Male (NMV J449, PPBES stn 974): d, pleopod 2. pereopod 1 with 3-5 spines; article 6 with 6-7 spines laterally along the cutting edge, a stout seta after the third or fourth spine and another after the last spine. Pereopods 2, 3 stouter than pereopod 1, article 2 particularly broader than that of pereopod 1, palm oblique. Article 5 of pereopods 2, 3 with 1 posterior spine; palm of article 6 with a proximal thumb and a row of 5-7 spines laterally. Pereopods 4-7 dissimilar to pereopods 1-3, subequal. Article 4 of pereopods 4-7 marginally setose, strongly lobed posteriorly, the lobe reaching distally midway along the posterior margin of article 6 on pereopod 4 and about | along article 6 on pereopod 7; article 5 triangular and with 4-5 strong posterior spines; article 6 with 5-6 stout posterior spines; article 7 barely tapering, with 1 short and 1 long terminal spine. Uropodal endopod setose, triangular, about twice as long as wide, about f as long as ped- uncle; exopod lanceolate, i as wide as long, not apically notched, folded over telson. Telson reaching halfway along endopod, dorsally con- vex, slightly tapering proximally but tip nar- rowing strongly to two acute lobes separated by a strong apical notch; statocyst opening by a proximal dorsal pore; 1 pair of simple setae and 1 pair of 'brush-setae' in the terminal notch and several setae dorsally on the telson. Male: Differs from the above description in slightly more elongate pleon; more setose and elongate palm on pereopod 1 ; setose flagellum on antenna 1 with 12 articles reaching back just beyond the posterior margin of the head. Appendix masculinis a simple rod just exceed- ing the inner ramus of pleopod 2. Material examined: 17 males, 16 females, 103 ju- veniles; 7-10 mm. Holotype: QM W7348, juvenile, 9-6 mm. Type locality: QUEENSLAND. Middle Banks, Moreton Bay, clean sand, 9-10 m, collected by S. Cook and S. Newlands for the University of Queensland benthic studies, June 1973. Paratypes: QUEENSLAND: Middle Banks, Moreton Bay, 9-10 m, QUBS samples: QM W6138 (19 specimens), QM W6140 (20). Other material: QUEENSLAND: Middle Banks, Moreton Bay, 9-10 m, QUBS samples: QM W6136 (14 specimens), QM W6137 (8), QM W6141 (16), QM W6143 (22), NMV J512 (17), AM P25457 (7). N.S.W.: Burwood Beach, 3 5 km S. of Hunter 162 GARY C. B. POORE Figure 18 — Ulakanthura lara> Holotype (NMVJ497): a, b, c, d, pereopods 1, 2, 4, 7. LEPTANTHURA AND RELATED GENERA 163 R., 14-16 m, HDWBS samples: AM P2403O- 24032 (3 specimens). Dudley Beach, 4-8 km S. of Hunter R., 10 an, HDWBS sample: AM P24039 (1). Belmont Beach, 16 km S. of Hunter R., 12-23 m, HDWBS samples: AM P24033- 24038 (8). Distribution: Southern Queensland to central New South Wales; clean coarse sand; 9-23 m. Remarks: Ulakanthura namoo is a clearly dis- tinguished species recognized by its lanceolate uropodal exopod and by the narrow acute api- cal lobes on the telson. U. wanda and [/. lara share with U. namoo a non-cleft exopod and also have a setose maxilliped basis but differ in the broader exopod and form of the telson. The aboriginal word 'namoo' means 'sand' and indicates the environment from which this species is taken. Ulakanthura wanda sp. nov. Figures 21, 22 Description: Head little longer than greatest width, tapering anteriorly, about as long as pereonite 1; rostrum broadly triangular; eyes absent. Pereonite lengths as follows: 1 < 2 = 3 = 4<5:=6>7. Pereon with dorso- lateral grooves, dorsal pits and paired rows of setae on dorsum of pereonites 4-6. Pleon about as long as pereonite 7, pleonites distinct. Antenna 1 flagellum of 3-4 articles, little longer than last article of peduncle. Antenna 2 flagellum of 4 setose articles. Mandible with an acute incisor, palp a single article, almost i as long as incisor. Max- illa a slender, serrated spine bearing a min- utely-hooked keel on its distal inner margin. Maxilliped basis not distinct from head, bear- ing 2 ventral and 8-10 lateral setae; maxilli- pedal palp with a barely-distinct terminal ar- ticle; palp articles together with 1 dorsal seta, 1 lateral seta and 7 ventral-distal setae. Pereopod 1 stout; palm axial and with a broad thumb proximally. Article 5 of pereopod 1 with 4-5 spines; article 6 with 10-11 spines laterally along the cutting edge, a stout seta after the fourth spine and another after the last spine. Pereopods 2, 3 stouter than pereopod l s articles 2 and 6 especially broader than those of pereopod 1; palm oblique. Article 5 of pereopods 2, 3 with 1 posterior spine; palm of article 6 with a proximal thumb and row of 12 spines laterally. Pereopods 4-7 dis- similar to pereopods 1-3, subequal. Article 4 of pereopods 4-7 setose, strongly lobed pos- teriorly, the lobe reaching about | way along the posterior margin of article 6 on pereopod 4 and just beyond end of article 5 on pereopod 7; article 5 triangular, with 4-5 strong pos- terior spines; article 6 with 6-7 stout posterior spines, article 7 barely tapering, with 1 short and 1 long spine terminally. Uropodal endopod setose, triangular, al- most 3 times as long as wide, about i as long as peduncle; exopod broad (about f as wide as long), not apically notched, folded over tel- son. Telson reaching almost halfway along en- dopod, dorsally concave, more or less parallel- sided, terminally two rounded lobes separated by a strong apical notch; statocyst opening by a proximal dorsal pore; 1 pair of simple setae and 1 pair of 'brush-setae' in the terminal notch and many setae on the dorsal surface of telson. Male: Differs from the above description in more elongate pleon; more setose and more elongate palm on pereopod 1; setose flagellum of antenna 1 with 11-14 articles reaching back to middle of pereonite 1 ; appendix mascu- linis a simple rod exceeding the inner ramus of pleopod 2 by about | its length and exceed- ing the outer ramus. Material examined: 4 males, 3 females, 12 ju- veniles; 7-11 mm. Holotype: QM W6124, juvenile, 10-7 mm. Type locality: QUEENSLAND. Middle Banks, Moreton Bay, clean sand, 9-10 m, collected by S. Cook and S. Newlands for the University of Queensland Moreton Bay benthic studies, December 1973. Paratypes: QUEENSLAND: Middle Banks, Moreton Bay, 9-10 m, QUBS samples: QM W6135 (1 speci- men), QM W6139 (1), QM W7349 (5). Other material: QUEENSLAND: Middle Banks, Moreton Bay, 164 GARY C. B. POORE Figure 19— Ulakanthura namoo. Holotype (QM Distribution: Southern Queensland, clean W7348): a, head; b, tail fan; c, mandible; cand 9 10 m d, maxilliped. Male (QM W6140, QUBS ' sample) : e, pleopod 2. Remarks: This species is closest to U. lara from Victoria differing in the form of the tel- son and length of the appendix masculinis of 9-10 m, QUBS samples: QM W7351 (3 sped- the male. mens), AM P25458 (4 specimens), NMV The aboriginal word 'wanda' means 'sand J513 (4). hills'. LEPTANTHURA AND RELATED GENERA 165 Figure 20 — Ulakanthura namoo. Holotype (QMW7348): a, b, c, d, pereopods 1, 2, 4, 7. 166 GARY C. B. POORE Figure 21 — Ulakanthura wanda. Holotype (QM W6124): a, tail fan; b, mandible; c, pereopod 1. Male (QM W6135, QUBS sample) : d, maxilliped; e, pleopod 2. LEPTANTHURA AND RELATED GENERA 167 Figure 22 — Ulakanthura wanda. Holotype (QM W6124): a, b, c, pereopods 2, 4, 7. M 168 GARY C. B. POORE NOMENDUBIUM Paranthura australis Haswell Paranthura australis Haswell 1881: 477-8, pi. 18, iig. 4.— Haswell, 1882: 304-5. Leptanthura australis. — Barnard, 1925: 151. — Nierstrasz, 1941: 242 (part), not Paranthura australis. Miers, 1884: 311. Haswell, 1884: 1012 (= Apanthura cop- pingeri Barnard, 1925). Remarks: Haswell (1881) described and illus- trated an anthuridean from Port Jackson in very general terms. None of the characters de- scribed by Haswell is sufficiently diagnostic to allow the name to be applied to any presently known species. Moreover, two specimens (AM P3315) in the Old Collection of the Australian Museum labelled 'Paranthura australis, Hasw Variety 524 Port Jackson' cannot be reconciled with Haswell's description and figures. Both are males, probably of the genus Paranthura. Haswell did not designate types (Barnard, 1974) although many of the specimens ex- amined by him remain in the Australian Mu- seum collections. It appears that AM P3315 was allocated to P. australis (Number 524 in Haswell's (1882) catalogue) in error at a later date. The name Paranthura australis Haswell is therefore nomen dubium, not certainly applic- able to any known taxon. Discussion Leptanthura, Bullowanthura, Ulakanthura, and possibly the monotypic genus Neoanthura, are separated from the remaining paranthurids by possessing a triangular article 5 on pereo- pods 4-7. They also share loss of eyes, some reduction of the mandibular palp, and con- sistent form of the maxilliped. A review of paranthurid genera in preparation by me shows that the group forms one of the major divisions of the family. It is noteworthy that southeastern Australia is a centre of distribution for this group of genera. Leptanthura now includes about 19 species mostly from bathyal regions. Three species are recorded here from Australia, two from sublittoral environments (less than 200 m). The monotypic genus Bullowanthura is endemic to the region as are all species of Ulakanthura. The latter genus is particularly significant in illustrating once again the high degree of speciation found in marine peracarids from southern Australia. Previously I have described ten species of Haliophasma (Anthuri- dae) from southern Queensland, New South Wales and Victoria (Poore, 1975) and Barnard (1972) has discussed at length adaptive radia- tion in Australian gammaridean amphipods. The total endemicity of this fauna at the species level and endemicity of two of the three genera makes discussion of its biogeo- graphical relationships difficult. The closest relatives of Australian species of Leptanthura seem to be South African species which is un- usual for the southeastern Australian fauna. Better appreciation of the origins of these species may be obtained by greater knowledge of the deep-sea isopods of the region. Acknowledgements I wish to thank the following individuals and institutions for the loan of the material on which this study was based: Diane Brown, Helen Fisher and Jim Lowry from the Austra- lian Museum, Sydney; Ronald Monroe from the Queensland Museum, Brisbane; Stephen Cook from the Department of Zoology, Univer- sity of Queensland; and Margaret Drummond from the Marine Studies Group, Victoria. I gratefully acknowledge the valuable com- ments of Brian Kensley from the South African Museum on the first draft of this paper. This paper is number 167 in the Ministry for Conservation, Victoria, Environmental Studies Series. References Barnard, J. L., 1972. Gammaridean Amphipoda of Australia, Part I. Smithson. Contr. Zool. 103: 1-333. , 1974. Gammaridean Amphipoda of Aus- tralia, Part II. Smithson. Contr. Zool. 139: 1-148. Barnard, K. H., 1925. A revision of the family An- thuridae (Crustacea Isopoda), with remarks on certain morphological peculiarities. /. Linn. Soc t 36: 109-160. LEPTANTHURA AND RELATED GENERA 169 Birstein, Y. A., 1963. 'Deepsea Isopods from the North-western Pacific.' [In Russian], p. 214. Aka- demii Nauk, USSR: Moscow. Haswell, W. A., 1881. On some new Australian marine Isopoda. Part I. Proc. Linn. Soc. N.S.W. 5: 471-481. , 1882. 'Catalogue of the Australian Stalk- and Sessile-eyed Crustacea.' p. xxiv, 326. Aus- tralian Museum: Sydney. 1884. A revision of the Australian Isopoda. Proc. Linn. Soc. N.S.W. 9: 1001-1014. Kensley, B., 1975. Marine Isopoda from the con- tinental shelf of South Africa. Ann. S. Afr. Mus. 67: 35-89- Menzies, R. J., 1962. The isopods of abyssal depths in the Atlantic Ocean. Vema Res. Ser. 1: 79-206. Miers, E. J., 1884. Crustacea, p. 178-322 in 'Report on the Zoological Collections made in the Indo- Pacific Ocean during the Voyage of H.M.S. 'Alert', 1881-2.' Brit. Mus. (Nat. Hist.): Lon- don. Nierstrasz, H. F., 1941. Die Isopoden der Siboga- Expedition. IV. Isopoda Genuina. III. Gnathii- dea, Anthuridea, Valvifera, Asellota, Phreati- coidea. Siboga Exped. 32d: 235-308. Poore, G. C. B., 1975. Australian species of Halio- phasma (Crustacea: Isopoda: Anthuridae). Rec. Aus.Mus. 29: 503-533. Sars, G. O., 1899. 'An Account of the Crustacea of Norway. Vol. II. Isopoda.' p. x, 270. Bergen Museum: Bergen. Stebbing, T. R. R., 1905. On the Isopoda. Rept Govt Ceylon Pearl Oyster Fisheries Gulf Manaar, Part 4, Suppl. Rept 23: 1-64. Wolff, T., 1956. Isopoda from depths exceeding 6000 metres. Galathea Rep. 2: 85-159. THE SPIONIDAE (POLYCHAETA) FROM SOUTHEASTERN AUSTRALIA AND ADJACENT AREAS WITH A REVISION OF THE GENERA 1 By James A. Blake Pacific Marine Station, University of the Pacific, Dillon Beach, California, USA 94929. and Jerry D. Kudenov Ministry for Conservation, Marine Studies Group 605 Flinders Street Extension, Melbourne. Victoria, Australia 3000. Abstract The Spionidae of southeastern Australia are described. Intertidal and subtidal collections come mostly from Victoria, New South Wales and southern Queensland. A few records from West Australia, South Australia, Tasmania and the Great Barrier Reef (Queensland) are also included. A total of 68 species in 19 genera have been identified. These include 4 new genera and 43 new species, with new synonyms and generic emendations also proposed. The genera with pointed prostomia include Scolelepis (9 species, 8 being new), Aonides (1 species) and Dispio (1 new species). Australospio trifida gen. et. sp. nov. is a unique spionid with both a distally pointed prostomium and lateral prostomial horns. The genera with frontal or lateral prostomial horns include Malacoceros (3 species, 2 being new), Rhynchospio (2 new species) and Scolecolepides (1 new species). The definition of Rhynchospio is expanded to include those species with branchiae free from dorsal lamellae. Laonice includes 3 new species, with 1 being a simultaneous hermaphrodite. Spiophanes includes 3 species, Spio with 3 species (2 being new) and Microspio with 1 new species. The genera of the Prionospio-compXex are reviewed and revised to include: Parapriono- spio (1 species), Orthoprionospio gen. nov. (1 new species), Streblospio (not represented) and Prionospio (9 species, 5 being new). A generic revision of the Poly dor a-comp\ex is presented with 6 genera recognized: Boccardia (3 species), Carazziella gen. nov. (4 new species), Polydora (15 species, 8 being new), Boccardiella gen. nov. (2 species, 1 being new), Tripolydora (not represented) and Pseu- dopolydora (5 species, 2 being new). Polydorella is synonymized with Pseudopolydora. Approximately I of the species reported herein are endemic to Australia; the remainder are species with more cosmopolitan distributions. Introduction Despite the voluminous global literature on The Spionidae is one of the largest and most spionids and the large number of species common of the polychaete families encoun- known to occur along some coastlines, there tered in intertidal and shallow subtidal habi- is ver Y little information on Australian spio- tats throughout the world. Species are typically nids - A review of the literature reveals only 4 marine bottom deposit feeders which occur in species which were originally described from all types of substrata, from soft muds to hard Australia. These are Polydorella prolifera Au- calcareous structures. Important systematic gener, 1914 and Spio mesnili Augener, 1914 treatments of the family or of certain genera fr ° m West Australia, Polydora polybranchta are by Mesnil (1896), Soderstrom (1920), Fau- Haswell, 1885 (now Boccardia) from New vel (1927), Hartman (1941), Pettibone (1962; South Wales and Boccardia limmcola Blake 1963), Woodwick (1964), Day (1967), Foster and Woodwick, 1976 from Victoria. In the (1971), Blake (1971), Blake and Woodwick southeastern region of Australia few species (1971-72), Rainer (1973), Read (1975) and have been identified below the generic level. Light (1977- 1978). Haswell (1885) reported Polydora ciliata from commercial oysters in New South Wales, Au- i Contribution No. 178 from the Ministry for gener (1927) reported Spiophanes kroeyeri Conservation, Melbourne, Victoria. from Victoria and Monro (1931) reported Seo- ul 172 JAMES A. BLAKE and JERRY D. KUDENOV lelepis (= Malacoceros) indicus from Queens- land. Several genera have been recorded by Hulchings and Recher (1974) from New South Wales and by Poore, et al. (1975) from Vic- toria, but species names were generally not assigned. There are no published records at all from Tasmania or South Australia. The purpose of the present paper is to des- cribe a new and highly endemic spionid fauna from southeastern Australia. As defined herein, southeastern Australia includes southern Queensland from Moreton Bay and Brisbane, all of New South Wales and Victoria, eastern South Australia and Tasmania. Materials and Methods Specimens have been accumulated from several sources. Extensive Victorian material comes from the Port Phillip Bay and Western- port Bay Environmental Surveys (PPBES, WPBES) performed by the Marine Studies Group (MSG) of the Ministry for Conservation. Location of MSG stations plus additional data are given by Poore, et al. (1975), Poore andKu- denov (1978 a-b) and Coleman, et al. (1978). Additional specimens from the Fisheries and Wildlife Crib Point Benthic Survey (CPBS) in Westernport were also examined. These ma- terials are deposited in the National Museum of Victoria (NMV). Other Victorian materials in- clude NMV collections obtained through Dr Brian Smith, Westernport epifauna collected by Mr Dave Staples and specimens from various lo- calities along the Victorian coast collected by us. Spionidae in the Australian Museum, Sydney (AM) were loaned by Dr P. A. Hutchings. Queensland benthic spionids from Moreton Bay were loaned by Dr W. Stephenson. Some of these specimens are deposited in the Queensland Museum (QM). Spionids from the New South Wales State Fisheries (NSWSF) were provided by Ms M. Skeel from shellfish surveys and through Dr Leon Collett from benthic surveys of Botany Bay. Mr C. Pregenzer (University of New South Wales) loaned specimens he had collected from shellfish. Mr R. Buttermore and M. Skeel provided specimens from Tasmania, which are now lodged in the Tasmanian Mu- seum (TM). Mr W. Zeidler, South Australian Museum (SAM), kindly allowed the second author access to their collections during August 1977. Dr G. Hartmann-Schroder provided Au- gener type materials from the Zoological Mu- seum of Hamburg (ZMH). Dr Marian H. Pettibone loaned comparative specimens from the collections of the National Museum of Natural History, Washington, D.C. (USNM). Since the materials examined in the present study were obtained from various sources, there is little uniformity in mode of collection and preservation. Specimens collected by us or in collections of the MSG were fixed in 10% formalin or Bouin's fluid and preserved in 70% ETOH to which glycerine had been added. Type material has been deposited in the NMV, AM and QM. We are grateful to the administration of the Marine Pollution Studies Group (Ministry for Conservation) for providing facilities and space to the first author during his stay in Melbourne in 1977. We are also grateful to MSG for financial support for this paper. The first author was supported by a sabbatical leave from University of the Pacific and by 2 grants from the National Science Foundation [NSF Grant Int76-11756 from the US-Australia Co- operative Science Programme and NSF Grant GA-30900 from the Biological Oceanography Section]. To these agencies, and to people who loaned or obtained materials on which this study is based, we extend our gratitude. A special debt of gratitude is extended to William J. Light, Oakland Museum, for critically re- viewing the manuscript, sharing his thoughts on many aspects of this study with the first author and by generously allowing us to use some of his unpublished observations. Family Spionidae Grube, 1850 Diagnosis: Prostomium variable: anteriorly rounded or incised, anteriorly expanded to acutely pointed, with or without frontal horns. Occipital tentacle sometimes present on prosto- mial caruncle. A pair of long prehensile peri- stomial palps arising on either side of caruncle near junction of peristomium and setiger 1. Setiger 1 reduced to well-developed, often fused with peristomium. Parapodia biramous, with parapodial lobes conical, cirriform or foli- ose; posterior neuropodia often form low SOUTHEASTERN AUSTRALIAN SPIONTDAE 173 ridges. Setae simple, including capillaries, aci- cular spines, modified setae (major spines of polydorids) and unidentate to multidcntate hooded or non-hooded hooks. Branchiae pre- sent or absent over a variable number of seg- ments. Pygidium reduced to well-developed, with or without cirri, or expanded into disc-like structure or forming several lobes. The following genera and species are covered in this report. 1. Scolelepis lamellicincta sp. nov. 2. S. canmculata sp. nov. 3. S. phyllobranchia sp. nov. 4. S. occipitalis sp. nov. 5. 5. towra sp. nov. 6. S. precirriseta sp. nov. 7. S. sp. 8. S, viridis sp. nov. 9. S. victor iensis sp. nov. 10. Aonides oxycephala (Sars, 1862) 11 . Dispio glabrilamellata sp. nov. 12. Australospio trifida gen. et. sp. nov. 13. Malacoceros indicus (Fauvel, 1928) 14. M. tripartitus sp. nov. 15. M. reductus sp. nov. 16. Rhynchospio glycera sp. nov. 17. R. australiana sp. nov. 18. Scolecolepides aciculatus sp. nov. 1 9. Laonice quadridentata sp. nov. 20. L. hermaphroditica sp. nov. 21 . L. bassensis sp. nov. 22. Paraprionospio pinnata (Ehlers, 1901) 23. Orthoprionospio cirrijormia gen. et nov. 24. Prionospio steenstrupi Malmgren, 1867 25. P. australiensis sp. nov. 26. P. queenslandica sp. nov. 27. P. ehlersi Fauvel, 1928. 28. P. paucipinnulata sp. nov. 29. P. tridentata sp. nov. 30. P. multipinnulata sp. nov. 31. P. aucklandica Augener, 1923 32. P. cirrijera Wiren, 1883 33. Spiophanes bombyx (Claparede, 1870) 34. 5. cf . Kroeyeri Grube, 1 860 35. 5. w/gfeyi Pettibone, 1962 36. Spio mesnili Augener, 1914 37. 5. pacifica sp. nov. 38. S. singularis sp. nov. 39. Microspio granulata sp. nov. sp. 40. Boccardia polybranchia (Haswell, 1885) 41. B. proboscidea Hartman, 1940 42. B. chilensis Blake and Woodwick, 1971 43. Carazziella victor iensis gen. et. sp. nov. 44. C. phillipensis sp. nov. 45. C. hymenohranchiata sp. nov. 46. C. hirsuiiseta sp. nov. 47. Polydora fiava Claparede, 1 870 48. P. socialis (Schmarda, 1861) 49. P. protuberata sp. nov. 50. P. tentaculata sp. nov. 51. P. giardi Mcsnil, 1896 52. P. aciculata sp. nov. 53. P. notialis sp. nov. 54. P. pilocollaris sp. nov. 55. P. armata Langerhans, 1880 56. P. few/ Webster, 1879 57. P. websteri Hartman, 1943 58. P. haswelli sp. nov. 59. P. latispinosa sp. nov. 60. P. woodwicki sp. nov. 61. P. hoplura Gaparcde, 1870 62. Boccardiella bihamata gen. et. sp. nov. 63. B. limnicola (Blake and Woodwick, 1976) comb. nov. 64. Pseudopolydora paucibranchiata (Okuda, 1937) 65. P. kempi (Southern, 1921) 66. P. glandulosa sp. nov. 67. P. stolonifera sp. nov. 68. P. prolifera (Augener, 1914) comb. nov. KEY TO GENERA OF SPIONIDAE (* denotes genera not yet found in Australia) Branchiae absent; setiger 1 with 1-2 large curved neuropodial spines in addition to normal capillariesi Spiophanes (p. 224) Branchiae present; setiger 1 without large neuropodial spines 2 2a. Setiger 5 modified, with specialized setae 18 b. Setiger 5 not modified, without spe- ialized setae 3 3a. Prostomium distally pointed (may appear conical with rounded apex in extremely contracted specimens), with or without subdistal lateral horns 4 la. b. 174 JAMES A. BLAKE and JERRY D. KUDENOV b. Prostomium not distally pointed, with distal lateral or frontal horns, broadly rounded or incised on an- terior margin 7 4a. Branchiae beginning on setiger 1 5 b. Branchiae beginning on setiger 2 6 5a. Prostomium with extra subdistal lateral horns in addition to apical point, appearing 3-parted; setigers 6-9 with dense setal fascicles; noto- setae of setiger 1 similar in length to those of succeeding setigers; ac- cessory branchiae absent Australospio (p. 193) b. Prostomium without extra lateral horns; setigers 6-9 without dense setal fascicles; notosetae of setiger 1 long, thin; accessory branchiae present Dispio (p. 191) 6a. Branchiae fused to dorsal lamellae at least basally, continuing to end of body Scolelepis (p. 175) b. Branchiae completely free from dorsal lamellae, present on variable number of anterior setigers, absent posteriorly Aonides (p. 189) 7a. Prostomium with lateral or fron- tal horns 8 b. Prostomium without lateral or fron- tal horns 10 8a. Branchiae beginning on setiger 1 9 b. Branchiae beginning on setiger 2 Rhynchospio (p. 198) 9a. Hooded hooks in both noto- and neuropodia of posterior segments Scolecolepides (p. 202) b. Hooded hooks only in neuropodia of posterior segments Malacoceros (p. 195) 10a. Branchiae limited to middle and posterior setigers except for a single pair on setiger 2 in males *Pygospio b. Branchiae beginning on setiger 1 or 2 and continuing for a variable number of setigers 11 11a. Branchiae concentrated in 1-22 an- terior setigers, absent posteriorly. . .12 b. Branchiae present over most of body length 15 12a. Branchiae beginning on setiger 1.13 b. Branchiae beginning on setiger 2 Prionospio (p. 211) 13a. One pair of cirriform branchiae; with dorsal collar across setiger 2 * Streblospio b. Three or more pairs of branchiae; no dorsal collar on setiger 2 14 14a. Three pairs of pinnate branchiae; with dorsal ridge across setiger 1 Paraprionospio (p. 209) b. Eighteen to 22 pairs of cirriform branchiae; no dorsal ridge on seti- ger 1 Orthoprionospio (p. 210) 15a. Branchiae beginning on setiger 1 .... 16 b. Branchiae beginning on setiger 2 ... 17 16a. Hooks in both noto- and neuropo- dia *Marenzellaria b. Hooded hooks only in neuropodia Spio (p. 226) 17a. Prostomium broad, bluntly rounded or squared on anterior margin; branchiae free from dorsal lamellae Laonice (p. 204) b. Prostomium narrow, entire or in- cised on anterior margin; branchiae fused to dorsal lamellae Microspio (p. 232) 18a. Branchiae beginning on setiger 2 ... 19 b. Branchiae beginning on setigers 6-12 21 19a. Setiger 5 only slightly modified, with prominent parapodia, major spines of 1 type; hooded hooks tri- dentate, with secondary teeth closely applied to main fang *Tripolydora b. Setiger 5 greatly modified, with re- duced parapodia and major spines of 2 types or 1 type with companion setae; hooded hooks bidentate with prominent angle between teeth 20 1 1 Poly dor a gttillei Laubier and Ramos, 1974 bears unidentate unhooded neuropodial hooks and does not agree with any known polydorid genus. See Remarks p. 234. 20a. Major spines of setiger 5 of 1 type, these simple, falcate, with smaller companion setae. Boccardiella (p. 264) SOUTHEASTERN AUSTRALIAN SPIONIDAE 175 b. Major spines of setiger 5 of 2 types, first with expanded ends bearing cusps or bristles, second simple, fal- cate Boccardia (p. 235) 21a. Setiger 5 slightly to moderately modified, usually with prominent parapodia; major spines of 2 types, first simple, acicular or falcate, second pennoned with both types usually arranged in U- or J-shaped row; hooded hooks with secondary tooth closely applied to main fang Pseudopolydora 2 (p. 267) 2 Pseudopolydora gibbsi Light, 1974 and P. stolonj- fera (this study) deviate somewhat from the generic definition in having more modification to setiger 5 than usually seen in more typical Pseudopolydora spe- cies. The hooded hooks are typical. b. Setiger 5 greatly modified, with re- duced parapodia; major spines of 1 or 2 types in curved row, not J- or U-shaped; hooded hooks with pro- minent angle between teeth 22 22a. Setiger 5 spines of 1 type, variously shaped, with or without companion setae Polydora (p. 245) b. Setiger 5 spines of 2 types, first type with expanded end, second type fal- cate, 1 or both types usually with bristles Carazziella (p. 240) Genus Scolelepis Blainville, 1828 Scolelepis Blainville, 1828. Type-species: Lumbricus squamatus Muller, 1806, by monotypy. Aonis Audouin and Milne-Edwards, 1833. Type- species: Aonis foliosa Audouin and Milne-Edwards, 1833, by monotypy. Nerine Johnston, 1838. Type-species: N. coniocephala Johnston, 1838, selected by Quatrefages 1843 (= Oonis foliosa). Pseudomalacoceros Czerniavsky, 1881. Type-species: Malacoceros longirostris Quatrefages, 1843 (= Lumbricus squamatus), by monotypy. Nerinides Mesnil, 1896. Type-species: Nerine longiro- stris Saint- Joseph, 1894 (= Nerinides cantabra Rio- ja, 1918), by original designation. Scolecolepis Michaelsen, 1897. Erroneous spelling of Scolelepis Blainville, 1828. [Not Malmgren, 18671- Pseudonerine Augener, 1926. Type-species: P. antipo- da Augener, 1926, by monotypy. [Synonymy fide Pettibone, 1963, p. 91]. 1 i Fauchald (1977a) has erroneously cited Nerinides cantabra Rioja, 1918, as the type-species of Pseudo- malacoceros Czerniavsky, 1881, and has replaced Nerinides with Pseudomalacoceros, Pettibone's sy- nonymy (see above) is the correct one and Pseudo- malacoceros is a synonym of Scolelepis (sensu s trie to). Diagnosis; Prostomium pointed on anterior margin, extending posteriorly as narrow car- uncle. Occipital tentacle present or absent. Caruncle posteriorly detached or attached to dorsal body wall. Peristomium well-developed, with or without lateral wings. Branchiae be- ginning on setiger 2, continuing to near pos- terior end of body; in anterior setigers, bran- chiae completely fused to dorsal lamellae or with distal portion free. Capillary noto- and neurosetae of anterior setigers usually ar- ranged in 2 tiers; notosetal capillaries of pos- terior setigers in a single tier. Most capillary setae with prominent sheaths imparting a uni- or bilimbate appearance. Neuropodial hooded hooks present. Notopodial hooded hooks pre- sent or absent. Hooks uni-, bi-, tri- or multi- dentate. Pygidium with oval disc or multi-lobed appendages. Remarks; Scolelepis as defined above is slightly different from that proposed by Pettibone (1963) in that no provision is made for sub- genera. The species are normally grouped into those having a distinct notch in the ventral lamellae (subgenus Scolelepis) and those hav- ing entire ventral lamellae (subgenus Neri- nides). Furthermore, Day (1967) pointed out that Scolelepis normally has both notopodial and neuropodial hooded hooks, whereas Neri- nides has only neuropodial hooded hooks. One or 2 species alternate in these characteristics, but in general they have held up. Light (1977), however, has determined that S. squamata in San Francisco Bay shows a continuous distribu- tion in the presence or absence of the neuro- podial cleft. He also notes that some specimens have notopodial hooks, whereas others do not (Light, 1978). Moreover, one of our spe- cies, 5. viridis (see below) has neuropodial hooded hooks and unhooded notopodial aci- cular spines. It is not clear whether such un- hooded spines, which are clearly of a different type than the hooded ones, have the same taxo- nomic importance as the notopodial hooks of other species. 176 JAMES A. BLAKE and JERRY D. KUDENOV We consider it to be increasingly difficult to 6a. Hooded hooks bidentate (Fig. 3d-e); maintain 2 subgencric or generic (see Fau- parapodia and branchiae of middle chald, 1 977a) categories based on such tenuous and posterior setigers vesiculate (Fig. and variable characteristics. Although we re- 3c) S. phyllobranchia gret lumping such a large group of species b. Hooded hooks multidentate (Fig. under one genus, we believe it to be the better 4h; 5c); parapodia and branchiae not course, pending a re-evaluation of the taxo- vesiculate 7 nomic characteristics of Scolelepis. A similar n n n , t • , , position was adopted by Light (1977). ^ ?™" c > e P^norfy elevated mo fo- The reader is referred to Pettibone (1963) i'°% ^ ^ "JF? n™! and Foster (1971) for historical and revision- S^JS'SSS T A t ary clarification of the genus Scolelepis, and l^l^t £ °> i^l ■ r for lists and keys to the tpecies. Eighteen spe- h %» £? 3t tipS Fl - g ' f C " d) , 5 " °?*** cies are listed as valid by Pettibone (1 963). Wu b ' £™f . PJ^S < m ' and Chen (1964), Hartmann-Schroder (1965), PO^ed not fohose (Fig. 5a-b); su- Foster (1971), Bcllan and Lagardere (1971 P en °r dorsal capillary setae with Gibbs (1971); (1972), and Day (1973) have dc- T^J^K^t T I Spm ° US scribed additional species. In our study, we Jff/ 1 f (Flg ; **** branchiae corn- have identified 9 species belonging to Scdlele- pletely fuS ° d t0 d ° rSal ,araeIIae S ' towra pis, of which 8 are new to science. ^a. Posterior notopodia with acicular spines (Fig. 8g) in addition to capil- KEY TO SPECIES OF SCOLELEPIS FROM lary setae; area posterior to caruncle southeastern Australia with nuchal papilla and raised rec- 1 a. Notosetae present on setiger 1 2 tangular patch (Fig. 7 a-b) S. viridis b. Notosetae absent from setiger 1 5 8b . Posterior notopodia without acicular 2a. Caruncle posteriorly free from dorsum 3 spines, with capillary setae only; car- b. Caruncle attached to dorsum 4 uncle with rounded lobe; nuchal pa- 3a. Setiger 1 with notopodial lamella in P 11 . 1 * located posterior to caruncle presetal position (Fig. 6a-b); hooded ( F '§- 9 a-b) S. victoriensis hooks multidentate, beginning in c ....... . neuropodia of setiger 15-90 (Fig Scolelepis Iamelhcincta sp. nov. 6g-f) S. precirriseta (Figure 1) b. Setiger 1 with notopodial lamella in Material examined: QUEENSLAND. Heron Island, DOStsetal nositinn CFio ?iV hnnApA sandy shore - NE of is1and > mid-low tide levels (1 EvEw ° f ? , f I ' specimen. AM W13000). VICTORIA. Westernport, hooks bl- or tndentate, beginning in Point Leo. coll. Mar. Studies Grp., 18 Jan. 1969 Sta neuropodia of setiger 24-25 (Fig. 57/01, 69/05 (HOLOTYPE, NMV G102- PARA- 2 h-k) S. carunculata TYPK - NMV G2987); Shorham, coll. R. Jensz, Sta An \1™A*A u~ n v -a * . V40 > 3 Dec - 1965 fPARATYPE, NMV G2988); So- 4a. Hooded hooks umdentate, aperture niers , coll. R. Jensz, Sta. V77, Jan. 1966 (PARA- of hood smooth (Fig. ll); occipital TYPE, NMV G2989): Walkerville, coll. R. Jensz, Sta. tentacle absent, but caruncle poster- V\1Q, Jan. 1966 (PARATYPE, NMV G2990). TAS- iorly formed into conical mount MANIA. Macquarie Island, Buckles Bay, dredged, (Fig. la) 5. lamellicincta Jffi», N f, u^f- ,f UT ,V u t t j j t. i -.1 /- • » , aumkalia. fclnston, in sand at base of jetty, coll. b. Hooded hooks with 5 apical teeth, I. M. Thomas, 18 Feb. 1975 (H SAM E1577) aperture of hood serrated (Fig. 6i-j); Description: Ho loty P e 32 mm long and 22 occipital tentaele present S. sp. mm wide across ^^ for ^ ^ ^ 5a. Caruncle posteriorly elevated or free paratypes up to 100 mm long and 8 mm wide from dorsum 6 for 140 setigers. Body widest anteriorly, rect- b. Caruncle posteriorly low, attached to angular in cross section. Salmon coloured in dorsum 8 alcohol. SOUTHEASTERN AUSTRALIAN SPIONIDAE 177 Figure 1 — Scolelepis lamellicincta sp. nov. — a, an- terior end, dorsal view; b, left setiger 11, anterior view; c, left setiger 33, anterior view; d, left setiger 51, anterior view; e, right setiger 120, anterior view; f, pygi- dium, dorsolateral view; g, capillary seta from posterior tier, setiger 10; h, capillary seta from anterior tier, setiger 10; i, uni- dentate hooded hook; j, sabre seta; k, superior capillary notoseta. [Scale 1 = 100 Mm; 2 = 200 Mm; 3 = 50 Mm; 4 = 20 Mm] Prostomium anteriorly projecting, elongate, distally pointed, tapering posteriorly to a blunt and attached, but elevated lobe (Fig. la). Two pairs of eyes, arranged trapezoidally, an- terior pair the larger. Palps extending pos- teriorly to setiger 10, with basal membrane. Peristomium well-developed, forming lateral wings partially overlapping prostomium. Setiger 1 reduced, with leaf-like dorsal la- mellae and rounded ventral lamellae (Fig. la). Dorsal lamellae from setiger 2 distally free from branchiae, flattened, narrow and distally pointed (Fig. lb-e). Postsetal neuropodial la- mellae small and rounded at first, increasing in size and becoming broadly rounded in sub- sequent anterior segments (Fig. 1 b-c) ; at setigers 19-35, dividing into a long interramal lamella and a small, rounded ventral lamella (Fig. ld-e). With slight interramal presetal 178 JAMES A. BLAKE and JERRY D. KUDENOV swelling in setigers 2-15 (Fig. lb); curved sub- podal lobe present in far posterior neuropodia (Fig. le). Notosetae arranged in superior and in- ferior fascicles in anterior setigers and in single fascicles in posterior setigers. All inferior an- terior capillary notosetae and neurosetae with granulations (Fig. lg-h); granulations becom- ing obscured in posterior notopodial capil- laries. Superior fascicle including capillaries with granulated shaft and clear, transparent sheath (Fig. Ik). Anterior capillary neurosetae similar to notosetae. Inferior neuropodial sabre setae from setiger 2-13, numbering 3-6 per fascicle; these distally pointed with finely granulated shaft and prolonged sheaths (Fig. lj). Unidentate hooded hooks in neuropodia from setiger 25-38, and in notopodia from seti- ger 75-80, numbering 5-6 per ramus. Hooks distally falcate, blunt, aperture of hood form- ing smooth circle (Fig. li). Branchiae fused to notopodial lamellae, but distally free (Fig. lb-e). Pygidium with in- cised ventral cushion (Fig. If). Anus dorsal, surrounded by crenulate margin. Remarks: Scolelepis lamellicincta is most closely related to S. bonneri (Mesnil) from France and to S. joliosa (Audouin Milne- Edwards) of cosmopolitan distribution; it dif- fers from those species and all others by the structure of the parapodia, especially by the long interramal lamella of the middle and pos- terior segments. The conical caruncle is also unique. Distribution: Queensland; Victoria; Tasmania, Macquarie Island; South Australia. Scolelepis carunculata sp. nov. (Figure 2) Material examined: QUEENSLAND. Moreton Bay, Jackson Creek, mangroves and sand flats, coll. P. Hutchings and C. Wallace, 12 July 1973 (1, AM W6057). NEW SOUTH WALES. Belmont Beach, HDWBS sample, Jan. 1976 (1, AM W8926). VIC- TORIA. Port Phillip Bay, Werribee MSG Sta. 1871, coll. G. Poore and J. D. Kudenov, Nov. 1975 (1, NMV G2991); Westernport, Shorham, Sta. V47, coll. R. Jensz, 20 Dec. 1965 (HOLOTYPE, NMV G2992; 5 PARATYPES NMV G2993); Port Fairy, Sta. V51, coll. R. Jensz, 10 Jan. 1966 (1, NMV G2994). Port- land, Sta. V62, coll. R. Jensz, 8 Jan. 1966 (1, NMV G2995); Somers, Sta. V77, coll. R. Jensz, Jan. 1966 (2, NMV G2996). WEST AUSTRALIA. Safety Bay, south of Perth, coll. 15 Sept. 1968 (2, NMV G2997). Description: A large species, measuring up to 48 mm long and 3 mm wide for 110 setigers. Body widest anteriorly, tapering gradually pos- teriorly; elliptical in cross section. Colour in alcohol: opaque white to dusky brown. Prostomium slightly fusiform, pointed an- teriorly and posteriorly, posterior lobe free un- attached, extending to anterior margin of seti- ger 2 (Fig. 2a-b). Two pairs of eyes, arranged trapezoidal Iy, the anterior pair slightly larger. Palps extending posteriorly to setiger 8-9. Pe- ristomium well-developed, forming lateral wings, but not overlapping prostomium. Setiger 1 reduced (Fig. 2a), with elliptical postsctal lamellae, dorsal lobes larger than ventral ones. Notopodial lamellae distally free from branchiae, entire, but greatly folded (Fig. 2c). Neuropodial lamellae in anterior setigers elliptical (Fig. 2c), becoming bilobed by setiger 30-35 (Fig. 2d). Parapodial lamellae of pos- terior setigers overlapping interramally (Fig. 2d)- Anterior setal tiers bear granulated capil- laries with transparent sheaths (Fig. 2f-g); su- perior capillaries of posterior notopodial tiers and inferior capillaries of posterior neuropo- dial tiers with smooth shafts and sheaths bear- ing large vesicles, appearing golden in reflected light (Fig. 2m); remaining capillaries of pos- terior tiers non-granulated; superior dorsal capillaries distally granulated, with thin, pro- longed distal process (Fig. 21). Two to 3 in- ferior sabre setae in neuropodia from setiger 3. Hooded hooks beginning in setiger 24-45 in neuropodia and in setiger 70-80 in notopodia. Figure 2 — Scolelepis carunculata sp. nov. — a, an- terior end, dorsal view; b, anterior end, lateral view; c, right setiger 10, anterior view; d, right setiger 30, anterior view; e, pygidium; f, anterior capillary seta; g, detail of shaft of f; h, neuropodial biden- tate hooded hook, frontal view; i, same, lateral view; j, notopodial tridentate hooded hook, lateral view; k, notopodial tridentate hooded hook, frontal view; 1, inferior capillary neuroseta; m, superior capillary notoseta. [Scale 1 = 500 Mm; 2 = 500 Mm; 3 = 20 Mm; 4 = 50 Mm; 5 = 20 Mm; 6 = 20 Mm] SOUTHEASTERN AUSTRALIAN SPIONIDAE 179 180 JAMES A. BLAKE and JERRY D. KUDENOV Hooks bidentate in neuropodia (Fig. 2h-i) and bi- and tridentate in notopodia (Fig. 2j-k); hooks with primary and secondary hoods, pri- mary hood bearing minute granules and sec- ondary hood clear. Branchiae stout, triangular, partially fused to dorsal lamellae. Pygidium with large ven- tral, entire cushion; anus surrounded by paired lateral and single dorsal lobes (Fig. 2e). Remarks: Scolelepis carunculata is most closely related to S. squamata (Miiller) in hav- ing notosetae on setiger 1 and in exhibiting bi- dentate hooded hooks. S. carunculata differs in the gills being fused to the dorsal lamellae for most of their length, the posterior notopo- dia are not incised, the dorsal and ventral la- mellae of posterior setigers overlapping inter- ramally and in the pygidium having dorsal and lateral lobes in addition to the ventral cushion. S. carunculata further differs from all pre- viously described species of the genus in hav- ing hooded hooks with a secondary hood. Distribution: Queensland; New South Wales; Victoria; West Australia. Scolelepis phyllobranchia sp. nov. (Figure 3) Material examined: QUEENSLAND. Moreton Bay, Middle Banks, Sta. 4B coll. W. Stephenson, Jan. 1976 (HOLOTYPE, NMV G2983; PARATYPE, NMV G2984); Sta. 48, Dec. 1972 (PARATYPE, NMV G2985); Sta. 4C, March 1976 (PARATYPE, QM G11586); Sta. 53, Sept. 1973 (PARATYPE , QM G11587). Description: All specimens are incomplete. Holotype the largest specimen, measuring 15 mm long and 5 mm wide for 27 setigers. An- terior end dorsoventrally flattened, wide, rec- tangular in cross section. Colour in alcohol: opaque white. Prostomium anteriorly elongate, distally pointed, continuing posteriorly as elevated mound with small apical nuchal papilla (Fig. 3a). Two pairs of subequal eyes, arranged rectangularly. Palps missing. Peristomium moderately developed, forming low lateral wings, partially encompassing prostomium. Setiger 1 reduced, bearing small rounded notopodial lamellae, lacking notosetae; sub- quadrate neuropodial lamellae (Fig. 3a). Noto- podial lamellae totally fused to branchiae from setiger 2 to about 18, thereafter notopo- dial lamellae from about setiger 20-25 dis- placed distally on elongated stalk (Fig. 3c), di- verging to form 2 leaf-shaped, vesiculate pro- cesses: a dorsal lamella and separate branchia. Anterior neuropodial lamellae rounded (Fig. 3b), becoming bilobed from about setiger 25, with vesiculate petaloid interramal lamella (Fig. 3c). Capillaries of anterior setal tiers mostly granulated, those of posterior row, and an- terior superior fascicle of 4-5 capillaries, non- granulated, sheathed, with sheath being dis- tally prolonged in superior capillaries (Fig. 3 f-g). Inferior sabre setae absent. Bidentate hooded hooks in neuropodia from setiger 23 (Fig. 3d-e); notopodial hooks lacking in frag- mentary material. Nature of pygidium unknown. Remarks: Scolelepis phyllobranchia is unique among spionids in having a stalk or trunk on posterior segments from which branch separate notopodial lamellae and gill processes. These parts as well as the neuropodial interramal la- mellae are vesiculated and resemble the dorsal cirri of phyllodocids. The fragmentary nature of the types precludes an assessment of the presence of notopodial hooks. Distribution: Queensland, Moreton Bay. Scolelepis occipitalis sp. nov. (Figure 4) Material examined: NEW SOUTH WALES. Bur- wood Beach, HDWBS samples, June 1975 (HOLO- TYPE, AM W8910 plus 8 PARATYPES from 8 stations, April, May, July, Sept. 1975, AM W8909, 8911-8917); Belmont Beach, HDWBS samples, 9 stations, June, July, Sept., Nov., Dec. 1975 and Feb. 1976 (9 PARATYPES, AM W8918-8925, 8928). Description: A moderate-sized species, up to 16 mm long and 3*0 mm wide for 68 setigers. Body widest anteriorly, rectangular in cross section. Colour in alcohol: opaque white. Prostomium anteriorly inflated with a distal conical protuberance (Fig. 4 a); posteriorly pro- duced into a flattened, irregularly-shaped, partly branched occipital process (Fig. 4a-b). Eyes absent. Palps missing. Peristomium mod- SOUTHEASTERN AUSTRALIAN SPIONIDAE 181 N0T0P0DIAL POSTSETAL LAMELLA a i c d l r I 4 e f g GILL INTERRAMAL Figure 3 — Scolelepis phyllobranchia sp. nov. — a, an- terior end, anterodorsal view; b, right setiger 10, anterior view; c, right setiger 25, anterior view; d, bidentate hooded hook, frontal view; e, bidentate hooded hook, lateral view; f, superior capillary notoseta; g, detail of same. [Scale 1 = 100 Mm; 2 = 20 /*m; 3 = 100 ^m; 4 = 10 /on] erately developed, with small lateral wings (Fig. 4a). Setiger 1 reduced, with small digitiform ase- tigerous notopodial lobes; neuropodial lamel- lae rounded, bearing capillary setae (Fig. 4 a-b). Anterior notopodial lamellae partly fused with branchiae from setiger 2, but distally free; lamellae elongate with rounded tips in ante- rior segments (Fig. 4c), becoming wing-shaped and distally pointed in posterior segments (Fig. 4d-e). Branchiae robust, wide and short at first, 182 JAMES A. BLAKE and JERRY D. KUDENOV Figure 4 — Scolelepis occipitalis sp. nov. — a, anter- ior end ,frontal view; b, anterior end, dor- sal view; c, right setiger 11, anterior view; d, right setiger 35, anterior view; e, right setiger 58, anterior view; f, pygidium, ter- minal view;g,quinquedentate hooded hook, lateral view; h, detail of quinquedentate hooded hook, hood omitted [not to scale]; i, quinquedentate hooded hook, frontal view; j, superior capillary notoseta; k, de- tail of shaft of same. [Scale 1 = 100 Mm; 2 =s 20™; 3 = 10 Mm] increasing in size over anterior region, decreas- ing in length posteriorly. Anterior neuropodial lamellae entire and rounded; lamellae divided in middle and posterior segments, forming a small ventral lobe and large interramal lamella (Fig. 4c-e). Dorsal and interramal lamellae of middle and posterior segments overlapping (Fig. 4d-e). Inferior presetal neuropodial lobe from setiger 19-20 (Fig. 4d-e). Capillary setae granulated in anterior tier, smooth in posterior tier; superior group of 2-5 capillaries with smooth shaft and tubercu- lated sheath (Fig. 4j-k). Neuropodial quinque- dentate hooded hooks from setiger 24-25 (Fig. 4g-i), notopodial hooks beginning about seti- ger 52. Hooks bearing 4 small apical teeth in 2 rows with a V-shaped medial gap surmount- ing large main fang (Fig. 4h-i); quinquedentate hooks appearing tridentate in certain views (Fig.4g-i); inferior sabre setae absent. Pygidium with ventral cushion, with 1 achae- SOUTHEASTERN AUSTRALIAN SPIONIDAE 183 tous preanal segment. Anus with deeply scal- loped margin (Fig. 4f). Remarks: Scolelepis occipitalis is readily re- cognized by the flattened, partly branched occi- pital lobe and the configuration of the hooded hooks, which more closely resemble those seen in the genus Prionospio than in other species of Scolelepis. Distribution: New South Wales. Scolelepis towra sp. nov. (Figure 5) Material examined: NEW SOUTH WALES. Botany Bay, Towra Beach, NSWSF Sta. 226, associated with Posidonia (HOLOTYPE, NMV G2982). Description: Holotype an incomplete anterior fragment, measuring 7 mm long and 0-8 mm wide for 28 setigers. Specimen a gravid female with hard membraned primary oocytes. Prostomium distally pointed, widest sub- terminally, with caruncle tapering posteriorly to form erect conical lobe extending to anterior margin of setiger 3, with slight constriction at level of eyes. Two pairs of eyes arranged tra- pezoidally, anterior pair the larger (Fig. 5a-b). Palps missing. Peristomium moderately en- larged, without lateral wings. Notopodial lobe of setiger 1 reduced, lack- ing notosetae; neuropodium rounded, with capillary setae. Notopodial lamellae completely fused with branchiae from setiger 2, rounded at first (Fig. 5c) then becoming triangular (Fig. 5d) in subsequent segments. Neuropodial la- mellae auricular and continuous with notopo- dial lamellae in anterior setigers (Fig. 5c), de- creasing in size posteriorly and shifting dor- sally to interramal position, becoming asym- metrical and overlapping notopodial lamellae (Fig. 5d). Low rounded presetal neuropodial lobe from setiger 15-18 (Fig. 5d). Anterior capillary notosetae and superior group of 3-5 longer capillaries with striated shaft and minutely spinous sheath (Fig. 5g-h), distally prolonged as fine tip. Smooth sabre setae in neuropodia only in setigers 10-14. Multidentate hooded hooks begin setiger 14- 15, secondary teeth arranged in 2 pairs, closely applied to elongated main fang (Fig. 5e-f). No- topodial hooks unknown. Nature of pygidiurn unknown. N Remarks: Scolelepis towra is closely related to S. texana Foster, 197 1 , from the Gulf of Mexi- co and S. occipitalis (this study), all of which have multidentate hooded hooks. The hooks of 5. occipitalis differ from those of 5. towra in exhibiting a large, medial, V-shaped gap be- tween the pairs of apical teeth; no such gap oc- curs in the hooks of S. towra, S. towra and S. texana have quite similar hooks, but the 2 spe- cies can be separated as follows: in 5. towra, the branchiae are totally fused to the dorsal lamellae and there is no occipital cirrus (al- though the posterior edge of the caruncle forms an erect lobe); in 5. texana branchiae are only fused basally, and an erect occipital an- tenna is present. In addition, the anterior dor- sal capillaries of S. towra bear striated shafts with finely spinous sheaths; those of S. texana exhibit punctate shafts with clear, entire, hya- line sheaths, similar to those seen in 5. squa- mata (W. J. Light, personal communication). Distribution: New South Wales, Botany Bay, Towra Beach. Scolelepis precirriseta sp. nov. (Figure 6a-h) Material examined: QUEENSLAND. Deception Bay, Burpengary Creek (1, AM W7129); Brisbane, Fine Station, Feb. 1972 (9, AM W7483): 14-2S m, May 1974 (HOLOTYPE, AM W13052; 2 PARA- TYPES, AM W13047). NEW SOUTH WALES. Wal- lis Lake, Zostera beds, coll. Dixon and O'Gower, Dec. 1970 (2, AM W5007). Description: A moderate-sized species, up to 17 mm long and 1-5 mm wide for 39 seti- gers. Body dorsoventrally flattened anteriorly, setigers 1-18 elliptical in cross section, sub- quadrate posteriorly. Collection includes males with sperm and females with hard membraned oocytes. Colour in alcohol; opaque white. Prostomium anteriorly pointed or blunt, de- pending on amount of contraction during pre- servation. Posterior aspect of prostomium pro- duced into elevated free lobe extending pos- teriorly to setiger 2 (Fig. 6a). Constriction pre- sent at level of eyes. Two pairs of eyes, trape- zoidally arranged, anterior pair the larger. Peristomium moderately developed, but with- out lateral wings. Regenerating palps inserted posterolaterally to prostomium in holotype 184 JAMES A. BLAKE and JERRY D. KUDENOV T 2 a b g h Figure 5 — Scolelepis towra sp. nov. — a, anterior end, dorsal view; b, anterior end, frontal view; c, right setiger 10, anterior view; d, right setiger 20, anterior view; e, quinque- dentate hooded hook, lateral view; f, quin- quedentate hooded hook, without hood showing 2 broken teeth and 2 entire teeth behind, lateral view; g, superior capillary notoseta; h, detail of sheath of same. [Scale 1 = 100 Mm; 2 = 500 Mm; 3 — 10 Mm; 4 = 20 Mm; 5 = 10 Mm] (Fig. 6a). Setiger 1 reduced, but with notosetae; notopodial lamellae fingerlike and presetal instead of postsetal in location; neuropodial lamellae rounded and postsetal (Fig. 6a-b). Dorsal lamellae basally fused with branchiae in anterior setigers, becoming free posteriorly. Lamellae of anterior setigers distally pointed (Fig. 6c), becoming hatchet-shaped in pos- terior setigers (Fig. 6d-e). Neuropodial lamel- lae continuous with notopodial lamellae in anterior setigers, the 2 being separated by a shallow notch (Fig. 6c); neuropodia becoming broadly rounded at about setiger 29, then mov- ing to interramal position (Fig. 6d-e). Anterior dorsal capillaries non-granulated or granulated; 4-5 superior dorsal capillaries with non-granulated shaft and tubercles on sheath (Fig. 6h). Hooded hooks first present SOUTHEASTERN AUSTRALIAN SPIONIDAE 185 Figure 6 — Scolelepis precirriseta sp. nov. — a, anterior end, dorsal view; b, anterior end, lateral view; c, right setiger 10, anterior view; d, right setiger 29, anterior view; e, right setiger 39, anterior view; f, multidentate hooded hook, lateral view; g, same, en- larged, hood omitted [not to scale]; h, superior capillary notoseta — Scolelepis sp. — i, hexadentate hooded hook, lateral view; j, hexadentate hooded hook, without hood showing arrangment of teeth, frontolateral view. [Scale 1 = 100 Mm; 2 = 300 Mm; 3 = 20 Mm] 186 JAMES A. BLAKE and JERRY D. KUDENOV in neuropodia from setiger 15-20, mostly re- placing capillaries. Hooks with relatively small primary fang and 3 pairs of small apical teeth (Fig. 6f-g), suggestive of a Prionospio setal type, appearing quadridentatc in lateral view. Notopodial hooks unknown. Sabre setae un- known. Nature of pygidium unknown. Remarks: Scolelepis precirriseta differs from all known species of the genus in having the dorsal lamella of setiger 1 in the presetal in- stead of postsetal position. The outer superior dorsal capillaries have minute tubercles on the sheath. Distribution: Queensland; New South Wales. Scolelepis sp. (Figure 6i-j) Material examined: VICTORIA. Westernport, Crib Point, CPBS Sta. 32N (1, NMV G2894; 1, NMV G2895). Remarks: The specimens are incomplete and measure up to 25 mm long and 2-5 mm wide for 44 setigers. They are poorly preserved. An occipital tentacle is present. There are no eyes. Setiger 1 bears notosetae. The branchiae are completely fused to the dorsal lamellae in at least in anterior segments. The dorsal lamellae appear highly glandular in anterior segments and are reddish in colour. Anterior fascicles have non-granulated capillary setae with sheaths. Multidentate hooded hooks begin on setiger 11. The margin of the hood aperture is serrated (Fig. 6i). Apical teeth are arranged in 2 pairs above the main fang, and underlie a small unpaired tooth (Fig. 6i-j). Sperm were observed in one specimen (NMV G-2894) and proved to be of the primitive type (Franzen, 1956). Specific identification cannot be made from the present, badly preserved specimens. Distribution: Victoria, Westernport Bay, in muddy sand. Scolelepis viridis sp. nov. (Figures 7-8) Material examined: QUEENSLAND. Great Bar- rier Reef, Opal Reef, coll. 9 Nov. 1976, subtidal, 2 m in coral sand under boulder on reef flats, coll. J. H. Dorsey (HOLOTYPE, NMV G2914). Description: Holotype complete, measuring 200 mm long and 9 mm wide for about 200 setigers. Body widest anteriorly, gradually ta- pering posteriorly, generally rectangular in cross section. Posterior segments each with raised glandular ridges surrounding body. Specimen a gravid female with flattened, hard membraned oocytes. Colour in life: bright green; in alcohol; dark grey. Prostomium distally pointed, widest sub- terminally, gradually tapering posteriorly (Fig. 7a-b). Flattened nuchal papilla present pos- terior to and separated from caruncle. A raised rectangular area occurring just posterior to pa- pilla, on setiger 3 (Fig. 7a-b). Palps missing. Two pairs of eyes in trapezoidal arrangement. Peristomium well-developed, forming lateral thickening about prostomium. Setiger 1 reduced, but with small rounded notopodial lamellae, lacking notosetae; neuro- podial lamella auricular, much larger than no- topodial lamella (Fig. 7a). From setiger 2 noto- podial lamellae marginally rounded, completely fused with branchiae, both elements forming strap-like structure (Fig. 8a-b); lamellae highly reduced in middle and posterior body region, where branchiae become separated and smaller (Fig. 8c-d), with notopodia elongating into cirriform lobes, barely fused basally to bran- chiae. Branchiae reduced to ciliated "mounds in far posterior setigers (Fig. 8e). Anterior neuro- podial lamellae entire (Fig. 8a); with inferior notch in middle setigers (Fig. 8b), becoming narrowly rounded (Fig. 8c-e) in posterior seti- gers, with small presetal lobe (Fig. 8d-e). Setal fascicles arranged as 5 tiers of capil- lary setae per ramus in anterior setigers, de- creasing to 2 tiers by setiger 40. From about setiger 50, notopodia each with superior and inferior tuft of capillaries (Fig. 8c). Anterior capillary notosetae of 2 types: (1) those of tiers 1-2 with granulated shaft and clear sheath (Fig. 7c-d); (2) those of tiers 3-5 and superior fas- cicles with non-granulated shaft and promin- ently bristled sheath (Fig. 7e-f). With simple, heavy acicular spines in notopodia of middle and posterior setigers (Fig. 8g) from setiger 50. Neurosetae including sheathed granulated and non-granulated capillaries in addition to bi- dentate hooded hooks. Two inferior sabre setae from setiger 2, number of sabre setae increas- ing to 20 per fascicle in middle segments; SOUTHEASTERN AUSTRALIAN SPIONIDAE 187 ^ L A a 2 C d 5 b g e f h _ A Figure 7 — Scolelepis viridis sp. nov. — a, anterior end, dorsal view; b, prostomium and peristo- mium, lateral view; c, superior capillary notoseta; d, detail of same; e, capillary notoseta from tier 3; f, detail of same; g, inferior neuropodial sabre seta; h, detail of same. [Scale 1 = 200 Mm; 2 = 100 Mm; 3 = 100 Mm; 4 = 20 Mm; 5 = 20 Mm] 188 JAMES A. BLAKE and JERRY D. KUDENOV Figure 8 — Scolelepis viridis sp. nov. — a, right seti- ger 10, anterior view; b, right setiger 38, anterior view; c, right setiger 100, anterior view; d, right setiger 150, anterior view; e, right setiger 190, anterior view; f, py- gidium, terminal view; g, notosetal spine; h, bidentate hooded hook, lateral view; i, same, frontal view. [Scale 1 = 200 Mm; 2 = 50 Mm] these with non-granulated shafts and minute bristles on sheath (Fig. 7g-h). Bidentate hooded hooks from setiger 28-29 in neuro- podia, secondary tooth forming angle of about 60° with main fang (Fig. 8h-i). Pygidium with 2 papillae: the first small, dorsomedial, the second ventromedial, conical and larger. Pygidium surrounded by thick glandular rings (Fig. 8f). Remarks; Scolelepis viridus is a unique species, characterized by having a nuchal papilla and peculiar raised nuchal area posterior to the caruncle; glandular ridges on posterior body segments; reduction of dorsal lamellae and cor- responding elongation of the notopodia in mid- dle and posterior segments; the presence of un- hooded acicular spines in notopodia; biden- tate neuropodial hooded hooks; 5 tiers of an- terior capillary setae; and a peculiar pygidial structure with dorsal and ventral papillae. SOUTHEASTERN AUSTRALIAN SPIONIDAE 189 Distribution: Queensland, Great Barrier Reef, with coral. Scolelepis victoriensis sp. nov. (Figure 9) Material examined: VICTORIA. Westernport, WPBES Stations 1718-1 (PARATYPE, NMV G2906); 1723-2 (PARATYPE, NMV G2907); 1734-2 (PARA- TYPE, NMV G2908); 1736-1 (PARATYPE, NMV G2909); 1737-1 (PARATYPE, NMV G2910); 1738-3 (PARATYPE, NMV G2911); 1740-1 (HOLOTYPE, NMV G2912); Crib Point, CPBS Station 300, Dec. 1970 (PARATYPE, NMV G2913). Description: All specimens incomplete anterior fragments measuring up to 15 mm long and 4 mm wide for 31 setigers. Body widest an- teriorly, elliptical in cross section. Colour in alcohol; opaque white. Prostomium distally pointed; posteriorly with rounded lobe just behind and above the 2 pairs of eyes. Anterior pair of eyes the larger. With nuchal papilla posterior to caruncle on setiger 2 (Fig. 9a). Palps missing. Peristomium well-developed, with lateral wings. Proboscis an eversible sac. Setiger 1 reduced with small, petaloid noto- podial lamellae, lacking notosetae; neuropodial about as lopg as notopodial lamellae (Fig. 9 a-b). Notopodial lamellae marginally rounded and straplike on all anterior setigers, entirely fused to branchiae. Neuropodial lamellae sub- quadrate in anterior setigers (Fig. 9c), low and broadly rounded in middle body segments (Fig. 9d). Anterior notopodia with well-developed presetal lobes (Fig. 9c-d). All notopodia bearing capillaries with granu- lations on shaft and broadly sheathed with minute hair-like fimbriations (Fig. 9e); sheath more closely applied in superiormost capil- laries, with fine bristles only apparent under oil immersion (Fig. 9g-h). Neuropodial capil- laries similar to those of notopodia. Inferior sabre setae beginning on setiger 2-3 with 2-3 per tuft at first, subsequently increasing to 4-5. Bidentate neuropodial hooded hooks from setiger 25, these hooks bearing a single distally pointed, rather long apical tooth closely ap- plied to main fang (Fig. 9f). Margin of hood aperture not serrated. Dorsal hooks lacking. Remarks: Scololepis victoriensis is a unique species in the nature of the overlapping apical tooth of the hooded hooks. The presence of a prostomial swelling and separate nuchal pa- pilla are also unusual. Distribution: Victoria, Westernport. Genus Aonides Claparede, 1864 Aonides Claparede, 1864. Type-species: A. auricu- laris Claparede, 1864 [= A. oxycephala (Sars, 1862)], by monotypy. Paranerine Czerniavsky, 1881. Type-species: Nerine oxycephala Sars, 1862, by monotypy. Diagnosis: Prostomium conical, tapered both anteriorly and posteriorly. Peristomium poorly developed. Branchiae cirriform, separated from dorsal lamellae, beginning on setiger 2 and present on a variable number of anterior setigers; absent posteriorly. Bi- or tridentate hooded hooks present in both notopodia and neuropodia. Pygidium with anal cirri. Remarks: A small genus, represented in our collections by only 1 species, A. oxycephaly an apparently cosmopolitan species (see Ra- mos, 1976). Aonides oxychephala (Sars, 1962) Nerine oxycephala Sars, 1862, p. 64. Aonides oxycephala: Ramos, 1976, p. 11 (Synonymy). Aonides californiensis Rioja, 1947, p. 205. Fide Ra- mos, 1976. Aonides mayaguezensis Foster, 1969, p. 393; 1971, p. 66. Fide Ramos, 1976. Material examined: NEW SOUTH WALES. Mer- imbula, Sta. MER 287N, Spencer Park Transect, sand, 5 Oct. 1975, coll. J. H. Day, et al. (1, AM W11735); Sta. MER 289V, channel core N. side cen- tral sand bank, Halophila, 4 Dec. 1975 (1, AM W11746); Sta. MER 297W, channel core opposite Spencer Park (1, AM W11747). VICTORIA. Port Phillip Bay, PPBES Station 985/2 (1, NMV G2915). Remarks: These specimens agree well with the description by Ramos (1976). There are at least 20 pairs of branchiae, and bidentate hooded hooks begin in the notopodia from about setiger 29 and in the neuropodia from about setiger 40. The usefulness of branchial distributions in the delineation of Aonides spe- cies needs careful consideration, particularly when dealing with what may be a widespread species. Similarly, the first occurrence of noto- and neuropodial hooks may not be taxono- mically important in this genus. An evaluation of this feature requires a more thorough un- derstanding of the larval development (see Hannerz, 1956, pp. 26-32). 190 JAMES A. BLAKE and JERRY D. KUDENOV Figure 9-ScoUlepis vgonenw sp. nov.-a, anterior end, lateral view; b, anterior end, dorsal view c, right setiger 8 anterior view; d, right setiger 21, anterior view; e, norma cap. ary notoseta, distal process; f, bidentate hooded hook, lateral view g superior cap.Hary notoseta; h, shaft, detail of same. [Scale 1 = 100 «n 2 = 30 7m 3 SOUTHEASTERN AUSTRALIAN SPIONIDAE 191 Distribution: New South Wales, Merimbula, associated with sandy channel sediments and Posidonia; Victoria, Port Phillip Bay, 9 m in sandy sediments. Genus Dispio Hartman, 1951 Type-species; D t uncinata Hartman, 1951, by mono- typy. Diagnosis: Prostomium fusiform, anteriorly pointed, with narrow caruncle extending pos- teriorly. Eyes present or absent. Peristomium moderately developed, forming low lateral wings. Anterior parapodial lamellae lobed or entire. Presetal notopodial and neuropodial lobes present or absent. Branchiae present from setiger 1, fused to notopodial lamellae for half or more of their length. With accessory branchiae on posterior face of notopodia in middle and posterior segments. With capillary notosetae only. Neurosetae include capillaries, hooded hooks and sabre setae. Pygidium with midventral flap and prominent anal cirri. Dispio glabrilamellata sp. nov. (Figure 10) Material examined: QUEENSLAND. Moreton Bay, Middle Banks, Sta. 3, Dec. 1973 (1, QM G11588); 6D, March 1976 (QM G11589), coll. W. Stephenson. NEW SOUTH WALES. Sydney, S. end Manly Beach, 7-2-8-7 m, coll. 30 June 1971 coll. P. A. Hutchings (1, AM W13001); Burwood Beach, HDWBS Stations, March, May, July, Nov. 1975 (4 PARATYPES, AM W8890-8893); McMasters Beach, HDWBS Stations, July, Nov. 1975 (8 PARATYPES, AM W8894-8896, 8946; HOLOTYPE AM W13048); Belmont Beach, HDWBS Station, Jan. 1976 (2 PARATYPES, AM W8897, 8833); Dudley Beach, HDWBS Station, July 1975 (PARATYPE, AM W8898). VICTORIA. West- ernport, West Head, Flinders Rock Platform, coll. R. Jensz, 16 Jan. 1968 (2 PARATYPES, NMV G2952). Description: All specimens incomplete. Lar- gest fragments up to 20 mm long and 3 mm wide for 110 setigers. Body widest anteriorly, tapering gradually posteriorly; trapezoidal to rectangular in cross section. Colour in alcohol: opaque white. Prostomium elongate, sharply pointed an- teriorly (Fig. 10a). Caruncle short, narrow, reaching only to posterior margin of setiger 1. Occipital tentacle absent, but prominent boss present behind eyes (Fig. 10b). Two pairs of subdermal eyes, often obscured, arranged tra- pezoidally. Palps missing. Peristomium well- developed, forming distinct lateral wings par- tially encompassing prostomium (Fig. lOa-b). Setiger 1 slightly reduced with notopodium shifted dorsally and bearing 4-5 long noto- setae; neurosetae much shorter than notosetae. Notopodial lamellae of setiger 2 and suc- ceeding setigers entire, long and narrow, fused with branchiae; lamellae distally free with pointed tips and entire margins (Fig. lOc-e). Neuropodial lamellae rounded to subrectangu- lar in anterior setigers (Fig. 10c), with pointed superior and inferior processes in posterior setigers (Fig. lOd-e). Presetal notopodial lobes from about setiger 30 as thickened margins (Fig. lOd), becoming quite elongate, subulate and maximally developed from around setiger 80 (Fig. lOe), decreasing in size thereafter. Pre- setal neuropodial lobes from about setiger 60, becoming broadly triangular from around seti- ger 80 (Fig. lOe). Setal fascicles arranged in 2 tiers in each ra- mus. Notosetae sheathed and with or without transverse partitions in the shaft (Fig. 10f). An- terior setigers with 30-35 capillaries and 3-4 longer superior setae in each notopodium, these gradually reduced to about 10 normal and 3 longer superior setae in posterior seti- gers. Capillaries of setiger 1, superior noto- setae and those in posterior tiers of both rami without transverse partitions; anterior tiers in each notopodium with transversely barred ca- pillaries, these partitions tending to disappear in posterior setigers. Neurosetal capillaries similar to notosetal. Neuropodial sabre setae present from setiger 9-11, these lacking par- titions initially, but becoming thicker and transversely barred from around setiger 25. Sabre setae numbering 2-5 per fascicle. With unidentate hooded hooks in neuropodia from setiger 22-26 (Fig. lOg); 1 juvenile specimen (AM W8891) with bidentate and unidentate hooded hooks intergrading in the same fascicle (Fig. lOh). Branchiae from setiger 1, smooth, not ser- rated, fused with dorsal lamellae. Accessory branchiae on posterior face of notopodia from about setiger 23, continuing to end of body; each with up to 4-6 digitate processes (Fig. 10 d-e). Nature of pygidium unknown. 192 JAMES A. BLAKE and JERRY D. KUDENOV Figure 10 — Dispio glabrilamellata sp. nov. — a, anterior end, dorsal view; b, anterior end, lateral view; c, left setiger 9, anterior view; d, left setiger 35, anterior view [broken lines indicate location of accessory branchiae on posterior side]; e, left setiger 80, anterior view [same with accessory branchiae]; f, anterior capillary notoseta; g, unidentate hooded hook; h, neuropodial fascicle of hooded hooks from juvenile showing bi- dentate (unworn) grading into unidentate (worn) hooded hooks [insets amplify details, not to scale]. [Scale 1 — 100 wn; 2 = 50 /^m] SOUTHEASTERN AUSTRALIAN SPIONIDAE 193 Remarks: Dispio was reviewed by Pettibone (1963) and Foster (1971). Four species were known at that time: D. uncinata Hartman, 1951 from the Gulf of Mexico; D. magna (Day, 1955) from South Africa; D. schusterae Fried- rich, 1956 from Central America; and D. re- mand Friedrich, 1956 from Central America. The two latter species were noted by Foster (197 1) to be incompletely described. She noted a wide range of variability in the number and distribution of serrations of the parapodial lamellae in specimens of D. uncinata from North America, the Caribbean Sea and Chile. Since these characters were ones which Fried- rich (1956) had used to justify his 2 species, Foster strongly suggested that they were prob- ably conspecific with D. uncinata, but did not formally place them into synonymy. We con- cur with Foster's implied synonymy of Fried- rich's 2 species with D. uncinata, but prefer to delay a formal referral until the type ma- terial can be located and examined. Foster considered D. magna to be a distinct species because it completely lacks parapodial serra- tions and its accessory branchiae are reduced to short nodules. Dispio maroroi Gibbs, 1971 from the Solomon Islands also falls within the limits of variability for D. uncinata (see Light, 1977; 1978) and should probably be referred to synonymy. Like D. magna, D. glabrilamel- lata lacks parapodial serrations, but like D. un- cinata has digitiform accessory branchiae. The transversely barred capillary setae of D. glabrilamellata would at first glance appear Light (1978), however, has found similar setae in D. uncinata from California. The occurrence of bidentate hooded hooks in juveniles of D. uncinata was reported by Foster (1971) and have been observed in lar- val stages of the same species (Blake, unpub- lished data). We have also found such hooks in juvenile D. glabrilamellata; in a single fas- cicle there is a gradation of wear from the bi- dentate to unidentate condition, suggesting that unidentate hooks can result from worn bidentate hooks (however, see Foster, 1971, p. 78). Distribution: Queensland; New South Wales; Victoria. Genus Australospio gen. nov. Type-species: Australospio trifida sp. nov. Gender, feminine. Diagnosis: Prostomium with both anterior pointed projection and sub-distal lateral horns. No occipital cirrus. Peristomium reduced, with poorly-developed lateral wings. Proboscis ever- sible, sac-like. Branchiae from setiger 1, con- tinuing almost to posterior end, basally fused with notopodial lamellae in anterior setigers, becoming nearly free posteriorly. Notopodial lamellae well-developed, entire; neuropodial lamellae well-developed, tending towards bilo- bate condition in middle setigers. Notosetae all capillaries. Neuropodial setae including capil- laries, hooded hooks and inferior sabre-like setae. Nature of pygidium unknown. Remarks: Australospio is closely allied to Scolelepis Blainville, Aonides Claparede and Dispio Hartman in having gills and a pointed prostomium. Australospio differs in also bear- ing lateral horns on the prostomium, a charac- teristic typical of Malacoceros Quatrefages, Rhynchospio Hartman and Scolecolepides Eh- lers; however, in the last 3 genera, the prosto- mium is not anteriorly pointed. The branchiae begin on setiger 1 in both Australospio and Dispio and on setiger 2 in Aonides and Scole- lepis. The notopodial lamellae are at least ba- sally fused to the branchiae in Dispio, Aus- tralospio and Scolelepis, whereas they are completely free in Aonides. The hooded hooks are bidentate in Australospio, bidentate to tri- dentate in Aonides, unidentate in adult Dispio species and range from unidentate to multiden- tate in species of Scolelepis. Only a single spe- cies, Australospio trifida sp. nov. is known. Australospio trifida sp. nov. (Figure 11) Scolelepis sp. Hutchings, 1974, p. 182. Material examined: NEW SOUTH WALES. Botany Bay, Towra Beach, NSWSF Stations 286 (3 PARA- TYPES, NMV G2902); 298 (6 PARATYPES, AM W13002); 300 (1, AM W13003); 303 (2, AM W13004); 307 (3, AM W13005); 310 (HOLOTYPE, NMV G2901; 5 PARATYPES, NMV G2903); Towra Point, NSWSF Stations 287 (6, AM W13006); 292 (3, AM W13007); 297 (2, AM W13008); Wallis Lake, Zoster a bed, Dec. 1970, coll. Dixon and O'Gower (PARATYPE, AM W5006). VICTORIA. Western- V)4 JAMES A. BLAKE and JERRY D. KUDENOV Figure 11 — Australospio trifida gen et sp. nov. — a, anterior end, dorsal view; b, prostomium, dorsal view; c, right setiger 1, anterior view; d, right setiger 7, anterior view; e, right setiger 15, anterior view; f, right setiger 22, anterior view; g, right setiger 40, anterior view; h, right setiger 50, an- terior view; i, neirroseta from setiger 7; j, bidentate hooded hook. [Scale 1 = 100 /*m; 2 = 50 Mm; 3 = 200,um; 4 = 30wn] port, WPBES Stations 1713-2 (PARATYPE, NMV G2904); 1716-3 (PARATYPE, NMV G2905); Port Phillip Bay, Hobsons Bay, 14 April 1977, coll. J. E. Watson (3, NMV G2998). Description: Only incomplete specimens avail- able. A moderate-sized species, up to 12 mm long and 3 mm wide for 55 sctigers. Holotype a 50 setiger fragment, 12 mm long and 18 mm wide. Body widest anteriorly and dorsoven- trally flattened, tapering posteriorly, becoming oval in cross section. Body opaque white in alcohol, but with paired mid- ventral black pig- ment spots on setigers 2-10. Prostomium anteriorly pointed, with 2 la- terally projecting horns (Fig. lla-b); car- uncle reduced, extending posteriorly just into setiger 1. Three to 5 pairs of red, circular eyes. Peristomium reduced ventrally, enveloping prostomium laterally as indistinct wings, palps basally thickened, extending posteriorly for 11-12 segments. Branchiae from setiger 1, basally fused to dorsal lamellae in anterior and middle seti- gers, free in posterior setigers; branchiae long and thin throughout with cilia on both inner and outer margins. Dorsal lamellae short and cirriform on seti- gers 1-3 (Fig. lie), becoming broad and leaf- like through setiger 7 (Fig. lld-e), smaller in middle and posterior setigers (Fig. llf-h); pos- terior setigers with prolonged finger-like noto- podial lobe (Fig. llf-h); ventral lamellae leaf- like, entire in anterior setigers, narrow at first (Fig. lie), becoming broader (Fig. lid) by setiger 7; lamellae bilobed in middle and posterior setigers (Fig. lle-g), again becoming entire in far posterior setigers (Fig. 1 lh). SOUTHEASTERN AUSTRALIAN SPIONIDAE 195 Setal fascicles arranged in 2 dorsoventral tiers on setigers 1-3, increasing to 4 tiers over setigers 4-11, subsequently diminishing to 2 tiers by setiger 14, with a single tier by seti- ger 15. Setigers 6-9 conspicuous, appearing modified with heavy thickened capillaries (Fig. lid) numbering up to 140 in both noto- and neuropodia. Notosetae all capillaries through- out; heavy notosetal capillaries of setigers 6-9 including both granulated and non-granulated types with sheaths only in the latter. Neuro- podial capillaries of setigers 6-9 including granulated and unusual geniculate setae bearing fine bristles (Fig. Hi). Bidentate neuropodial hooded hooks (Fig. llj) from setiger 15-22, numbering 2-3 per neuropodium and accom- panied by capillaries and 3-4 granulated in- ferior sabre setae. Nature of pygidium unknown. Remarks: Australospio trifida is a unique spio- nid in having both a distally pointed prosto- mium and subdistal lateral prostomial horns. The species is also unusual in having an an- terior modified region with thickened capil- laries. Some of those capillaries are of a pe- culiar hirsuted geniculate type, heretofore unknown in the Spionidae and reminiscent of similar setae reported from species of the family Heterospionidae. Distribution; New South Wales; Victoria. In mud, Zostera and Posidonia beds. Genus Malacoceros Quatrefages, 1843 Malacoceros Quatrefages, 1 843 . Type-species: Spio vulgaris Johnston, 1827, designated by Pettibone, 1963. Colobranchus Schmarda, 1861. Type-species: C. tet- racerus Schmarda, 1861, by monotypy. Uncinia Quatrefages, 1865. Type-species: Colobran- chus ciliatus Keferstein, 1862 (= C. tetracerus Schmarda, 1861), by monotypy. Scolecolepis Malmgren, 1867. Type-species: Spio vul- garis Johnston, 1827, by original designation. Diagnosis: Prostomium with frontal horns; eyes present or absent; branchiae from setiger 1, partly fused with notopodial lamellae, extend- ing for most of body length. With only capil- lary notosetae. Neurosetae include capillaries, hooded hooks and sabre setae. Pygidium with anal cirri. Remarks: The present definition of Malacoce- ros conforms closely to that of Pettibone (1963) except that we prefer to elevate Rhyn- chospio to a full genus, thus separating out those species with branchiae from setiger 2. Three species of Malacoceros have been identified in our collections, of which 2 are newly described. KEY TO SPECIES OF MALACOCEROS FROM AUSTRALIA la. Neuropodial lamellae of middle and posterior segments with nipple-like projections M, indicus b. Neuropodial lamellae of middle and posterior segments rounded, without projections (Fig. 12c) 2 2a. Setiger 1 markedly reduced, with short branchiae (Fig. 13a); body without pigment M, reductus b. Setiger 1 well-developed, with long branchiae (Fig. 12a); body with dark brown pigment M. tripartitus Malacoceros indicus (Fauvel, 1928) Scolelepis inclica Fauvel, 1928a, p. 93; 1930, p. 35: 1953, p. 313; Monro, 1931, p. 25. Malacoceros indicus: Pettibone, 1963, p. 99; Day, 1967, p. 477; Foster, 1971, p. 50. Material examined: QUEENSLAND. Great Barrier Reef Expedition, 1929 (I, AM W2953). Remarks: The single specimen agrees well with the description given by Foster (1971); it is probably one of the specimens studied by Monro (1931). The prostomium is T-shaped, and the caruncle extends posteriorly to setiger 2. There are no eyes. The branchiae are ba- sally fused to the notopodial lamellae and be- gin on setiger 1. Notosetae are found on seti- ger 1 and tri- and quadridentate neuropodial hooded hooks are present from setiger 50. The middle and posterior neuropodial lamellae have nipple-like projections. Inferior tufts of neurosetal capillaries are present, each with about 20 setae per bundle. The pygidium was missing. Distribution: Queensland, Great Barrier Reef; circumtropical. 196 JAMES A. BLAKE and JERRY D. KUDENOV Figure 12 — Malacoceros tripartitus sp. nov. — a, anterior end, dorsal view; b, right setiger 8, an- terior view; c, right setiger 40, anterior view; d, right setiger 65, anterior view; e, pygidium, terminal view; f, tridentate hooded hook, frontal view; g, tridentate hooded hook, lateral view [Scale 1 = 200 Mm; 2 = 30 m] SOUTHEASTERN AUSTRALIAN SPIONIDAE 197 Malacoceros tripartitus sp. nov. (Figure 12) Material examined: VICTORIA. Port Phillip Bay, Werribee, Little River, Station 6, MSG Monitoring Program (P609), coll. J. D. Kudenov, 11 Jan. 1977 (HOLOTYPE, NMV G2916; 9 PARATYPES, NMV G2917, 2919-2923); 14 April 1977 (2 PARATYPES, NMV G2918); Yarra River, MSG Benthic Survey, 10 Oct. 1975, coll. G. Poore and J. D. Kudenov, black mud, 7 m, Sta. 128/1 (42, NMV G2924); 128/4 (76, NMV G2925); 128/5 (341, NMV G2926). Description: A moderately-sized species, up to 17 mm long and 11 mm wide for 85 setigers. Body widest anteriorly, gradually tapering pos- teriorly. Colour in alcohol dark brown, with dusky brown pigment granules distributed throughout epidermis in distinct banding pat- terns over dorsum, ventrum and all along body (Fig. 12a). Prostomium T-shaped, with distinct lateral horns, continuing posteriorly as a peculiar tri- lobed caruncle to anterior margin setiger 2 (Fig. 12a). Three pairs of round eyes, trape- zoidally arranged, posterior pair the larger; an- terior pair formed of 2 partially fused spots (Fig. 12a). Palps inserted posterior to level of eyes, extending posteriorly to setigers 9-10. Peristomium reduced, lacking lateral wings. Setiger 1 well-developed, with notosetae. Dorsal lamellae elliptical in anterior and mid- dle body setigers (Fig. 12b-c), becoming elon- gate, and narrow in posterior setigers (Fig. 12d). Anterior neuropodial lamellae subquad- rate (Fig. 12b), becoming elliptical in middle setigers (Fig. 12c), and subtriangular in pos- terior setigers (Fig. 12d). Anteriormost seg- ments with thickened presetal noto- and neuro- podial lobes (Fig. 12b). Notosetae forming spreading fascicles of sheathed, non-granulated capillaries. Neuro- setae including non-granulated capillaries, tri- dentate hooded hooks from setiger 3 1 and in- ferior sabre setae from setiger 25 (Fig. 12c-d); hooded hooks distinctly tridentate in frontal view (Fig. 12f), but appearing bidentate in lateral aspect (Fig. 12g). Branchiae elongate, strap-like, fused to no- topodial lamellae. Those on setiger 1 not as well-developed as on succeeding segments. All branchiae marginally pigmented with brown granules. Pygidium terminal with 6 anal cirri (Fig. 12e). Remarks: Malacoceros tripartitus most closely resembles M. tetracerus (Schmarda) from Europe in having a similarly shaped prosto- mium. M. tripartitus has tridentate hooded hooks, a trilobed caruncle and a heavily pig- mented body. M. tetracerus has bidentate hooded hooks, an entire caruncle and lacks body pigmentation. Foster (1971) pointed out that the hooded hooks of Malacoceros need to be examined under oil immersion in order to elucidate their structure; a re-examination of species such as M. tetracerus is necessary in order to improve our understanding of the systematics of the genus. Distribution: Victoria, Port Phillip Bay, in mud. Malacoceros reductus sp. nov. (Figure 13) Material examined; NEW SOUTH WALES. Bur- wood Beach, HDWBS Stations, Sept., Oct., Nov. 1975 (HOLOTYPE, AM W8906; 13 PARATYPES, AM W8882, 8904, 8905, 8907, 8908). Description: A moderately-sized species, up to 16 mm long and 8 mm wide for 75 setigers. Body widest anteriorly, evenly tapering pos- teriorly; subrectangular in cross section. Colour in alcohol: whitish-brown. Prostomium with anterior medial incision and very pronounced lateral horns; posteriorly a faint medial protuberance present between lateral nuchal processes that encompass palps (Fig. 13a). Two pairs of eyes, anterior pair the larger, each consisting of 2 fused ocelli; posterior pair round and small. Peristomium moderately developed, forming lateral wings. Palps extending posteriorly to setiger 6-7. Setiger 1 reduced, but with notosetae. An- terior notopodia fused with branchiae (Fig. 1 3b-c) ; both rami of posterior parapodia emerging from raised basal mounds (Fig. 13d). Notosetae with superior group of capillaries overlying main fascicle, the latter arranged in 2 tiers bearing sheathed, granulated capillaries (Fig. 13f), as well as non-granulated ones. An- terior setigers with about 20 setae per fascicle, 198 JAMES A. BLAKE and JERRY D. KUDENOV Figure 13 — Malacoceros reductus sp. nov. — a, an- terior end, dorsal view; b, left setiger 11, anterior view; c, left setiger 38, anterior view; d, left setiger 65, anterior view; e, pygidium; f, capillary notoseta; g, sabre seta; h, tridentate hooded hook, lateral view; i, tridentate hooded hook, frontal view. [Scale 1 = 200 Mm; 2=100 Mm; 3 = 20 Mm] and about 5 in superior tuft; fewer setae per fascicle in middle and posterior setigers. Neuro- setae include both granulated and non-granu- lated capillaries of which some are faintly spinous; with tridentate hooded hooks from setiger 20, replacing capillaries, and a single inferior sabre seta from setiger 20. Sabre setae with sheaths forming long distal processes (Fig. 13g). Hooded hooks with distinct, readily vis- ible apical teeth (Fig. 13h-i). Branchiae of setiger 1 very short, incon- spicuous (Fig. 13a), those of setiger 2 more elongate, strap-like and fused to dorsal lamel- lae; branchiae becoming reduced in length by setiger 35; absent from last 10-12 setigers. Pygidium terminal, small, with 3 pairs of short, digitiform anal cirri and a pair of dorsal swellings (Fig. 13e). Remarks: Malacoceros reductus most closely resembles M. juliginosa (Claparede) in having a similarly shaped prostomium. In M. reductus, setiger 1 is distinctly reduced, tridentate hooded hooks are present from setiger 20-21 and pig- mentation is lacking. In M. juliginosa, the parapodia of setiger 1 are only slightly re- duced, bidentate hooded hooks occur from seti- ger 30-40 and the body is heavily pigmented. Distribution: New South Wales in beaches. Genus Rhynchospio Hartman, 1936 emended Type-species; R, arenincola Hartman, 1936, by monotypy. Diagnosis: Prostomium with frontal horns, car- uncle variously developed. Occipital tentacle absent. Eyes present. Branchiae from setiger 2, free from dorsal lamellae or only fused ba- sally. With only capillary notosetae. Neuro- setae including capillaries, hooded hooks and sabre setae. Pygidium with cirri or lobes. Remarks: Rhynchospio Hartman has been con- sidered both as a full genus (Hartman, 1959; SOUTHEASTERN AUSTRALIAN SPIONIDAE 199 Day, 1967; Fauchald, 1977a) and as a sub- genus of Malacoceros (Pettibone, 1963; Fos- er, 1971). However, we consider the first ap- pearance of the branchiae to be an important axonomic characteristic for the family Spioni- iae. We therefore prefer to treat Malacoceros and Rhynchospio as full genera. Foster (1971) and others have included the fusion of branchiae and lamellae in their defini- tion of Rhynchospio. However, a basal fusion :of these 2 elements is barely evident in the 2 *valid species discussed by Foster (1971). In (this study, we have found 2 previously unde- ■scribed species in which the branchiae and la- mellae are entirely free from one another. The definition of Rhynchospio is accordingly emended herein. According to Hartman (1966) and Fauchald (1977a), spionids having bran- chiae and lamellae free from each other should be referred to the genus Mesospio Gravier, 191 1. Such a conclusion is unacceptable, how- ever, since Gravier (1911) clearly states in his discussion that M. moorei Gravier, 1911, the type-species, lacks frontal horns. His figures depict a spionid with an incised prostomium. Mesospio was referred to the genus Microspio Mesnil, 1896 by Foster (1971). We agree with that synonymy since Gravier's species, Meso- spio moorei, agrees well with the generic de- finition of Microspio. Foster (1971) reduced the number of valid species of Rhynchospio to 2. We are pre- sently re-examining some of her proposed synonyms as part of another study, but for the time being are following her revision. The following species are assigned to Rhyn- chospio 1 : l Rhynchospio microcera Dorsey, 1977 is herein referred to the genus Microspio. 1. R. glutaea, Ehlers, 1897), p. 83. Straits of Magellan. Scolecolepis cornifera Ehlers, 1913, p. 509. Fide Pettibone, 1963. Rhynchospio arenincola Hartman, 1936, p. 51. Fide Foster, 1971. Rhynchospio arenincola asiaticus Khle- bovitsch, 1959. Fide Foster, 1971. 2. R. inflates Foster, 1971, p. 57. Bahamas. o 3. R. glycera sp. nov. (see below). New South Wales. 4. R t australiana sp. nov. (see below). West Australia. Rhynchospio glycera sp. nov. (Figure 14) Material examined: NEW SOUTH WALES. Bur- wood Beach, HDWBS Station, Dec. 1975 (HOLO- TYPE, AM W8903). Description: Holotype incomplete, coiled, measuring approximately 8 mm long and 0-5 mm wide for 47 setigers. Body widest an- teriorly, elliptical in cross section. Colour in alcohol: opaque white. Prostomium well-developed, forming large, conical anterior projection bearing a pair of apical horns (Fig. 14a); with raised medially- incised ridge about half-way between anterior and posterior ends, bearing 2 pairs of trape- zoidally arranged eyes. Caruncle posteriorly merging with a prominent occipital lobe bear- ing 2 pairs of digitiform nuchal organs arising from a common base (Fig. 14a). Palps missing. Peristomium greatly reduced, not visible in dorsal view. Muscular pharynx visible through body wall. Setiger 1 reduced, with bluntly conical noto- podial and quadrate neuropodial lamellae; both notosetae and neurosetae present (Fig. 14a). Notopodial lamellae free from branchiae, anterior ones quadrate, those of posterior re- gion subclavate (Fig. 14b-c). Anterior neuro- podial lamellae quadrate, posterior ones tri- angular (Fig. 14b-c). Setal fascicles arranged in 2 parallel tiers. Notosetae all sheathed, non-granulated capil- laries, with somewhat opaque, lightly bristled sheaths (Fig. 14d-e). Anterior notopodia with about 20 short and stout and 3 longer superior capillaries per fascicle (Fig. 14b); posterior notopodia with about 15 narrow capillaries and no superior ones (Fig. 14c). Capillaries of an- terior setigers increase in length ventrally. Neu- rosetae sheathed, non-granulated, opaque, lightly bristled capillaries. No evidence of sabre setae. Anterior neuropodia with about 15 short and stout and 3 longer inferior capil- laries (Fig. 14b). Posterior neuropodia with narrow capillaries and hooded hooks (Fig. 200 JAMES A. BLAKE and JERRY D. KUDENOV Figure 14 — Rhynchospio glycera sp. nov. — a, an- terior end, dorsal view; b, right setiger 8, anterior view; c, right setiger 45, anterior view; d-e, anterior capillary setae [inset, cross section of same]; f, quadridentate hooded hook, lateral view; g, same, an- terolateral view; h, same without hood. [Scale 1 = 200 Mm; 2 = 200 Mm; 3 = 20 Mm) 14c); quadridentate hooded hooks from setiger 27, these appearing bidentate in lateral view, with inconspicuous, medial quaternary tooth inserted between 2 longer intermediate-length apical teeth (Fig. 14f-h), all 3 in 1 plane over- lying enlarged main fang. Branchiae on setiger 2, slightly shorter than subsequent ones, not fused to dorsal lamellae, and continuing to end of specimen. Inner mar- gins ciliated; no evidence of transverse ciliated dorsal ridges. Nature of pygidium not known. Remarks: Rhynchospio glycera differs from other species of the genus in having a conical prostomium with apically-mounted frontal horns resembling the prostomial antennae seen in the errant family Glyceridae, in having 4 nuchal organs arising from the caruncle and in the highly unusual arrangement of anterior notopodial capillaries, where they increase in length from the superior to inferior the posi- tion. Distribution: New South Wales. Rhynchospio australiana sp. nov. (Figure 15) Material examined: WEST AUSTRALIA. Perth, Cottesloe Beach, in calcareous algae and Idanthrysus tubes -3-66 m, coll. H. Paxton, 14 Feb. 1970 (HOLO- TYPE, AM W4362). Description: Holotype complete, measuring 5 mm long and 0-3 mm wide for 31 setigers. Body widest anteriorly, tapering gradually pos- teriorly, trapezoidal in cross section. Pigmen- tation in patterns of single red chromatophores dorsomedially on setigers 6-13; colour in alco- hol: light brown. Prostomium anteriorly prolonged, blunt bearing 2 broad lateral horns (right one miss- SOUTHEASTERN AUSTRALIAN SPIONIDAE 201 Figure 15 — Rhynchospio australiana sp. nov. — a, an- terior end, dorsal view; b, left setiger 5, an- terior view; c, left setiger 23, anterior view; d, pygidium, dorsal view; e, su- perior capillary notoseta, setiger 5; f, tri- dentate hooded hook, lateral view; g, inferior sabre seta [inset, detail of same, not to scale]. [Scale 1 = 200 Mm; 2 = 20 Mm] ing from the type), inflated, slightly incised where it overlies anterior margin of peristo- mium; with raised boss posteriorly (Fig. 15a). Two pairs of eyes, arranged in shallow cres- cent, distal pair the larger. No occipital cirrus; palp scars on either side of prostomium be- hind eyes. Peristomium greatly reduced; la- teral wings absent. Setiger 1 with short notopodial lobes, lack- ing notosetae (Fig. 15a). Neuropodia laterally reduced, with inflated lamellae and capillary neurosetae. Both pre- and postsetal lobes of both rami inflated (Fig. 15b) in anterior seti- gers; the presetal lobes becoming reduced in posterior se tigers (Fig. 15c); notopodial la- mellae cylindrical, distally blunt, conical in setigers 2-13; thereafter digitate. Neuropodial 202 JAMES A. BLAKE and JERRY D. KUDENOV lamellae inflated, elliptical to irregularly rounded. Notosetal capillaries distally sheathed, sub- lanceolate, granulated, arranged in 2 tiers per fascicle. Anterior tiers with 15-16 capillaries and 4-5 slightly isolated superior ones; middle setigers with 10 such capillaries, no superior tufts. Superior notopodial capillaries of setigers 2-6 with sheaths bearing large vesicles (Fig. 15e), inferior capillaries lacking such vesicles in sheaths; capillaries from posterior tiers with lightly bristled sheaths. Neurosetae arranged in 2 tiers; anterior tiers bearing 6-7 sheathed granulated capillaries per tier; posterior seti- gers with 2 superior capillaries, 3 hooded tri- dentate hooks and inferior sabre seta. Anterior capillaries similar to those of notopodia. Hooded hooks and sabre setae from setiger 8; the hooks with an inconspicuous tooth in- serted above the more prominent apical tooth; margin of hood uneven, not serrated (Fig. 15f); sabre setae granulated, distally tapering with prolonged sheath (Fig. 1 5g). Branchiae of setiger 2 slightly smaller than those of succeeding setigers; not fused to dor- sal lamellae. Branchiae overlapping middor- sally, continuing to within last 10 body seg- ments; inner and outer margins ciliated, no transverse ciliary ridges. Pygidium terminal, trilobate (Fig. 1 5d). La- teral lobes flattened, plate-like, subequal, the single middorsal lobe a small knob. Remarks: Rhynchospio australiana differs from other species of the genus by the shape of the prostomium, which has a subterminal inflated, incised area and 2 broad lateral horns on the terminal portion, by lacking notosetae on seti- ger 1, the form of the vesiculated sheaths of the anterior capillary setae and the unique struc- ture of the pygidium. Distribution: West Australia. Genus Scolecolepides Ehlers, 1907 Type-species: Scolecolepides benhami Ehlers, 1907, by monotypy. Diagnosis: Prostomium with frontal or lateral horns. Branchiae present from setiger 1 , limited to anterior setigers or continuing throughout body. Anterior setae mostly capillary, with ventral acicular spines present. Bidentate hooded hooks present in both neuropodia and notopodia. Pygidium with cirri surrounding anus. Remarks: The following species of Scolecole- pides are herein recognized: 1. S. benhami Ehlers, 1907, p. 14. New Zea- land. 2. S. viridis (Verrill, 1873), p. 600 [as Scole- colepis]. New Jersey. S. arctius Chamberlin, 1920, p. 17. Fide Foster, 1971. 3. S. aciculatus sp. nov. (see below). Victoria. Scolecolepides aciculatus sp. nov. (Figure 16) Material examined: VICTORIA. Westernport, Flin- ders, in sand on rock platform, 15 Dec. 1974, coll. J. D. Kudenov (HOLOTYPE, NMV G2891; 100+ PARATYPES, NMV G2892); Port Phillip Bay, Wer- ribee, MSG Monitoring Programme (P609), coll. J. D. Kudenov, 11 Jan. 1977, Stations 16 (24, NMV G2999); 18 (81, NMV G30OO-3OO1); 20 (39, NMV G3OO2-3003). Description: A large species, measuring up to 37 mm long and 1-5 mm wide for 140 seti- gers. Body dorsoventrally flattened anteriorly, becoming cylindrical and narrower posteriorly. Colour in alcohol: tan to brown. Prostomium broadly rounded on anterior margin, bearing 2 lateral horns (Fig. 16a). Posteriorly, prostomium tapering as blunt car- uncle terminating at posterior margin of seti- ger 1; eyes absent; palps short, thick, extending posteriorly to setiger 8. Setigers 1 and 2 well-developed, but noto- podial lamellae and branchiae shorter than on succeeding setigers. Anterior notopodial la- mellae elongate, foliaceous, fused basally with branchiae (Fig. 16c); posterior notopodial la- mellae shorter, similarly fused to branchiae (Fig. 16d). Neuropodial lamellae much smaller, less foliaceous than those of notopodia. Capillary notosetae lacking granulations or sheaths. Capillary neurosetae with minute granulations near tips in setigers 1-6, mostly re- placed in setigers 6-8 with 2 dorsoventral tiers of acicula, but with a few inferior capillaries remaining (Fig. 16c); acicula sharply pointed or distally blunt (Fig. 16e), blunted acicula thicker and darker than pointed ones; acicula becoming partially replaced by capillary setae SOUTHEASTERN AUSTRALIAN SPIONIDAE 203 Figure 16 — Scolecolepides aciculatus sp. nov. — a, anterior end, dorsal view; b, posterior end, lateral view; c, right setiger 13, anterior view; d, right setiger 60, anterior view; e, acicular neurosetae from anterior setigers showing different forms; f, hooded hook. [Scale 1 = 200 wtn; 2 = 200 /*m; 3 = 20 ^m; 4 = 20 nm] 204 JAMES A. BLAKE and JERRY D. KUDENOV before first bidentate hooded hooks on setiger 39-55; hooded hooks with a greatly reduced inconspicuous secondary tooth (Fig. 16f). No- topodial hooded hooks beginning on setieers 80^85. Branchiae begin setiger 1, long, basally fused to postsetal lobes, continuing to near posterior of body. Pygidium with 2 long su- perior cirri and up to 6 smaller ones surround- ing terminal anus (Fig. 16b). Remarks: Scolecolepides aciculatus and S. ben- hami closely resemble each other in bearing lateral horns on the prostomium; they differ in this respect from 5. viridis, which has short frontal horns. 5. aciculatus has 20 or more neuroacicula of 2 types (pointed and blunt) from setiger 6-8, whereas S. benhami has only 4-5 pointed acicula beginning on setiger 9. The pygidium of S. aciculatus has 2 long su- perior cirri and 6 smaller ventral cirri, while S. benhami has numerous short cirri. Species of Scolecolepides appear to be limited to waters of reduced salinity (George, 1966; Estcourt, 1967). 5. aciculatus has been taken near sew- age outfalls and may also tolerate low salini- ties. Distribution: Victoria, Westernport Bay, Port Phillip Bay. Genus Laonice Malmgren, 1867 Laonice Malmgren, 1867. Type-species: Nerine cir- rata M. Sars, 1851, designated by Malmgren, 1867. Spionides Webster and Benedict, 1887. Type-species: S. cirrata Webster and Benedict, 1887 [Homonym, = L. cirrata (Sars)], by monotypy. Aricideopsis Johnson, 1901. Type-species: A. mega- lops Johnson, 1901, monotypy. Diagnosis: Prostomium anteriorly rounded to slightly incised, lacking frontal horns; caruncle extending posteriorly for variable number of setigers; occipital tentacle present. Peristomium reduced, generally without lateral wings. Bran- chiae from setiger 2, separated from dorsal la- mellae, continuing for at least i of body. Noto- podia with capillaries only; neurosetae including capillaries, hooded hooks and sabre setae. Inter- parapodial genital pouches present. Pygidium with anal cirri. Remarks: A considerable degree of confusion persists regarding this genus because of 2 dif- ferent approaches to species identification: So- derstrom (1920) emphasized the distribution of genital pouches and recognized that a close relationship exists between their first appear- ance and the end of the atokous region. This approach was followed by Banse and Hobson (1968) who indicate that although the brood pouches of their newly described species, L. pugettensis, first appear from setiger 2-7, oocytes could be seen through the body wall at or just behind setiger 25 (Banse and Hobson, 1968, pp. 26-27); they recognized 5 species of Laonice in their paper. A second approach (Fauvel, 1927; Foster, 1971) questions the reliability of this relationship and points out that the distribution of genital pouches is highly variable. Foster (1971) considers Laonice cir- rata (Sars, 1851) as the only valid species. It is essential, however, to determine the role of genital pouches in reproduction in order to understand the taxonomic significance of their distribution on the body. At present there is little information available on repro- duction in Laonice. Morphology of the genital pouches before and after spawning has been described by Soderstrom (1920) and Orrhage (1964). Hannerz (1956) cultured and de- scribed larvae isolated from plankton and de- termined that the eggs and sperm were spawned into sea water with development being entirely planktonic. No mention was made of the role genital pouches might play in re- production. We are herein describing a species with an anterior region bearing eggs and geni- tal pouches and a posterior region bearing sperm. The occurrence of anterior brood pouches may reflect the type of reproduc- tion. In our simultaneous hermaphrodite, the genital pouches are more anteriorly located than in most forms. Forms with brood pouches beginning more posteriorly may have separate sexes or possibly represent stages in sequential hermaphroditism. We prefer to re-examine the Laonice-com- plex utilizing the distribution of genital pou- ches, together with the configuration of the caruncle, capillary setae, hooded hooks and parapodia. In the present survey we have found 3 distinct forms of Laonice, all of which differ from the description of L. cirrata as given by Foster (1971). Although a thorough SOUTHEASTERN AUSTRALIAN SPIONIDAE 205 study of Laonice from worldwide areas is needed, we feel justified in describing each form as a distinct species. Each is compared to L. cirrata as defined by Foster (1971); other known species of Laonice are insufficiently de- scribed. KEY TO SPECIES OF LAONICE FROM SOUTHEASTERN AUSTRALIA la. Genital pouches from between seti- gers 1-2 or 2-3; hooded hooks bi- or quadridentate 2 b. Genital pouches from between seti- gers 6-7; hooded hooks quadriden- tate (Fig. 17f) L. quadridentata 2a. Hooded hooks bidentate, hood mar- gins smooth (Fig. 18g); nuchal organs paired, extending to setiger 7; geni- tal pouches beginning between seti- gers 2-3 L. bassensis b. Hooded hooks quadridentate (Fig. 17i), hood margin serrated (Fig. 17j); nuchal organ single, extending to setigers 50-55; genital pouches be- ginning between setiger 1-2 , L. hermaphroditica Laonice quadridentata sp. nov. (Figure 17a-g) Material examined: VICTORIA. Port Phillip Bay, PPBES Stations 930/1 (HOLOTYPE, NMV G2954) and 37 PARATYPES as follows: 908/5 (1, NMV G2960); 909/2 (1, NMV G2976); 910/5 (1, NMV G2966); 925/5 (1, NMV G2961); 926/1 (2, NMV G2971); 926/2 (1, NMV G2957); 930/4 (1, NMV G2977); 931/3 (1, NMV G2964); 931/4 (1, NMV G2981); 932/2 (2, NMV G2973); 932/3 (1, NMV G2963); 932/4 (1, NMV G2967); 932/5 (3, NMV G2970); 933/4 (2, NMV G2959); 933/5 (1, NMV G2980); 936/1 (5, NMV G2958); 944/4 (1, NMV G2974); 946/1 (1, NMV G2962); 954/3 (1, NMV G2969); 957/2 (1, NMV G2972); 957/5 (1, NMV G2953); 965/4 (1, NMV G2979); 972/4 (1, NMV G2978); 982/4 (1, NMV G2955); 983/1 (1, NMV G2975); 983/2 (1, NMV G2956); 983/5 (1, NMV G2968); 985/5 (1, NMV G2965). Description: A large species, up to 50 mm long and 3 mm wide for 136 setigers. Colour in al- cohol: pink with faint grey pigment on la- mellae. Prostomium bluntly rounded, 2 pairs of eyes, all fused together into transverse patch; caruncle extending to setiger 18 (Fig. 17a); with long cirriform occipital tentacle; peris- tomium forming slight lateral wings emphasiz- ing grooves next to prostomium, with thick palps extending posteriorly for 6 setigers. Setiger 1 about i as large as setiger 2, with subrectangular to rounded parapodial lamellae, containing prominent setal fascicles. Noto- podial lamellae from setiger 2 distally pointed, becoming elliptical (Fig. 17b); anterior neuro- podial lamellae elliptical, distally blunt, sub- sequently becoming subtriangular and distally pointed (Fig. 17b). Notosetae and neurosetae including sheathed, granulated capillaries. Quadriden- tate hooded hooks bearing 3 small, obscure apical teeth surmounting larger main fang (Fig. 17c-d) from neuropodium of about seti- ger 40, best seen in lateral view; inferior sabre setae from setiger 16, these heavily granulated, sheathed, distally prolonged (Fig. 17e). Branchiae reduced at first, smaller than and free from notopodial lamellae, these gradually increasing in length in succeeding setigers, be- coming longer than dorsal lamellae. Genital pouches first present between setigers 6-7 (Fig. 17f). Pygidium with 11-12 anal cirri (Fig. 17g). Remarks: Laonice quadridentata differs from L. cirrata in having quadridentate, rather than bidentate, hooded hooks and heavily granu- lated inferior sabre setae. Distribution: Victoria, Port Phillip Bay. Laonice hermaphroditica sp. nov. (Figure 17h-k) Material examined; QUEENSLAND. Moreton Bay, Middle Banks, Sta. 29, Sept. 1972 (HOLOTYPE, NMV G2893); Sta. 35, Sept. 1972 (PARATYPE, QM G11591), Dec. 1972 (2 PARATYPES, QM G11592); Sta. 57, Dec. 1973 (PARATYPE, QM G11593), coll. W. Stephenson. Description: A moderate-sized species, up to 24 mm long and 0-8 mm wide for 80 setigers. Body widest anteriorly, somewhat flattened, elliptical in cross section, becoming cylindrical posteriorly. Colour in alcohol: opaque white with diffuse grey on dorsal lamellae. Prostomium bluntly rounded on anterior margin (Fig. 17h); caruncle extending pos- teriorly to setiger 50, present between dorsal transverse segmental lamellae; occipital ten- 206 JAMES A. BLAKE and JERRY D. KUDENOV Figure 17 — Laonice quadridentata sp. nov. — a, an- terior end, dorsal view; b, right setiger 10, anterior view; c, quadridentate hooded hook, lateral view; d, quadridentate hooded hook, frontal view; e, sabre seta; f, lateral view showing brood pou- ches and dorsal lamellae; g, pygidium, dorsolateral view — Laonice hermaphrodi- tica sp. nov. — h, anterior end, dorsal view; i, quadridentate hooded hook, lateral view; j, same, without hood, anterolateral view; k, oocyte [250 Mm]. [Scale 1 — 100 Mm; 2 = 20 Mm; 3 =z 200 Mm; 4 = 50 Mm; 5 = 100 Mm]. tacle short, thin; 2 pairs of eyes, posterior pair with 1 member often missing, darker in colour, irregularly shaped and placed; anterior pair lighter in colour. Peristomium poorly-de- veloped; palps missing. Setiger 1 reduced with short, narrow noto- podial lamellae bearing only 2-3 capillary setae (Fig. 17h); subsequent notopodial lamellae broad, bluntly rounded. Anterior neuropo- dial lamellae foliaceous, distally pointed, sub- sequently becoming reduced and rudimentary. SOUTHEASTERN AUSTRALIAN SPIONIDAE 207 Notopodia with sheathed capillaries; quad- ridentate hooded hooks in neuropodia from setigers 24-25, often appearing bidentate in lateral view (Fig. 17i-j); aperture of hood distinctly serrated (Fig. 17i). Nature of pygidium unknown. Genital pouches first present between setigers 1-2, these large, delicate and easily detached. Large pri- mary oocytes, up to 300 ^m in diameter pre- sent in anterior 54-55 setigers (Fig. 17k). Short headed sperm present in coelomic cavity from about setiger 56. Remarks: Laonice hermaphroditica differs from L. cirrata in having quadridentate, rather than bidentate hooded hooks, the caruncle is prolonged over 50 setigers instead of 12-30 and genital pouches first occur from setiger 1 rather than from setiger 4 or thereafter. L. hermaphroditica differs from L. cirrata and L. quadridentata in its apparent simultaneous hermaphroditism and in having hooded hooks with serrated apertures. Distribution: Queensland, Moreton Bay. Laonice bassensis sp. nov. (Figure 18) Material examined: VICTORIA. Bass Strait, ESSO- GIPPS Sta. 8, 106 m, dredged sandy clay bottom, 148° 43' 50'E. 38° 50' 8'S, May 1969 (HOLOTYPE, AM W13009); Sta. 9, 144 m, May 1969 (PARATYPE, AM W13010). Description: Largest specimen, the holotype, measuring 13 mm long and 1-2 mm wide for 42 setigers. Body widest anteriorly, elliptical in cross section, becoming cylindrical posteriorly. Colour in alcohol: opaque white. Prostomium medially incised on anterior margin (Fig. 18a-b); caruncle continuing to setiger 2, surrounded laterally by 2 nuchal or- gans extending posteriorly to setiger 7; with small digitiform occipital tentacle; no eyes. Peristomium reduced, with grooves separating prostomium from peristomium; palps missing. Setiger 1 reduced, dorsally displaced, abut- ting against prostomium (Fig. 18a-b); anterior notopodial and neuropodial lamellae foliose (Fig. 18c), with rounded apices; posterior dor- sal lamellae triangular, elliptical, with pointed inferior processes (Fig. 18d). All setae arranged in 2 tiers. Capillaries sheathed, shafts with or without granulations distributed along striae (Fig. 18f); anterior notopodia with about 50 capillaries; posterior setigers with about 35 per ramus; neuropodial capillaries similar to those of notopodia, but fewer in numbers. Bidentate hooded hooks from setigers 22-23 (Fig. 18g). Inferior sabre setae from setigers 9-10. Branchiae shorter than dorsal lamellae for first 4-5 segments, thereafter larger, not fused with lamellae. Genital pouches commencing between setigers 2-3 (Fig. 18e). Nature of pygidium unknown. Remarks: Laonice bassensis differs from L. cirrata in having an incised, rather than an en- tire prostomium, paired rather than single nu- chal organs, ventrally directed rather than tri- angular dorsal lamellae in posterior setigers and genital pouches beginning from setiger 2 rather than from setiger 4-50. L. bassensis differs from L. hermaphroditica and L. quad- ridentata in having bidentate rather than quad- ridentate hooded hooks. Distribution: Victoria, Bass Strait. General Remarks on the Prionospio-complex Foster (1969; 1971) reorganized Prionospio Malmgren into 5 genera: Prionospio, Para- prionospio Caullery, Apoprionospio Foster, Aquilaspio Foster and Minuspio Foster. With exception of Paraprionospio, which has pinnate branchiae beginning on setiger 1, the other 4 genera have branchiae from setiger 2 and are separated on the basis of branchial form. Thus, variation in a single characteristic has been used to set aside 4 genera. Perhaps anticipating objections to the scheme, Foster (1971, p. 80) suggested that 5 subgenera of Prionospio might be more acceptable. Day (1973), however, dis- agreed with the need for Apoprionospio and Light (1977) preferred to use subgenera, but synonymized Aquilaspio with Paraprionospio. Other authors have used all 5 as full genera or as subgenera. In the present study, we have found 2 here- tofore undescribed Australian species which cause considerable confusion to Foster's ar- rangement. One represents a new genus: it has a well-developed peristomium, setiger 1 is 208 JAMES A. BLAKE and JERRY D. KUDENOV Figure 18 — Laonice bassensis sp. nov. — a, anterior end, dorsal view; b, prostomium, antero- lateral view; c, right setiger 8, anterior view; d, right setiger 41, anterior view; e, genital pouches from about setiger 50, lateral view; f, anterior capillary seta [in- set, detail of shaft, not to scale]; g, biden- tate hooded hook. [Scale 1 = 200 Mm; 2 = 50 Mm; 3 = 50 Mm; 4 = 20 Mm] distinct and lacks a dorsal ridge between the parapodial bases, bears 18-22 pairs of bran- chiae from setiger 1 and has an unusual py- gidium and lacks anal cirri. The second spe- cies belongs to the genus Prionospio, the first and third pairs of branchiae are pinnate and the second and fourth pairs cirriform. The pin- nate branchiae have very few pinnules, how- ever so few, in fact that some specimens are missing pinnules on one or both members of a gill-pair. Individual specimens might be thus classified with either Prionospio (sensu stricto) or a modified version of A poprionospio, based upon Foster's generic definitions. We consider that the form of branchiae is not an adequate justification for separating genera when used as the sole criterion. We include a suite of what appear to be more con- servative characteristics in arranging the Prio- nospio-complox. The position of the branchiae, degree of development of the peristomium, the distinctness and degree of development of setiger 1, structure of the hooded hooks and the form of the pygidium are considered the major generically important characteristics. Bran- chial form is considered to be of secondary importance. Any discussion of this complex must include the genus Streblospio Webster (see also Foster, 1971; Light, 1977). The fol- lowing genera and subgenera are recognized in the Prionospio-complcx: 1 . Paraprionospio Caullery, 1914 SOUTHEASTERN AUSTRALIAN SPIONIDAE 209 2. Orthoprionospio gen. nov. 3. Streblospio Webster, 1879a 4. Prionospio Malmgren, 1867 (sensu lato) a. Subgenus Aquilaspio Foster, 1971 b. Subgenus Minuspio Foster, 1971 c. Subgenus Prionospio (sensu stricto) [includes Apoprionospio Foster, 1971] Characteristics separating these genera may be seen in Table 1 . The first 3 genera have branchiae beginning on setiger 1 and well-developed peristomia with lateral wings developed to various degrees (see Table 1. In Paraprionospio and Orthopriono- spio setiger 1 is well-developed and distinct from the peristomium, while in Streblospio seti- ger 1 is reduced and fused with the peristomium. Streblospio and Paraprionospio share peculiar- ities of hooded hook structure and development of anterior ridges and collars, while Orthoprio- nospio has hooded hooks approaching the con- dition seen in Prionospio and lacks anterior dorsal ridges. Paraprionospio, Orthoprionos- pio and Streblospio all have different pygidial structures. In Prionospio, the branchiae begin on setiger 2, with the peristomium and setiger 1 both reduced and fused. The hooded hooks have inconspicuous secondary hoods. Various kinds of dorsal ridges or crests may be present or absent, but normally only on the postbran- chial setigers. The pygidium in Prionospio is similar to that of Paraprionospio; it includes a medial and 2 ventrolateral cirri. About 30 species of Prionospio are currently recogni2ed (Foster, 1971; Fauchald, 1972; this study). We follow Foster (1971) in recognizing groups of species according to whether the branchiae are all pinnate (subgenus Aquilaspio), all cirri- form (subgenus Minuspio) or a combination of cirriform and pinnate (subgenus Prionospio). We do not consider these taxa as full genera since their separation is presently based on only a single character. However, we do consider that they are useful categories to separate spe- cies groups. Apoprionospio cannot be justified as even a subgenus because it is based on a particular ordered arrangement of pinnate and cirriform branchiae. We consider such a character to be important only at the species level. In this regard we agree with Day (1973). We cannot agree with Light (1977) that Aquil- aspio is a synonym of Paraprionospio because all of the major generic characters of Prionos- pio (Table 1) apply equally to species assigned to Aquilaspio and are very different from the generic characters of Paraprionospio. Streblospio has not been encountered in our studies. Paraprionospio pinnata^ Orthoprionos- pio cirriformia gen. et. sp. nov. and 9 species of Prionospio, including 5 new species, are des- cribed below. Genus Paraprionospio Caullery, 1914 Type-species: P. pinnata (Ehlers, 1901), designated by Caullery (1914). Diagnosis: Prostomium wedge-shaped, slightly curved on anterior margin, without posterior caruncle; peristomium well-developed forming erect lateral wings often completely overlap- ping prostomium. Setiger 1 well-developed, dis- tinct from peristomium. Notopodial lamellae of setigers 1-5 enlarged, thereafter becoming smaller, rounded. Distinct transverse dorsal ridge present between branchial bases of seti- ger 1; 3 pairs of branchiae from setiger 1, all pinnate. Anterior setae all capillaries in both rami; posterior noto- and neuropodia with multidentate hooded hooks; hooded hooks with secondary teeth closely applied to main fang; with prominent secondary hood. Ventral sabre setae present. Pygidium with 1 long medial cirrus and 2 shorter ventrolateral cirri. Paraprionospio pinnata (Ehlers, 1901) Prionospio pinnata Ehlers, 1901, p. 163; 1908, p. 110; Augener, 1927, p. 351; Okuda, 1937, p. 247; Fauvel, 1953, p. 323; Hartman, 1960, p. 114; Ima- jima and Hartman, 1964, p. 286; Hartmann- Schroder, 1965, p. 211; Day, 1967, p. 488. Paraprionospio pinnata: Foster, 1971, p. 102 (Sy- nonymy). Material examined: NEW SOUTH WALES. Stock- ton Bight, dredge, 5-9-9 m, coU. N. Ruello, July 1970 (2, AM W4410); Sydney, Malabar, 28 m, AMSBS Station CA+5 March 1974 (4, AM W6501); B3S1 (1, AMW6502). Description: Prostomium curved on anterior margin; spindle-shaped, no caruncle; 0-4 eyes. Segment 1 achaetous, together with peristo- mium forming lateral wings overlapping pro- 210 JAMES A. BLAKE and JERRY D. KUDENOV TABLE 1 Taxonomic Criteria Separating Genera of the Prionospio-complcx Genus Peristomium Setiger 1 Paraprionospio Enlarged with prominent wings Well-devel., free from peristomium Branchiae a. Begin b. Structure a. Setiger 1 b. 3 pairs, all pinnate Dorsal ridges or collars Hooded hooks Pygidium Prominent ridge across setiger 1 Large apical teeth, closely applied to main fang; secondary hood prominent With 1 long dorsomedial cirrus and 2 short ventro- medial cirri Orthoprionospio Enlarged with moderate wings Well-devel., free from peristomium a. Setiger 1 b. 18-22 pairs all cirriform Absent Smallapicalteeth With 1 ventral not closely lobe and 4 small applied to main lateral papillae fang; secondary hood not apparent Streblospio Enlarged with low wings Reduced, a. Setiger 1 fused to b. 1 pair, peristomium cirriform Inconspicuous ridge across setiger 1; prominent hood or collar across dorsum of setiger 2 Large apical Simple, with 2 teeth not closely rounded lobes; applied to appendages main fang; absent secondary hood prominent Prionospio Reduced, with or without lateral wings Reduced, fused to peristomium a. Setiger 2 b. 3-40 pairs, all pinnate, cirriform, or combination of both types on post- Ibranchial segments Absent. Some species with mem- branous crests across dorsum Small secondary With 1 long teeth not closely dorsomedial and applied to main fang; secondary hood inconspicuous 2 short ventro- lateral cirri stomium; palps long with conspicuous mem- branous basal sheath. Setiger 1 well-developed with prominent parapodial lobes and a con- spicuous transverse dorsal ridge connecting bases of first pairs of branchiae. Anterior para- podial lamellae of setigers 1-5 foliose, those of neuropodia less foliaceous and smaller than in preceding segments; notopodia becoming pro- gressively longer and thinner in postenor seg- ments. Anterior setae in both rami all capillaries. Notosetae anteriorly arranged in 3 tiers; capil- laries appearing bilimbate, with prominent sheaths bearing granulations. Multidentate hooded hooks and sabre setae in neuropodia from setiger 9, accompanied by several capil- laries- notopodial hooded hooks commencing posterior to setiger 20, these hooks with 8 secondary teeth in 2 rows above main fang, with a clear primary hood and striated internal secondary hood, the latter tightly applied to shaft w . _ Pinnules of branchiae numerous, subject to various configurations depending upon stage of growth and or regeneration {fide Foster, 1971). tt , - Pygidium bearing 1 long medial cirrus and 2 short lateral cirri, similar to those seen in the genus Prionospio. Distribution: New South Wales; cosmopolitan. Genus Orthoprionospio gen. nov. Type-species: Orthoprionospio cirriformia sp. nov. Gender, feminine. _. Diagnosis: Prostomium rounded on anterior SOUTHEASTERN AUSTRALIAN SPIONIDAE 211 margin; caruncle absent. Peristomium well-de- veloped, forming prominent lateral wings along- side prostomium. Setiger 1 well-developed, dis- tinctly separate from peristomium, parapodia indistinguishable from those of succeeding seti- gers; notopodial lamellae of all anterior setigers foliose, becoming reduced in posterior segments; dorsal ridges lacking. Branchiae beginning on setiger 1, 18-22 pairs, all cirriform. An- terior setae all capillary; multidentate hooded hooks in both neuropodia and notopodia of posterior segments; hooks with apical teeth not closely overlapping main fang; without second- ary hood; inferior sabre setae present in mid- dle and posterior segments. Pygidium with 1 reduced, conical ventral lobe and 4 very small lateral papillae. Remarks: See Table 1 and foregoing discussion. A single species, O. cirriformia is known. Orthoprionospio cirriformia sp. nov. (Figure 19) Material examined: NEW SOUTH WALES. Bot- any Bay, Georges River, Picnic Point, coll. NSWSF, 5 July 1972 (7, AM W13011). VICTORIA. Yarra River, MSG Benthic Survey, 10 Oct. 1975, coll. G. Poore and J. D. Kudenov, black mud, 7 m, Stations 128/1 (HOLOTYPE, NMV G2846; 12 PARATYPES, NMV G2847); 128/2 (3, NMV G3004); 128/4 (7, NMV G3005); 128/5 (7 PARATYPES, NMV G2848). TAS- MANIA. Derwent Estuary, 5 Jan. 1977, coll. R. But- termore (TM K842). Description: A moderate-sized species, up to 22 mm long for 100 segments. Body light tan to brown in alcohol with black pigment present on palps and between some anterior parapo- dia. Prostomium broadly rounded on anterior margin, bell-shaped (Fig. 19b), without car- uncle. Three pairs of eyes, anteriormost pair cup-shaped followed by more rounded pairs, all arranged in straight rows. Peristomium with raised lateral peristomial wings (Fig. 19a-b). Setiger 1 well-developed with enlarged dor- sal lamellae; those of all anterior setigers en- larged, subtriangular in shape (Fig. 19f); pos- terior notopodial lamellae reduced to short lobes; neuropodial lamellae smaller, more rounded. Notosetae including fascicles of deli- cate capillaries lacking granulations; neuro- setae similar to notosetae but fewer in number; 2-3 granulated inferior sabre setae per neuro- podium, and beginning setiger 18-22 (Fig. 19e); multidentate hooded hooks in neuro- podia from setiger 29-34, and in notopodia from setiger 62-68; neuropodial hooks num- bering 4-6 per fascicle accompanied by capil- laries; hooks bearing 8 apical teeth in 2 rows above main fang (Fig. 1 9c-d) ; notopodial hooks in far posterior setigers long, projecting prominently from body and accompanied by long capillaries (Fig. 19g). Branchiae long, cirriform, beginning setiger 1, continuing for 18-22 segments (Fig. 19a, f). Pygidium as described for genus (Fig. 19g-h). Ecology: The species is common in areas of re- duced salinity and according to R. Buttermore is \ . . the most numerous species in the Der- went Estuary [Tasmania]' (personal communi- cation). The species is also common in the Yarra River (Poore and Kudenov, 1978 b). Distribution: New South Wales; Victoria; Tas- mania. Genus Prionospio Malmgren, 1867 Prionospio Malmgren, 1867. Type-species: P. steen- strupi Malmgren, 1867, by monotypy. Ctenospio M. Sars, 1867. Type-species: C. ptumosus M. Sars, 1867, by monotypy. Apoprionospio Foster, 1969. Type-species: A. dayi Foster, 1969, by monotypy. Diagnosis: Prostomium more or less straight, medially incised or rounded on anterior mar- gin, without frontal horns; caruncle variously elongated. Peristomium fused in varying de- grees with setiger 1, often forming low lateral wings. Parapodia of setiger 1 reduced, noto- podia on branchial bearing segments enlarged; postbranchial notopodia becoming smaller, in- conspicuous; dorsal folds or crests present or absent on postbranchial segments, rarely on branchial segments. Branchiae all pinnate (sub- genus Aquilaspio Foster, 1971), all cirriform (subgenus Minuspio Foster, 1971) or both pin- nate and cirriform (subgenus Prionospio [sensu stricto]), branchiae always limited to anterior setigers. Anterior setae all capillaries; hooded hooks in posterior noto- and neuropodia; hooks bi-, tri- or multidentate; inferior sabre setae present. Pygidium with 1 long medial SOUTHEASTERN AUSTRALIAN SPIONIDAE 213 3a. Three pairs of pinnate branchiae; nil, 1897, p. 90; Soderstrom, 1920, p. 232; Fauvel, low dorsal crest on setiger 7 I 92 ?, P- 60; Day, 1963, p. 418; 1967, p. 489; Hart- P aucklandica man ' 1965 k, P* 152 > Hartmann-Schroder, 1971, p. b. Four pairs of pinnate branchiae; dor- fso*^ ^ P * 84 {Synonymy); Light ' 1977 ' sal crests absent P. multipinnulata Spiophanes tenuis Verrill, 1879, p. 176; 1881, p. 4a. First pair of branchiae long, pinnate, 320. Fide Pettibone, 1954, p. 282. second and third pairs shorter, cirri- Prionospio tenuis Verrill, 1881, p. 370. Fide Foster, form, fourth pair long, thin, cirri- u ) 91hv / u ; . ; . „ „ fr»rtr» (Ti\rr ofww „,uu ~!Lu«i J P- 84 (Synonymy). [But see Fauchald, 5a. First and third pairs of branchiae p ? 972 ' f 197] ; . , . . ^ 10£1 , . , r x t - Prionospio malmgrem var. dubia Day, 1961, p. 489. pinnate (pinnules sparse), second and Fi ^ e Foster, 1971, p. 84. fourth pairs cirrifonn (Fig. 22a); car- Prionospio malmgreni: of authors [partim]. [Not uncle long, reaching to setiger 5 Claparede, 1870]. Fide Fauchald, 1972, p. 197; P. paucipinnulata Light > 1977 > P- 80 - b. Branchiae otherwise; caruncle reach- r "«?% --^ NEW SOUTH WALES-Wal- 9 w *w* rtV *wv« j is Laifg^ Forster, boatsheds, 0-3 m, Zostera beds, 24 ing to setiger 2 6 May 1968, coll. H. Paxton (1, AM W4242; 2, AM 6a. First and fourth pairs branchiae pin- W4254); Wallis Lake, weed bed mixed, Dec. 1970, nate, second and third pairs cirrifonn coll. Dixon and O'Gower (l, AM W5018; 2, AM (Fig. 20a-b); hooded hooks multi- J* 5020 * ?^ tte Bay ^ thid J Q ^; m ^f^^l' A ♦ * (vr oiv\ n DlX0n and O'Gower, Dec. 1970 (1, AM W5021); aentate ^lg. Zin; / Sydneyj North Head> AMSBS Station, 32 m, May b. First 3 pairs of branchiae cirrifonn, 1972 (AM W6505). VICTORIA. 113 km south Lakes fourth pair pinnate (Fig. 23a); Entrance, 93-6 m, 148° 24' 50"E-39° 00' 00"S, sand, hooded hooks tridentate (Fig. 23d) co11 - c - Pni PP s & AM W13012). P< tridentata Description: A small species, up to 15 mm 7a. Peristomium forming lateral peristo- i ong f or 170 setigers. Prostomium wedge- mial wings (Fig. 20a) P. steenstrupi shaped, broadly rounded anteriorly, caruncle b. Peristomium not forming lateral to posterior margin of setiger 1 (Fig. 20a); 2-3 wm 8 s 8 pairs of eyes. Peristomium reduced, fused dor- 8a. Prostomium smoothly rounded on sal i y to set i g er 1. Parapodia of setiger 1 well- anterior margin (Fig. 20b); promin- developed with notopodial lamellae forming ent dorsal crests on setigers 11-18 erect peristomial wings; notopodial lamellae (Fig. 20d); first 3 pairs of branchiae thereafter more rounded, forming low trans- short, fourth pair long (Fig. 20b) verse dorsal crests on some specimens. An- P* ctustrahensis terior setae all capillaries; neuropodial sabre b. Prostomium broad or slightly incised setae beginning on setiger 12; multidentate on anterior margin (Fig. 21a); dorsal hooded hooks from setiger 15-18 ventrally fold on setiger 5 and low dorsal crests and from setiger 30 _ 40 dorsaliy; these hooks on variable number of setigers from multidentate with 10-12 apical teeth arranged setiger 9 (Fig. 21c); first and fourth in 2 vertical rows above main fang. Four pairs pairs of branchiae longest, second of branchiae with first and fourth pinnate, sec- and third pairs short (Fig. 21a) ond ^ t hi r d cirriform. Pygidium with 1 long F. queenslandica and 2 short cirri. Prionospio (Prionospio) steenstrupi Remarks: See comments below for P. queens- Malmgren, 1867 landica ' (Figure 20a) Distribution: New South Wales; Victoria; cos- Prionospio steenstrupi Malmgren, 1867, p. 202; Mes- mopolitan. 212 JAMES A. BLAKE and JERRY D. KUDENOV Figure 19 — Orthoprionospio cirriformia gen. et sp. nov. — a, anterior end, lateral view; b, anterior end, dorsal view; c, hooded hook, lateral view; d, hooded hook, frontodorsal view; e, sabre seta; f, right setiger 8 in anterior view; g, posterior end, dorsal view; h, posterior end, ventral view. [Scale 1 = 200 Mm; 2 = 200 tm; 3 = 20 Am; 4 = 10 (im\ cirrus and 2 short ventrolateral cirri or thick- ened lobes. KEY TO SPECIES OF PRIONOSPIO FROM SOUTHEASTERN AUSTRALIA la. Branchiae all cirriform (Fig. 25a) P. cirrifera b. Branchiae all pinnate or pinnate and cirriform 2 2a. Branchiae all pinnate (Fig. 24a; 25b) 3 b. Branchiae both pinnate and cirriform (Fig. 20a-b) 4 214 JAMES A. BLAKE and JERRY D. KUDENOV Figure 20— Pnonospto steenstrupi Malmgren-a, anterior end, dorsal vkw-Prionospio austra- iwnsis nov. sp.—b, anterior end, dorsal view; c, left setiger 4, anterior view: d, right setiger 13, anterior view; e, capillary notoseta, setiger 4 from posterior tier; f, capil- ary notoseta, setiger 4 from anterior tier; g, sheathed neuroseta, setiger 4- h capil- lary neuroseta, setiger 13; i, sabre seta; j-k, multidentate hooded hooks; 1, posterior end dorsal view- Prionospio ehlersi Fauvel-m, anterior end, dorsolateral view. [Scale 1 = 200 nm; 2 = 200 m; 3 = 50 Am; 4 = 20 m ] SOUTHEASTERN AUSTRALIAN SPIONIDAE 215 Prionospio (Prionospio) australiensis sp. nov. (Figure 20b-l) Material examined: QUEENSLAND. Moreton Bay, Middle Banks, Sta. 8, Sept. 1976 (HOLOTYPE, NMV G2842; 14 PARATYPES, NMV G2843), coll. W. Stephenson. Description: A moderately-sized species, up to 34 mm long and 0-75 mm wide for 50 seg- ments. Colour in alcohol: opaque white. Prostomium bluntly rounded on anterior margin, continuing posteriorly as narrow car- uncle to middle setiger 2 (Fig. 20b). Two pairs of eyes, anterior pair cup-shaped, posterior pair irregularly shaped, both pairs about equal in size. Peristomium fused with setiger 1, not forming peristomial wings. Setiger 1 well-developed with enlarged no- topodia and notosetae, but smaller than suc- ceeding setigers; setigers 2-5 with enlarged no- topodial lamellae (Fig. 20c), thereafter la- mellae rounded, extending across dorsum as prominent dorsal crests on setigers 11-18 (Fig. 20d); dorsal lamellae pronounced on remain- ing segments; neuropodia of anterior setigers small, rounded at first (Fig. 20c), gradually be- coming wider. Capillary notosetae of setigers 1-11 arranged in 3 tiers, with first tier contain- ing shorter lanceolate setae with prominent distal granulations and sheaths (Fig. 20f) and second and third tier containing longer setae with granulations throughout length of shafts and indistinct sheaths (Fig. 20e); thereafter dorsal capillaries delicate, arranged in 2 poorly differentiated tiers. Capillary neurosetae small, unilimbate (Fig. 20g-h). Inferior neuropodial sabre seta from setiger 11 (Fig. 20i); multiden- tate hooded hooks from neuropodia setiger 17 and from notopodia from about setiger 45; neuropodia with about 8 hooks accompanied by capillaries; these hooks with 8-12 apical teeth arranged in 2 rows above main fang (Fig. 20j-k). Four pairs of branchiae with the first and fourth pairs pinnate, the second and third pairs cirriform (Fig. 20b). First 3 pairs short, the fourth long; pinnules numerous on posterior face of pinnate branchiae. Pygidium with 1 long medial cirrus and 2 rounded lateral lobes (Fig. 201). Remarks: See comments below following P. queenslandica. Distribution: Queensland, Moreton Bay. Prionospio (Prionospio) queenslandica sp. nov. (Figure 21) Material examined: QUEENSLAND. Moreton Bay, Middle Banks, Sta. 4, Sept. 1976 (HOLOTYPE, NMV G2839; 27 PARATYPES, NMV G2840), coll. W. Stephenson. Description: A moderately-sized species, up to 23 mm long for 65 setigers. Body generally opaque white in alcohol. Prostomium broad, with 4 small peaks on anterior margin, appearing slightly incised in some specimens (Fig. 21a); caruncle continu- ing posteriorly to posterior border of setiger 2; 2 pairs of eyes, anterior pair cup-shaped, posterior pair irregular, consisting of several individual ocelli fused together. Peristomium fused with setiger 1, lacking peristomial wings. Setiger 1 well-developed, with notosetae but with smaller parapodial lobes than succeeding setigers. Setigers 2-5 with enlarged foliose no- topodial lamellae (Fig. 21b), with well-de- veloped transverse dorsal fold on setiger 5; subsequent dorsal lamellae low, merging with dorsum in setigers 6-8, but extending across dorsum on setigers 9-13 forming low dorsal crests (Fig. 21c), these low crests indistinct, but continuing on setigers posterior to 14 (Fig. 2 Id); neuropodia smaller, rounded in anterior setigers, becoming more elongated from about setiger 10. Notosetae and neurosetae of an- terior setigers include spreading fascicles of stout, lanceolate granulated capillary setae (Fig. 21g) and delicate smooth capillary setae (Fig. 2 If). One inferior neuropodial sabre seta per fascicle from setiger 10 (Fig. 21e), sabre setae bearing a prominent granular sheath; multi- dentate hooded hooks in neuropodia from seti- ger 14 and in notopodia from setiger 35-40, hooks numbering 7-8 in neuropodia, accom- panied by capillaries; these hooks with 10-12 apical teeth arranged in 2 rows above main fang (Fig. 21h-i). Four pairs of branchiae, first and fourth pairs long and pinnate, second and third pairs short, cirriform (Fig. 21a); with pinnules paired along axis of branchia, but absent from tip. 216 JAMES A. BLAKE and JERRY D. KUDENOV Figure 21 — Prionospio queenslandica sp. nov. — a, anterior end, dorsal view; b, right setiger 5, anterior view; c, right setiger 12, anterior view; d, right setiger 20, anterior view; e[ sabre seta; f, smooth capillary notoseta from setiger 5; g, granulated capillary notoseta from setiger 5; h, hooded hook, lateral view; i, same, frontal view [Scale 1 = 200 Mm; 2 = 300 Mm; 3 = 50 Mm; 4 = 20 Mm; 5 = 10 Mm] SOUTHEASTERN AUSTRALIAN SPIONIDAE 217 Pygidium with a single long medial cirrus and 2 lateral bulges. Remarks: Foster (1971) redescribed P. steen- strupi and compared it with P. fallax Soder- strom, 1920. She noted that P. fallax possesses a high dorsal crest on setiger 7, which P. steen- strupi presumably lacks. P. steenstrupi, how- ever, may possess or lack low or high dorsal crests on some postbranchial setigers (Foster, 1971; Light, 1977). Light (1977) found that specimens which he referred to P. steenstrupi from Canada, San Francisco Bay and southern California possess moderate to well-developed dorsal folds on a variable number of segments from setiger 7. He also suggested that P. cris- tata Foster, 1971 might be a juvenile of P. steenstrupi, since it has dorsal crests on seti- gers 7 and 9 and overlaps characteristics on P. steenstrupi from the northeastern Pacific. None the less, after a study of Australian materials it is apparent to us that several closely related species forming a suite of sib- lings may eventually need to be sorted from populations currently lumped by Foster (1971) and others under the name P. steenstrupi. There is no question that at least 3 species in the steenstrupi group occur in Australia. What we call P. steenstrupi has a reduced peristo- mium, which forms peristomial wings in con- junction with notopodia of setiger 1. P. aus- traliensis and P. queenslandica on the other hand, have well-developed peristomia, but lack any development of lateral peristomial wings. P. australiensis is distinguished from P. queenslandica in having the prostomium smoothly rounded on the anterior margin in- stead of broad or slightly incised, by having prominent dorsal crests on setigers 11-18 in- stead of low crests from setiger 9, in lacking instead of having a dorsal fold on setiger 5 and in having a short pinnate gill on setiger 2, instead of a long one. Distribution: Queensland, Moreton Bay. Prionospio (Prionospio) ehlersi Fauvel, 1928 (Figure 20m) Prionospio ehlersi Fauvel, 1928b, p. 10; 1936, p. 61; Day, 1967, p. 490; Hartman, 1965a. p. 151; Hart- man and Fauchald, 1971, p. 105; Gibbs, 1971, p. 170. Prionospio krusadensis: Lee, 1976, p. 65. [Not Fauvel, 1929 = P. aucklandica Augener], Material examined: QUEENSLAND. Moreton Bay, Middle Banks, Sta. 9, Sept. 1976 (2, NMV G2841), coll. W. Stephenson. Description: A small species, up to 8 mm long and 0-8 mm wide for about 35 setigers. Colour in alcohol: opaque white. Prostomium broadly rounded on anterior margin, caruncle short, terminating at posterior margin of setiger 1 (Fig. 20m). Two pairs of eyes, anterior pair small, inconspicuous, con- sisting of several small individual spots, pos- terior pair enlarged, irregularly shaped. Peri- stomium reduced, fused with setiger 1, without lateral wings. Setiger 1 reduced, notosetae present with small noto- and neuropodia. Setigers 2-6 with enlarged subtriangular notopodial lamellae, these largest on setigers 2-4; following noto- podia low, occasionally forming low ridges across dorsum, but prominent dorsal crests lacking; neuropodia small rounded through- out. With genital pouches between successive neuropodia of anterior segments beginning between setigers 2-3. Anterior segments with dense fascicles of granular unilimbated capillary notosetae ar- ranged in 3 tiers; neurosetae fewer, less dense; neuropodia with a stout inferior sabre seta and multidentate hooded hooks from setiger 20; 5-6 neuropodial hooks accompanied by capil- laries; hooks with 6-8 apical teeth in 2 rows above main fang and a conspicuous secondary hood. Four pairs of branchiae on setigers 2-5. First pair long, pinnate, second and third pairs short, cirriform with fourth pair long, thin, cirriform. Pygidium typical for genus. Remarks: Specimens from Queensland agree well with descriptions from other areas. P. ehlersi is the only Prionospio observed in Aus- tralia to date with genital pouches. The bran- chial arrangement is unique. Distribution: Queensland; Morocco; South Af- rica; eastern North America; South America; Solomon Islands; Korea. Prionospio (Prionospio) paucipinnulata sp. nov. (Figure 22) Material examined: VICTORIA. Port Phillip Bay, 218 JAMES A. BLAKE and JERRY D. KUDENOV Figure 22 — Prionospio pausipinnulata sp. nov. — a, anterior end, dorsal view; b, left setiger 4, anterior view; c, left setiger 20, anterior view; d, left setiger 10, anterior view; e, posterior end, dorsal view; f, hooded hook, lateral view; g, same, frontal view; h, sabre seta. [Scale 1 = 200 nm\ 2 = 200 Mm; 3 = 300 Mm; 4 = 20 Mm] PPBES Stations 985 (HOLOTYPE, NMV G2837; 21 PARATYPES, NMV G2838; 80, NMV G3006); 907 (2, NMV G3007); 918 (1, NMV G3008); 944 (4 NMV G3009); 945 (6, NMV G3010). Description: A small species, up to 7 mm long and 0-4 mm wide for 53 setigers. Colour in alcohol: opaque white. Prostomium rounded on anterior margin, continuing posteriorly as long caruncle to pos- terior setiger 4 (Fig. 22a); 2 pairs of eyes; an- terior pair smaller, cup-shaped; and a larger, irregularly shaped posterior pair. Peristomium broad with short lateral wings. Setiger 1 well-developed, smaller than seti- ger 2; setigers 2-5 with well-developed sub- triangular notopodial lamellae (Fig. 22b); no- topodia of setiger 7 fused, forming large dor- sal crest (Fig. 22c); similar crest also present on setiger 8 but less-developed; notopodia thereafter smaller, appearing subtriangular (Fig. 22d); neuropodia small, rounded through- out. Notosetae of first and subsequent setigers including fascicles of granulated capillaries, those on setiger 1 being thinner; neurosetae of setigers 1-9 in fascicles of granular capillaries; a single inferior sabre seta in each neuropodium from setiger 1 (Fig. 22h). Multidentate hooded hooks from neuropodia of setiger 13 with 1-3 hooks at first, thereafter increasing to 6-7 in middle body segments, those hooks with 6-8 apical teeth arranged in 2 rows above main fang (Fig. 22f-g); similar hooks in notopodia from setiger 34-35. SOUTHEASTERN AUSTRALIAN SPIONIDAE 219 Four pairs of branchiae in setigers 2-5; first and third pairs pinnate, second and fourth pairs cirriform; short second pair (Fig. 22a); branchial pinnules sparse, usually no more than 4-7 on a single branchia; in some cases, pinnules reduced to 1-3 per gill or may even be lacking from 1 or both members of a bran- chial pair. Pygidium with 1 long medial cirrus and 2 lateral lappets (Fig. 22e). Remarks; Prlonospio paucipinnulata is unique in the arrangement of pinnate (first and third) and cirriform (second and fourth) branchiae and the reduced number of pinnules on the pinnate branchiae. The long caruncle is also unusual among species of Prionospio. Distribution: Victoria. Prionospio (Prionospio) tridentata sp. nov. (Figure 23) Material examined: NEW SOUTH WALES. Botany Bay, Towra Point, coll. NSWSF, April 1973 (2, NMV G2844; 1, NMV G2845); Newcastle, Burwood Beach, HDWBS Station, March 1975 (HOLOTYPE, AM W8829), April, 1975 (PARATYPE, AM W8900). Description: A small species, up to 10 mm long and 0-5 mm wide for 60 segments. Colour in alcohol: light tan. Prostomium trapezoidal, anteriorly broad to narrow with medial incision, tapering pos- teriorly to narrow caruncle terminating mid- way onto setiger 2 (Fig. 23a); 2 pairs of eyes, anterior pair widely spaced, cup-shaped, pos- terior pair rounded, close together. Peristo- mium reduced, without lateral wings. Setiger 1 well-developed, with notosetae, parapodial lamellae smaller than those of fol- lowing setigers; notopodia of setigers 2-6 with enlarged triangular lamellae, those of seti- ger 7 merging across dorsum as a dorsal crest; dorsal lamellae of succeeding setigers low, merging with dorsum, not forming crests; neu- ropodial lamellae rounded throughout. Setigers 2-5 with dense fascicles of bilimbate granulated capillaries (Fig. 23b). Neuropodia bearing an inferior sabre seta with partial distal sheath from setiger 11 (Fig. 23c); tridentate hooded hooks having 2 secondary teeth in line above main fang (Fig. 23d), in neuropodia from seti- ger 19 and in notopodia from setiger 28; neu- ropodial hooks number 4-5 with capillaries. Four pairs of branchiae on setigers 2-5, first pair narrow, thin, cirriform, second and third pairs thick, cirriform, fourth pair thick, long, pinnate (Fig. 23a) with numerous pinnules on posterior face. Pygidium with short medial cirrus and 2 lateral bulges. Remarks: Six species of Prionospio have been previously described as having the first 3 pair of branchiae cirriform and the fourth pinnate (Fauchald, 1972). Five of those species and P. tridentata are compared in Table 2. P. nova Annenkova, 1938 is poorly known and based upon incomplete specimens. Since there is no type material, or any specimens, for that mat- ter, of P. nova available (Light, personal com- munication), the species is here considered a nomen dubium. P. tridentata is most closely related to P. caspersi (Table 2). P. tridentata differs by having tridentate, instead of biden- tate, hooded hooks, and the prostomium is dis- tinctly bilobed to incised instead of entire. Foster (1969) removed this entire species group from Prionospio to her newly erected genus Apoprionospio. We do not consider that the ordered arrangement of cirriform and pin- nate branchiae is a character of sufficient weight to warrant generic or subgeneric rank and have included Apoprionospio in the sy- nonymy of Prionospio (see foregoing discussion above, p. 209). Distribution; New South Wales. Prionospio (Aquilaspio) multipinnulata sp. nov. (Figure 24) Material examined: NEW SOUTH WALES. Mer- imbula, Sta. MER 404L-S, core sample, Posidonia, op- posite Spencer Park, 18 March 1976, coll. J. H. Day, et al. (1, AM W11736); MER 278M, netting over weed beds, 3 Dec. 1975 (1, AM W11737). VIC- TORIA. Port Phillip Bay, PPBES Station 953 (HOLO- TYPE, NMV G2836; 5 PARATYPES, NMV G2833- 2835), Description; Only incomplete specimens avail- able. A moderately-sized species, up to 36 mm long for 85 setigers. Colour in alcohol: opaque white. 220 JAMES A. BLAKE and JERRY D. KUDENOV Figure 23 — Prionospio tridentata sp. nov. — a, an- terior end, dorsal view; a', prostomium from holotype; b, anterior capillary noto- seta; c, sabre seta; d, hooded hook. [Scale 1 = 200 fim; 2 = 200 Mm; 3 = 20 Am] Prostomium broadly rounded on anterior margin, tapering posteriorly as short caruncle to anterior margin of setiger 2 (Fig. 24a); 4-6 pairs of eyes, anterior pair largest, posterior pair often divided. Peristomium with short lateral wings. Setiger 1 well-developed, notosetae present, notopodial lobe enlarged, neuropodium small, rounded; setigers 2-5 with enlarged notopodia (Fig. 24d), thereafter notopodia smaller, of low profile, merging with dorsum but not forming dorsal crests (Fig. 24e-f); neuropodia smaller, rounded throughout body. Anterior capillary notosetae and neurosetae granulated; neuropo- dia with an inferior sabre seta lacking a sheath from setiger 10; multidentate hooded hooks accompanied by capillaries in neuropodia from setiger 18; 5-6 such hooks and capillaries in notopodia from setiger 30; hooded hooks with 6-8 apical teeth arranged in 2 rows above main fang (Fig. 24b-c), surmounted by a single api- cal tooth (Fig. 24c). Four pairs of pinnate branchiae on setigers 2-5, first pair the largest (Fig. 24a). Numerous pinnules irregularly arranged on posterior side. Nature of pygidium unknown. Remarks: Prionospio multipinnulata is closely related to P. peruana Hartmann-Schroder, 1962 from Peru and to P. tetelensis Gibbs, 1971 from the Solomon Islands in having 4 pairs of pinnate branchiae. P. multipinnulata SOUTHEASTERN AUSTRALIAN SPIONIDAE 221 TABLE 2 Some Taxonomic Characteristics of Species of the Prionospio pygmaea Group Species Anterior Peristomium Sabre Hooded Distribution Occurrence References margin of setae hooks: num- of branchial of dorsal prostomium begin be of apical teeth pinnules crests dayi Broad with With low Setiger 2-3 2 rows Setiger Foster, 1969; Foster small distal peak lateral wings 11 7 only 1971 caspersi Laubier Broad to weakly iconcave Reduced; no lateral wings Setiger 11 1 Irregular iSe tiger 7 only Laubier, 1962 pygmaea Hartman Broad with small distal peak With low lateral wings Setiger 11 6-8 2 rows Setiger 7 & subsequent segments* Hartman, 1961; Foster, 1971; Light, 1978 saldanha Rounded Well-de- Setiger Day veloped; with 12 low lateral wings Irregular Setiger Day, 1961; 19 to end 1967 of body vermilliensis Fauchald Broad Well-de- veloped with low lateral wings 2 rows Fauchald, 1972 tridentata Concave to Reduced; Setiger n. sp. deeply no lateral 1 1 incised wings Irregular Setiger This study 7 * The presence of very thin dorsal membraneous cresthave only recently been discovered in P. pygmaea (see Light, 1978). differs from both species in lacking dorsal crests on postbranchial setigers; from P. tetelen- sis in having a shorter caruncle and lacking notosetae on setiger 1; and from P. peruana in having the prostomium broadly rounded on the anterior margin, rather than straight across with an apical peak, and further differs in hav- ing instead of lacking short peristomial wings. Distribution: New South Wales; Victoria. Prionospio (Aquilaspio) aucklandica Augener, 1923 (Figure 25b-g) Prionospio aucklandica Augener, 1923, p. 69. Prionospio krusadensis Fauvel, 1929, p. 182; 1930, p. 38; 1953, p. 236. Fide Foster, 1971. Material examined: NEW SOUTH WALES. Wallis Lake, sandy mud, Dec. 1970, coll. Dixon and O'Gower (1, AM W5017); Merimbula, Sta. MER 197A, transect on S. bank of estuary 200 m seaward of bridge, coll. J. H. Day et ah 6 Oct. 1975 (1, AM W11739; 2, AM W11740); Botany Bay, Towra Point, Towra Beach, NSWSF Stations 317 (4); 207 (2); 321 (6); 325 (5); 329 (1); 331 (6); 332 (6); 333 (1); 335 (7); 344 (2) (NMV G3 135-3 144). VICTORIA. Port Phillip Bay, PPBES Stations 919 (3); 929 (7); 932 (1); 940 (1); 941 (1); 944 (87); 953 (41); 971 (1) (NMV G3092-3099); Werribee; MPSG Sta. 1833, coll. G. Poore and J. D. Kudenov, Nov. 1975 (1, NMV G3091); Westernport, WPBES Stations 1701 (2); 1703 (5); 1705 (13); 1706 (6); 1708 (3); 1709 (1); 1710 (4); 1715 (1); 1716 (9); 1718 (4); 1720 (1); 1721 (12); 1724 (1); 1735 (2); 1736 (6); 1737 (16); 1738 (16); 1740 (22); 1741 (15) (NMV G3O72-3O90). Description: A moderately-sized species, up to 23 mm for 110 segments. Colour in alcohol: opaque white. Prostomium rounded on anterior margin with 3 small prominences, these obscured if 222 JAMES A. BLAKE and JERRY D. KUDENOV Figure 24 — Prionospio multipinnulata sp. nov. — a, an- terior end, dorsal view; b-c, hooded hooks; d, left setiger 4, anterior view; e, left seti- ger 10, anterior view; f, right setiger 20, anterior view. [Scale 1 = 200 ^m; 2 = 200 Am; 3 = 20 Mm] poorly preserved (Fig. 25b); caruncle thick, continuing to posterior end of setiger 1; 2 pairs of eyes, the anterior pair somewhat further apart, oval, posterior pair more closely spaced, cup-shaped. Peristomium narrow, with- out lateral hood; palps thick, extending pos- teriorly for 6-7 segments. Setiger 1 well-developed, notosetae present; dorsal lamellae of setigers 2-5 enlarged, mem- branous, thereafter notopodia reduced; setiger 7 with low dorsal crest; neuropodia smaller, more rounded throughout body. Noto- and neuropodia of setigers 1-9 with heavy bilim- bate capillaries exhibiting prominent granula- tions (Fig. 25g). Capillaries thereafter smaller; inferior sabre setae in setiger 10 (Fig. 25f); multidentate hooded hooks appearing in neuro- podia from setiger 17-18 and in notopodia from setiger 27-30; hooks bearing 5 tiers of apical teeth arranged in multiple rows above main fang; lower tier with 3-4 teeth, second and third tiers with 2-4 teeth each, fourth tier with 2 teeth and fifth with a single apical tooth (Fig. 25c-e); primary hood long and inflated; secondary hood present, but incon- spicuous. Three pairs of branchiae on setigers 2-4, all pinnate, first pair the largest (Fig. 25b). Py- gidium with 1 long cirrus and 2 short lobes. Remarks: The large number of teeth on the hooded hooks reported for this species appears to be unique for the genus Prionospio. It is the only member of the genus known to bear 3 pairs of pinnate gills. Distribution: New South Wales; Victoria; In- dia; Auckland Islands. Prionospio (Minuspio) cirrifera Wiren, 1883 (Figure 25a) Prionospio cirrifera Wiren, 1883, p. 409; Fauvel, SOUTHEASTERN AUSTRALIAN SPIONIDAE 223 Figure 25 — Prionospio cirrifera Wiren — a, anterior end, dorsal view — Prionospio aucklandica Augener — b, anterior end, dorsal view; c, hooded hook, lateral view; d, same frontal view; e, diagram of hooded hook tooth pattern at maximal development of secondary teeth; f, sabre seta; g, anterior capillary notoseta. [Scale 1 = 200 ^m; 2 = 20 fim] 1927, p. 62; 1953, p. 324; Soderstrom, 1920, p. 237; Day, 1967, p. 486; Hartmann-Schroder, 1971, p. 324. Prionospio multibranchiata Berkeley, 1927, p. 414; Pettibone, 1967, p. 12. HOMONYM [not Fauvel, 1928a]. Prionospio delta Hartman, 1965a, p. 11, 46, 147. Fide Foster, 1971, p. 108. Minuspio cirrifera: Foster, 1971, p. 108 (Synonymy). Prionospio {minuspio) cirrifera: Light, 1977, p. 82. Material Examined: QUEENSLAND. Deception Bay, Burpengary Creek, Jan. 1975 (50+, AM W7127- 7128, 7130-7131). NEW SOUTH WALES. New- castle, Hunter River region, April, June 1971, coll. P. A. Hutchings and N. Ruello (20+, AM W13013- 13015); Tuncurry Creek, Site 25-3, Aust. Littoral Soc., Jan. 1973 (1, AM W5673); Merimbula, Sta. MER 120G, Oyster lease transect, Oct. 1975, coll. J. H. Day et ah (3, AM W11723); MER 117F Oct. 1975 (3, AM W11724); MER 148W, Spencer Park tran- sect, Zostera, muddy sand, Oct. 1975 (1, AM W11728); MER 278M, netting over weed beds, Dec. 1975 (1 AM W11727); MER 366N-Z, grab sample from channel, March 1976 (1, AM W11726). VIC- TORIA. Port Phillip Bay, PPBES Stations 901 (93); 913 (2); 921 (11); 932 (2); 945 (54); 952 (215); 962 (1); 978 (2); 983 (5) (NMV G3146-3154); Hobsons Bay-Yarra River, MSG Stations, Feb.-Nov. 1975, coll. G. Poore and J. D. Kudenov, 128 (1554, NMV G3155); 131 (165, NMV G3156); 134 (109, NMV G3157); Paynesville, Gippsland Lakes, from jetty, 2 m, sand, coll. J. D. Kudenov (20, NMV G3145). Description: Prostomium anteriorly rounded, with caruncle extending to posterior end of setiger 2 (Fig. 25a). Peristomium with low lateral wings partially enclosing prostomium. Six to 12 pairs of cirriform branchiae from setiger 2; these varying in length. Low dorsal crests present on a variable number of post- branchial setigers. Anterior setae all capillaries; multidentate hooded hooks in neuropodia from setiger 12-19 and in notopodia from setiger 15-40; these hooks with 8-10 apical teeth ar- 224 JAMES A. BLAKE and JERRY D. KUDENOV ranged in 2 rows over main fang. Pygidium with long medial cirrus and 2 lateral cirri. Remarks: The variability seen in different populations of P. cirrifera suggest that a de- tailed global analysis of the morphology of this species is needed. We have not observed genital pouches on any of our specimens, nor did Light (1977) in his San Francisco Bay study. Foster (1971), however, indicated that they did occur in some of her material. Distribution: Queensland; New South Wales; Victoria; cosmopolitan. Genus Spiophanes Grube, 1860 Type-species: Spiophanes kroeyeri Grabs, 1860, by monotypy. Diagnosis: Prostomium anteriorly broad, rounded or bell-shaped with frontal horns de- veloped to varying degrees; eyes present or ab- sent; with or without occipital tentacle. Bran- chiae entirely lacking. Setigers 1-4 shifted dor- sally with well-developed dorsal and ventral lamellae; setigers 5-14 generally with intcrra- mal thread glands forming supernumerary bacil- lary setae. Neuropodia of sctiger 5 and sub- sequent segments poorly-developed, with re- duced neuropodia; notopodia also reduced, more laterally placed. Anterior notosetae all capillaries, hooks or crochets only in neuro- podia, generally from about setiger 15, these hooks bi-, tri-, or quadridentate, with or with- out hood; 1-2 large curved spines in neuropodia of setiger 1, inferior sabre setae generally oc- curring in middle and posterior neuropodia. Pygidium with 2 or more anal cirri, with or without other processes or papillae. KEY TO SPECIES OF SPIOPHANES FROM SOUTHEASTERN AUSTRALIA 1 a. Occipital tentacle present; prosto- mium bell-shaped, broad anteriorly, without frontal horns; nuchal organs single (Fig. 27a) S. kroeyeri b. Occipital tentacle absent; prosto- mium rounded anteriorly or with dis- tinct frontal horns; nuchal organs paired 2 2a. Prostomium rounded anteriorly; nu- chal organs extend posteriorly to setiger 4 then curve back anteriorly toward prostomium (Fig. 26a) . S. wigleyi b. Prostomium straight across an- teriorly, with prominent frontal horns; nuchal organs extend pos- teriorly and terminate about setiger 4 S. bombyx Spiophanes bombyx (Claparede, 1870) Spio bombyx Claparede, 1870, p. 485. Spio crenaticornis: Giard, 1881, p. 600. Not Montagu, 1813. Fide Mesnil, 1896, p. 249; Foster, 1971, p. 41. Spiophanes verrilli Webster and Benedict, 1884, p. 728. Fide Pettibone, 1962, p. 85; Foster, 1971, p. 41. Spiophanes bombyx: Mesnil, 1896, p. 249; Fauvel, 1927, p. 41; Soderstrom, 1920, p. 243; Okuda, 1937, p. 222; Day, 1967, p. 474; Foster, 1971, p. 41 (Synonymy); Light, 1977, p. 80. Material examined: QUEENSLAND. Moreton Bay, Middle Banks, Sta. 5A, Jan. 1976, coll. W. Stephen- son (2, QM G11594J. VICTORIA. Gippsland, Mc- Gaurans Beach, 9-18 m, sandy with shell fragments and silt, June, Aug. 1977, coll. J. E. Watson (2, NMV G305O). Description: Prostomium with 2 prominent frontal horns, lacking occipital cirrus. Neuro- podia of setiger 1 with 1-2 large curved spines. Hooded hooks tridentate, beginning on se- tiger 15-16. Setiger 5-15 with parapodial thread glands, from which supernumerary setae extend through an interramal opening. Pygi- dium with 2 anal cirri. Remarks: Spiophanes bombyx is the best known species of the genus, occurring through- out the world in sand substrata. Our specimens agree well with the recent description by Fos- ter (1971). Day (1967) described 5. bombyx from South Africa as having an occipital ten- tacle, but subsequently reported this to be in error (Day, 1973). Distribution: Queensland; Victoria; cosmopoli- tan. Spiophanes wigleyi Pettibone, 1962 (Figure 26) Spiophanes wigleyi Pettibone, 1962, p. 83; Hartman 1965a, p. 147; Foster, 1971, p. 43. Material examined: VICTORIA. Bass Strait ESSO-GIPPS Sta. 9, sand, 148° 35'S-38° 8'E. (1, AM W13016); Sta. 17, 419 m, sand, 148° 34'S-38°' 59'E (3, AM W13017); Sta. 20, 93.6 m, sand, 148° 24' 50"S-39° 00' 02"E (1, AM W13018). SOUTHEASTERN AUSTRALIAN SPIONIDAE 225 Figure 26 — Spiophanes wigleyi Pettibone — a, an- terior end, dorsal view; b, hooded hook; c, right setiger 3, anterior view. [Scale 1 = 200 Am; 2 = 20 M Description: A small species up to 10 mm long for about 40 segments. Prostomium rounded anteriorly, lacking frontal horns and occipital tentacle (Fig. 26a). Caruncle absent; usually 2 pairs of eyes, the anterior pair cup-shaped, posterior pair oval; a pair of recurved dorsal sense organs extending to setiger 4, then turn- ing forward and returning to setiger 1 forming W-shaped pattern (Fig. 26a). Setiger 1 well- developed but with shorter dorsal cirri than 2-4; notopodia of setigers 1-4 basally broad, tapering distally (Fig. 26c), neuropodia broadly triangular, longest on setigers 2-3; parapodia reduced from setiger 5, neuropodia pad-like; notopodia shorter. Anterior noto- and neurosetae capillaries with granulations arranged in 2 tiers, neuro- setae reduced to 1 tier by setiger 9; tridentate neuropodial hooks with reduced hood (Fig. 26b) from setiger 15-16. Dorsal segmental ridges or crests present from setiger 15, these prominent on setigers 18-30 thereafter inconspicuous. Setiger 5-14 with parapodial thread glands bearing bacillary setae. Genital pouches absent. Pygidium with 4-6 anal cirri. Remarks: Specimens agree well with the orig- inal description of S. wigleyi. The apparent dis- junct range of this species is probably a reflec- tion of the lack of samples along continental shelf margins. Distribution: Victoria; North America, New England [Type locality] to Gulf of Mexico. Spiophanes cf. kroeyeri Grube, 1860 (Figure 27) Spiophanes krdyeri Grube, 1860, p. 88; Pettibone, 1962, p. 85 (Synonymy). Spiophanes kroeyeri: Fauchald, 1972, p. 29; Light, 1977, p. 79 (Synonymy). Material examined: QUEENSLAND. Moreton Bay, Middle Banks, Sta. 4, Sept. 1976, coll. W. Stephen- son (11, NMV G2849-2850). NEW SOUTH WALES. Sydney, Malabar, AMSBS Sta. B3 SI (12, AM W6499); Same, Sta. CA + 5 (AM W6500); Newcastle, Burwood Beach, HDWBS Station, April 1975 (1. AM W8947). VICTORIA. Bass Strait, ESSO-GIPPS Stations 11, 738 m, 148° 39'S-38° 18' 20"E (1, AM 226 JAMES A. BLAKE and JERRY D. KUDENOV WI3019); Sta. 13, 1440 m, ooze, 148° 40' 10"S-38° 26' 26"E (I, AM W J 3020); Sla. 20, 93-6 m, sand, 148° 24' 50"S-39° 00' 02"F, (AM W13021). Description: A moderate-sized species up to 22 mm long and 2 mm wide for 47 segments. Body generally opaque white in alcohol with dark brown parapodial glands on setigers 6-12. Body flattened, thereafter narrower, cylindrical. Prostomium bell-shaped, with occipital ten- tacle, caruncle extending to setiger 1-2. Eyes variable, usually 2 pairs, anterior pair rounded, posterior pair with several distinct ocelli (Fig. 27a). Peristomium well-developed, but not en- closing prostomium. Narrow nuchal ciliary tracts extending posteriorly along dorsum be- hind caruncle. Setiger 1 with finger-like, pointed neuropo- dial lamella (Fig. 27b); ventral lamellae of seti- gers 2-4 rounded, elongate; neuropodia there- after reduced to low ridge (Fig. 27e); noto- podia of setigers 1-4 cirriform, thereafter re- duced, with broad base and narrow finely ta- pered cirrus (Fig. 27c-e). Setiger 1 with large, curved non-granulated spine (Fig. 27b). Granulated inferior sabre seta appearing on setiger 4, 1 per fascicle, continu- ing throughout body (Fig. 27g); notosetae of setigers 1-4 in 3 tiers, setae of first tier with granulations, rest non-granulated, all unilirn- bate; notosetae of setiger 5 and subsequent setigers fewer, shorter; capillary neurosetae of setigers 1-4 smooth, these replaced over follow- ing setigers by broadly sheathed pointed setae with granulated shafts (Fig. 27f), numbering up to 12-15 per fascicle by setiger 9 (Fig. 27c); all capillaries replaced by setiger 15 with 6-7 quadridentate hooks (Fig, 27d) without hoods (Fig. 27h); hooks increasing to 11-12 per ra- mus in posterior neuropodia. Dark glands present between noto- and neuropodia on setigers 6-12, each gland with a cleft (Fig. 27c); no bacillary setae (see Re- marks). Genital pouches from setiger 15 oc- curring between successive parapodia for vari- able number of segments; dorsal crests with cilia from setiger 18 (Fig. 27e). Pygidium cylindrical, surrounded by 8 thin cirri. Remarks: The present specimens are closely re- lated to S. kroeyeri Grube, 1860 known from cosmopolitan cold water areas (Light, 1977). They differ somewhat in having a bell- shaped instead of subtriangular prostomium, and by lacking, instead of having, bacil- lary setae emerging from parapodial thread glands. Although bacillary setae are often dif- ficult to observe, we did examine each speci- men carefully, and in no case were the super- numerary setae detected. Several segments were removed and squashed under coverslips, but in no case were threads or bacillary setae ob- served. The glandular material appeared as clumps of globular cells suggestive of mu- cous glands rather than thread glands. We ten- tatively refer our material to S. kroeyeri. General Remarks on the Genera Spio and Microspio The separation of the genera Spio and Mic- rospio has been vague in the previous litera- ture. Microspio is currently considered a sub- genus of Spio (see Foster, 1971). The bran- chiae begin on setiger 1 in Spio and on setiger 2 in Microspio. However, some species of Spio, have very reduced branchiae on setiger 1 and have been considered by some authors as species of Microspio (see Foster, 1971 for re- view). We consider that the first occurrence of the branchiae is absolute, regardless of size, and concur with Foster (1971), but prefer to raise both taxa to full generic rank. Species of both Spio and Microspio are nu- merous and in need of review (Holmquist, 1967; Foster, 1971). Our studies have con- firmed that the dorsal sense organs of the Aus- tralian species are important to species con- cepts, provided that specimens are well-pre- served. Such an approach was previously utilized by Soderstrom (1920). Any revision of Spio and Microspio should thus consider dorsal sense organs at the species level. Genus Spio Fabricius, 1785 Spio Fabricius, 1785. Type-species: Nereis filicornis Miiller, 1776, designated by Soderstrom, 1920. Paraspio Czerniavsky, 1881. Type-species: Spio de- coratus Bobretzky, 1870, by monotypy. Diagnosis: Prostomium anteriorly rounded or incised, without frontal horns (although may be anteriorly inflated); eyes present or absent. SOUTHEASTERN AUSTRALIAN SPIONIDAE 227 1 5 . 3 b f a L e \{ Figure 27 — Spiophanes cf. kroeyeri Grube — a, an- terior end, dorsal view; b, left setiger 1, anterior view; c, left setiger 9, an- terior view; d, right setiger 15, anterior view; e, left setiger 21, anterior view; f, broad sheathed neuroseta from setiger 9; g, sabre seta, setiger 15; h, quadridentate hook. [Scale 1 = 200 Mm; 2 = 200 Am; 3 = 20 Mm] Branchiae from setiger 1, continuing through- out body, fully developed or greatly reduced at first, increasing in length thereafter, often partially fused to dorsal lamellae in anterior, and free in posterior setigers. Notosetae capil- laries only; neurosetae including capillaries, hooded hooks and sabre setae. Pygidium with anal cirri. KEY TO SPECIES OF SPIO FROM AUSTRALIA 1 a. Prostomium incised on anterior margin S. mesnili b. Prostomium entire on anterior margin . . 2 2a. Branchiae reduced on setiger 1, fully developed by setiger 2 (Fig. 29a); hooded hooks multidentate (Fig. 29g) from setiger 15-20. . . .S. singularis 228 JAMES A. BLAKE and JERRY D. KUDENOV b. Branchiae fully developed on setiger I (Fig. 28a); hooded hooks triden- tate (Fig. 28i) from setiger 9-11 S. pacifica Spio mesnili Augener, 1914 (Figure 281) Spio mesnili Augener, 1914, p. 10. Material examined: WEST AUSTRALIA. Sharks Bay, Surf Point, outer bar, 16 June 1905, S.W. Australia Exped. 1905, Sta. 25 (1, TYPE, ZMH V-8222). Description of Holotype: The type specimen is a poorly preserved anterior fragment approxi- mately 4 mm long and 1-1 mm wide for 17 setigers. Body elliptical in cross section. Colour in alcohol: opaque white. Prostomium anteriorly incised; no eyes; oc- cipital tentacle lacking; nuchal organs obscured by poor preservation; peristomium poorly-de- veloped; proboscis a flattened funnel. Setiger 1 reduced dorsally, with notosetae; notopodial lamellae fused with branchiae; neu- ropodial lamellae small, elliptical; parapodia of setiger 2 slightly displaced dorsally, inter- mediately between those of setigers 1 and 3, subsequent segments similar to setiger 3. Anterior setae sheathed capillaries, arranged in 2 tiers in both rami, first tier with short, granulated capillaries in all parapodia, second tier with non-granulated capillaries in all no- topodia and in neuropodia of setigers 1-10; tridentate hooded hooks with secondary hoods first appearing in neuropodia of setiger 11, these hooks bearing main fang overlain by elongate secondary tooth and a third small apical tooth, poorly-defined (Fig. 281). Setal distribution as follows: notopodia of setigers 4-14 with 9-10 capillaries in first tier and 10- 12 in second tier; neurosetae of setiger 4 as in notopodia; neurosetae from setiger 14 with 7-8 granulated capillaries in first tier and 6 hooded hooks in second tier, plus 3 inferiorly located granulated capillaries. Branchiae from setiger 1 reduced, i as long as on setiger 2, branchiae arching dorsally, reaching midline at about setiger 10-15, bases connected by transverse dorsal ciliated ridge continuous with ciliated inner gill margin. Pygidium absent on type, but according to Augener (1914) it consists of 2-4 anal cirri. Remarks: The new description of the type as presented above, generally agrees with the original description by Augener (1914, pp. 10-12). He notes, however, that 2 pairs of eyes were present and arranged in a square. The absence of those eyes on the type today, probably represents fading after 60 years in alcohol. The type specimen of S. mesnili is from sta- tion 25 (S.W. Australian Expedition of 1905) and was noted by Augener (1914) to differ from other specimens from stations 1 and 3. The latter specimens were said to bear bran- chiae on setiger 1 which were as long as those on setiger 2, they apparently lacked granu- lated notosetae, and the tridentate hooded hooks began on setiger 14 and numbered only 3-4 per fascicle. In contrast, the type specimen has branchiae on setiger 1 which are only i as long as on setiger 2, the anterior notosetae are granulated and hooded hooks begin on setiger 11 and number up to 7-8 per fascicle. The specimens from stations 1 and 3 possibly represent a separate species, but final con- firmation and possible identification must await either the location of those specimens or the collection of new materials from Sharks Bay. Spio mesnili bears some similarity to S. pacifica (see below), but differs in having an incised prostomium and hooded hooks with secondary hoods and in lacking granulated sheaths in anterior capillaries. The presence or absence of inferior sabre setae in 5. mesnili is not known. Distribution: West Australia, Sharks Bay. Spio pacifica sp. nov. (Figure 28a-k) Material examined: QUEENSLAND. Moreton Bay, Middle Banks, Sta. 6E, May 1976, coll. W. Stephen- son (1, QM Gl 1590). NEW SOUTH WALES. Botany Bay, Towra Point, coll. NSWSF, April 1973, Sta. 211 (HOLOTYPE, NMV G2939 and 27 PARATOPES, NMV G2940); 107 PARATYPES from Botany Bay as follows: Sta. 213 (3); 332 (11); 337 (16); 340 (11) (NMV G2941-2944); 203 (3); 207 (2); 215 (4); 218 (6); 320 (5); 324 (4); 325 (10); 329 (13); 331 (11); 344 (8) (AM W13022-13031); Stockton Beach, HDWBS Station, Nov. 1975, (1, AM W8899). VIC- SOUTHEASTERN AUSTRALIAN SPIONIDAE 229 Figure 28 — Spio pacifica sp. nov. — a, anterior end, dorsal view; b, anterior end, lateral view; c, right setiger 4, anterior view; d, right setiger 11, anterior view; e, right setiger 22, anterior view; f, right setiger 36, an- terior view; g, pygidium, dorsal view; h, anterior capillary seta; i, tridentate hooded hook; j, inferior sabre seta; k, detail of j [not to scale] — Spio mesnili Augener — 1, hooded hook. [Scale 1 = 100 Mm; 2 = 200 Mm; 3 = 20 Mm; 4 = 50 Mm; 5 = 20 Mm] TORIA. Westernport, Crib Point, CPBS Station 32S Dec. 1969 (1, NMV G2896); Westernport, WPBES Stations 1722 (2, NMV G2929-2930); 1724 (4, NMV G2927-2928); 1732 (1, NMV G2931). Port Phillip Bay, PPBES Stations 123/3 (1, NMV G2938); 920/5 (1, NMV G2934); 945 (2, NMV G2932-2933); 967 (5, NMV G2935-2937). Description: A small species, up to 10 mm long and 1 mm wide for 50 setigers. Body widest an- teriorly, tapering gradually posteriorly. Speci- mens heavily pigmented anteriorly with brown pigment on prostomium, peristomium, dorsum, ventrum and intersegmental regions (Fig. 28 a-b); brown transverse band mid- way along gill filaments (Fig. 28b) and paired mid- ventral spots on setigers 2-10; no pigment in middle and posterior setigers. Prostomium rounded on anterior margin, with a median furrow (Fig. 28a). Caruncle divides into 2 lobes surrounded laterally by 230 JAMES A. BLAKE and JERRY D. KUDENOV paired, curved ciliated nuchal organs extend- ing to middle of setiger 3. Eyes 2 pair; an- terior pair largest. Palps extend posteriorly to setiger 8-10. Peristomium moderately de- veloped to form lateral wings. Setiger 1 reduced, notopodial lamellae small, elliptical, dorsolateral^ positioned; neu- ropodial lamellae 2-3 times larger, subquad- rate (Fig. 28a-b); notopodial lamellae of an- terior setigers elliptical to subquadrate (Fig. 28c-d); those of middle and posterior setigers elongate, rounded (Fig. 28e-f); neuropodial la- mellae rounded (Fig. 28c-f), with presetal no- topodial lobes in setigers 1-17 (Fig. 28c-d). Anterior notosetae and neurosetae sheathed capillaries, granulated and non-granulated, ar- ranged in 2 tiers, notopodia with additional 3 long smooth superior capillaries; capillaries of anterior tier golden with large granules in setigers 1-6 (Fig. 28h); granulations becoming small, and setae appearing light brown by seti- ger 11; granulations only distal from about setiger 22 and completely gone by setiger 36; anterior neurosetae include sheathed granu- lated and non-granulated capillaries arranged in 2 tiers; inferior sabre setae from setigers 1 1-33, these sabre setae unique, granulated, dis- tally falcate with partial hood formed by exten- sion of sheath (Fig. 28j-k); neuropodial tri- dentate hooded hooks first present from seti- ger 9-11, with third tooth inconspicuous, then becoming more pronounced in posterior seti- gers (Fig. 28i). Margin of hood aperture smooth. Branchiae beginning setiger 1, continuing to end of body; well-developed, fused with dorsal lamellae in anterior segments, free in middle and posterior segments, ciliated on both inner and outer margins, connected by transverse ciliated bands across dorsum; ciliated band of setiger 2 interrupted by paired nuchal organs (Fig. 28a). Pygidium terminal, with 2 pairs of thick anal cirri (Fig. 28g), ventral pair H times longer and wider than dorsal pair; pygidium equal in length to last 5-6 setigers. Remarks: Spio pacifica and Spio pettlboneae Foster, 1971 have rounded prostomia, large branchiae on setiger 1 and tridentate hooded hooks. 5. pacifica differs in having presetal notopodial lobes in the anterior segments, an inconspicuous third tooth on the hooded hooks instead of a prominent one, acicular and fal- cate sabre setae with distal partial hoods and a different pygidial structure. Distribution: Queensland; New South Wales; Victoria. Spio singularis sp. nov. (Figure 29) Material examined: QUEENSLAND. Moreton Bay, Middle Banks, Sta. 57, Dec. 1972, coll. W. Stephen- son (HOLOTYPE, NMV G2884); Sta. 5\, Dec. 1973 (PARATYPE, NMV G2945): Sta. 48, Dec. 1972 (PARATYPE, NMV G2946): Sta. 29, Sept. 1972 (3 PARATYPES, QM G11595); Sta. 28, Dec. 1973 (PARATYPE, QM GU596). Description: Only incomplete specimens avail- able. A small species, up to 10 mm long and 2 mm wide for 30 setigers. Anterior body very wide, subrectangular in cross section. Body brownish red with brown pigment patches on prostomium, and brown interramal spots (Fig. 29 a-b). Prostomium bluntly rounded, with raised in- flated boss at level of setiger 1 (Fig. 29a); car- uncle extending posteriorly in 1 or 2 descend- ing steps to middle of setiger 2; no eyes; peris- tomium reduced, forming slight lateral wings; nuchal organs paired, extending to anterior margin setiger 3. Setiger 1 reduced, with notosetae, notopodia shifted dorsally, notopodial lamellae elliptical, becoming subtriangular in middle segments (Fig. 29c-e); neuropodial lamellae elliptical to broadly rounded (Fig. 29c-e). Notosetae sheathed capillaries, arranged in 2 tiers, those of anterior setigers granulated, becoming non- granulated posteriorly; neurosetae similarly arranged in 2 tiers; sabre setae first present from about setiger 13; hooded hooks from setiger 15, numbering up to 6 per neuropo- dium, appearing tridentate in lateral view (Fig. 29f); but with 4-5 secondary teeth when seen in frontal view (Fig. 29g). Branchiae short on setiger 1, reaching full- size by setiger 2, but gills narrower than those of setiger 3, branchial ciliation present on inner gill margin, continuous with dorsal trans- verse ciliary bands on body wall (Fig. 29a). Pygidium unknown. SOUTHEASTERN AUSTRALIAN SPIONIDAE 231 Figure 29 — Spio singularis sp. nov. — a, anterior end, dorsal view; b, anterior end, lateral view; c, right setiger 6, anterior view; d, right setiger 13, anterior view; e, right setiger 27, anterior view; f, quinquedental hooded hook, lateral view; g, hexadentate hooded hook, hood omitted [not to scale]. [Scale 1 = 100 /^m; 2 = 20 Mm] 232 JAMES A. BLAKE and JERRY D. KUDENOV Remarks: The presence of hooded hooks bear- ing 4 and 5 apical teeth is unique for the genus Spio. Distribution: Queensland, Moreton Bay. Genus Microspio Mcsnil, 1896 Microspio Mesnil, 1896. Type-species: Spio meczniko- wianus Claparede, 1869, designated by Soder- strom, 1920. Mesospio Gravier, 1911. Type-species: Mesospio moo- rei Gravier, 1911, by monotypy. Fide Foster, 1971. Diagnosis: Prostomium rounded to bilobed on anterior margin, without frontal horns; eyes present or absent. Occipital cirrus present or absent. Branchiae beginning on setiger 2. Ca- pillary notosetae only; neurosetae including ca- pillaries, hooded hooks and sabre setae. Py- gidium with anal cirri. Microspio granulata sp. nov. (Figures 30^31) Material examined: NEW SOUTH WALES. Botany Bay, Towra Point, NSWSF Stations 329, April 1973 (HOLOTYPE, NMV G2947), 6 PARATYPES as follows: Sta. 211 (1, NMV G2950); Sta. 311 (1, NMV G2951); Sta. 325 (1, NMV G2948); Sta. 338 (1, AM W13049); Sta. 342 (1, AM W13050); Sta. 335(1, AM W13032). Description: Holotype and paratypes incom- plete, up to 10 mm long and 15 mm wide for 31 setigers. Body very wide anteriorly, el- liptical in cross section; intensely pigmented with brown patches on prostomium, peristo- mium and branchiae, brown paired rectangu- lar patches on dorsum continuous laterally and ventrally (Fig. 30a-b); a dorsomedial strip lacking pigment in anterior 6-7 segments (Fig. 30a); from setiger 8, segmentally paired me- dial pigment areas, occurring on each side of transverse ciliary band. Prostomium bilobed, deeply incised on an- terior margin with medial furrow continuing on frontal lobe, caruncle bluntly terminating at middle of setiger 2, posteriorly rounded, with prominent occipital papilla (Fig. 30a-b); 2 pairs of eyes, arranged trapezoidally, anterior pair the larger. With transverse hood reminis- cent of dorsal collar seen in genus Streblospio present posterior to prostomium, surrounded laterally and posteriorly by curved nuchal grooves. Peristomium weakly developed. Palps extend posteriorly 6-7 setigers. Proboscis an eversible sac. Setiger 1 reduced, with small digitiform no- topodial lamellae shifted dorsally, lacking notosetae; postsetal neuropodial lamellae re- duced, inserted ventrolaterally (Fig. 30a-b); an- terior notopodial lamellae broadly to narrowly rounded, partially fused to branchiae (Fig. 30 c-e); anterior neuropodial lamellae broadly rounded, reaching maximum size at about seti- ger 20. Notosetae include sheathed capillaries ar- ranged in 2 tiers; notopodia with 20-30 capil- laries in main fascicles plus 2-3 long superior capillaries; anterior tiers of setigers 2-5 with granulated capillaries, these granulations lost in subsequent setigers, only non-granulated ca- pillaries remaining in posterior setigers; pos- terior tiers with distally granulated capillaries throughout body; sheaths of setae on setigers 2-6 strongly granulated (Fig. 31a). Neurosetae of anterior setigers with about 10-1 1 capillaries in each of 2 tiers, plus 2 long, tapered, in- ferior capillaries; capillaries of anterior tiers of setigers 1-4 granulated throughout their lengths, remaining setae of both neurosetal tiers granulated only on distal half. Capillary sheaths of posterior tiers distally bristled (Fig. 31b); tridentate hooded hooks beginning from setiger 9, 8-9 hooks per fascicle (Fig. 31c-d); sabre setae first appearing in setigers 15-17, these distally mucronate, heavy and spinous (Fig.31e). Branchiae fully developed from setiger 2, continuing to end of body; with inner ciliated gill margins connected by transverse ciliated ridges on dorsum from setiger 3. Pygidium unknown. Genital pouches present from setiger 12, de- creasing in size to setiger 29 (Fig. 30d-e). Remarks: Microspio granulata is similar to M. mecznikowianus (Claparede) in lacking notosetae on setiger 1 and in bearing triden- tate hooded hooks. M. granulata differs in hav- ing an occipital cirrus and a transverse hood across setiger 2; such a hood is otherwise known only in the genus Streblospio. Distribution: New South Wales, Botany Bay. SOUTHEASTERN AUSTRALIAN SPIONIDAE 233 a b c I e Figure 30— Microspio granulata sp. nov.— a, an- General Remarks on the Polydora-complex terior end, dorsal view; b, anterior end, Tne generic arrangement of the numerous vSVSt Stigef ; 2 , Se Sior'vKt r cieS K ° f * e ^^--omplex has usually right setiger 29, anterior view. [Scale been based u P on the fir st appearance of 1 = 100 nm; 2 = 200 Mm] hooded hooks and the presence of branchiae 234 JAMES A. BLAKE and JERRY D. KUDENOV Figure 31 — Microspio granulosa sp. nov. — a, su- perior capillary notoseta; b, anterior capil- lary neuroseta, distal tip; c, tridentate hooded hook, frontal view; d, tridentate hooded hook, lacking hood, lateral view; e, inferior sabre seta. [Scale 1 = 20 Am; 2 = 20 fim] anterior or posterior to setiger 5. Five genera are normally recognized: Polydora Bosc, Boc~ cardia Carazzi, Pseudopolydora Czerniavsky, Tripolydora Woodwick and Polydorella Au- gener. A sixth genus, Paraboccardia Rainer has lately been regarded as a subgenus of Boccardia (Read, 1975). Many species, how- ever, demonstrate characteristics of one or more of these genera and the system has re- mained unstable (Woodwick, 1964; Foster, 1971; Read, 1975) with little agreement be- tween workers on which genera (or subgenera) are valid. In the present study we introduce a new generic format based on the degree of modifi- cation of setiger 5, structure of modified spines in setiger 5, first appearance of branchiae and to a lesser degree, structure and first appear- ance of hooded hooks. The polydorids are herein arranged into 6 full genera: Pseudopolydora, Polydora, Caraz- ziella nov. gen., Tripolydora, Boccardiella nov. gen. and Boccardia. Paraboccardia is a new sy- nonym of Boccardia and Polydorella is a new synonym of Pseudopolydora. Details for each of these genera and reasons for the synony- mies are discussed under each genus in the text. Table 3 is a summary of characteristics of each of the 6 genera. As seen in this ar- rangement, 3 genera have branchiae beginning on setiger 2 and 3 have branchiae beginning posterior to setiger 5. In each of these groups there is a genus bearing a close relationship to one in the other group. For example, Pseudo- polydora and Tripolydora are similar, but the first has branchiae posterior to setiger 5 while the second has branchiae from setiger 2. Both have relatively undeveloped fifth setigers and similar hooded hook structures, but a different arrangement and structure of major spines of setiger 5. In a similar manner, Polydora and Boccardiella show close relationships as do Carazziella and Boccardia. Polydora remains the most heterogeneous of the genera and it contains the greatest number of species. Poly- dora guillei Laubier and Ramos, 1974 from the Mediterranean, described from fragmentary specimens, differs from all 6 polydorid genera and all known spionids in having unhooded unidentate neuropodial hooks from setiger 15. The species probably represents a separate genus. In the present study only the genus Tripoly- dora was not encountered. Species making up the newly proposed genera Carazziella and Boccardiella are, in part, removed from other pre-existing genera, but, several new SOUTHEASTERN AUSTRALIAN SPIONIDAE 235 TABLE 3 Some Taxonomic Characteristics of the Genera of the Polydora-complex Genus Setiger 1 Setiger 5 modification Setiger 5 major spines Branchiae begin Hooded hooks beginning; no. teeth Hooded hooks Structure Pseudopolydora Normal to achaetous Slight to moderate 2 types, usually in J- or U-shaped double rows Posterior to setiger 5 Setiger 8; bidentate Secondary tooth closely applied main fang; with constriction on shaft Polydora With or without notosetae Great 1 type, with or without companion setae Posterior to setiger 5 Setigers 7-17; bidentate With prominent angle between teeth; with or without constric- tion on shaft Carazziella With or without notosetae Great 2 types in 2 rows, 1 or both may bear bristles Posterior to setiger 5 Setigers 7-14; bidentate With prominent angle between teeth; without constriction on shaft Tripolydora* Without notopodia and (notosetae Slight 1 type, with companion setae Setiger 2, present on setiger 5 Setiger 9; tridentate Secondary teeth closely applied to main fang; no constriction on shaft Boccardiella With or without notosetae Great 1 type, with companion setae Setiger 2, present or absent on setiger 5 Setiger 7; bidentate With prominent angle between teeth; without constriction on shaft Boccardia With or without notopodia and notosetae Great 2 types in 2 rows, 1 may be bristle- topped Setiger 2, absent on setiger 5 Setigers 7-8; bidentate With prominent angle between teeth; without constriction on shaft * Holotype of Tripolydora spinosa Woodwick, 1964 [Type-species] from Eniwetok was examined (USNM 254881). species are also described. Fifteen new poly- dorid species are described below from southeastern Australia. Further reviews and descriptions of additional new species from world-wide areas will be published in a sub- sequent paper (Blake, in preparation). Genus Boccardia Carazzi, 1893 emended Boccardia Carazzi, 1893. Type-species: Polydora (Leucodore) polybranchia Haswell, 1885, by mono- typy. Paraboccardia Rainer, 1973. Type-species: Paraboc- cardia syrtis Rainer, 1973, by original designation. Diagnosis: Prostomium rounded or incised, ex- tending posteriorly as caruncle. Setiger 1 with or without notosetae. Setiger 5 modified with 2 types of major spines, few capillary noto- setae, or these absent. Bidentate hooded hooks with conspicuous angle between teeth, without constriction or manubrium on shaft; first ap- pearing on setiger 7-8. Posterior notopodial spines present or absent. Branchiae beginning on setiger 2, absent from setiger 5, present thereafter and continuing for variable number of segments. Pygidium disc-like with or with- out separate lobes or reduced to small lobes or cuffs. Remarks: The genus Boccardia is herein re stricted to those species having 2 types of ma- 236 JAMES A. BLAKE and JERRY D. KUDENOV Jar spines on setiger 5. Species formerly as- signed to Boccardia on the basis of gills being present anterior to setiger 5, but having a single type of major spine on setiger 5 are here- in transferred to Boccardiella (sec below). Paraboccardia was erected by Rainer (1973) for a group of New Zealand species, in which the hooded hooks were first present on setiger 8 instead of 7; Read (1975) treated Paraboc- cardia as a subgenus of Boccardia. The dis- tribution of hooded hooks has been found to be of marginal importance in defining spionid genera, and for that reason Paraboccardia is here considered a synonym of Boccardia. Boccardia is left with 16 species, 3 of which occur in Australia. KEY TO SPECIES OF BOCCARDIA FROM SOUTHEASTERN AUSTRALIA la. One of the 2 types of major spines on setiger 5 densely bristle-topped with- out distal cavity (Fig. 32d); occipital tentacle absent 2 b. One of the 2 types of major spines on setiger 5 with distal concavity and central cone, bristles few, only around rim (Fig. 33d); occipital ten- tacle present B. chilensis 2a. Prostomium deeply incised (Fig. 33a); notosetae absent from setiger 1 B. polybranchia b. Prostomium rounded or weakly in- cised; short notosetae on setiger 1 B. proboscidea Boccardia polybranchia (Haswell, 1885) emended (Figure 32) Polydora polybranchia Haswell, 1885, p. 275; Soder- sirom, 1920, p. 256. Boccardia polybranchia: Carazzi, 1893, p. 16; Mesnil, 1896, p. 221; Fauvel, 1927, p. 58; Hartman, 1948, p. 63; Berkeley and Berkeley, 1952, p. 16; Tmajima and Hartman, 1964, p. 279; Hartman, 1966, p. 15; Day, 1967, p. 463. [Major references only]. Polydora euryhalina Hartmann-Schroder, 1960, p. 33. Fide Hartman, 1965b. Material examined: NEW SOUTH WALES. Coffs Harbour, amongst Galeolaria tubes and coralline al- gae, on rocks, in pools, 28 Feb. 1971, coll. P. Hutch- ings (2, AM W13033). VICTORIA. Kilcunda, amongst algae in tide pools, 13 Feb. 1977, coll. J. A. Blake and J. D. Kudenov fl, NMV G3011). Tasmania, Macquarie Island (5, AM W4742-4743). Description: A moderate-sized species up to 15 mm long for 80 segments. Specimens light tan in alcohol, with dark pigment along sides of prostomium and on palps. Prostomium anteriorly bifid, continuing pos- teriorly as caruncle to posterior margin of seti- ger 3 (Fig. 32a); no occipital tentacle; 2 pairs of eyes, anterior pair more widely separated than posterior pair. Palps thick extending pos- teriorly for 9-10 setigers. Setiger 1 reduced, with small notopodial lobe, without notosetae. Capillary neurosetae short; setigers 2, 3, 4, -, 6 and succeeding seg- ments with prominent fascicles of sheathed, unilimbate capillary notosetae arranged in 3 tiers: first tier with short, thick setae, the sec- ond gradually elongating, setae of third tier longest and thinnest. Number of capillary setae gradually reduced along body, with only a few long pointed setae remaining, in posterior seg- ments; without posterior notopodial spines. Neuropodia of setigers 2, 3, 4, - and 6 bearing fascicles of unilimbate capillary setae. Biden- tate hooded hooks beginning setiger 7; hooks numbering 7-8 at first with 3-4 capillaries, in- creasing to 12-13 in middle body segments, the capillaries dropping out; hooks exhibiting wide angle between teeth; hood bearing striae and serrations at hood opening (Fig. 32b). Setiger 5 with ventral fascicle of unilimbate capillary setae; major spines of 2 types: (1) simple falcate with weak subdistal concavity along one edge (Fig. 32c); (2) bristle-topped, with tip laterally produced into smooth boss (Fig. 32d-e). Branchiae on setigers 2, 3, 4, -, 6 and suc- ceeding segments, but absent from posterior i of body; branchiae longest from setigers 8-9, rarely overlapping at midline. Pygidium with 4 equal lobes (Fig. 32f). Remarks: The original description of B. poly- branchia by Haswell (1885) (as Polydora) was brief and lacked figures. There have been no reports of the species from Australia since the original description. The type locality has changed drastically since 1885 and oyster grow- ing centres, once so numerous along the Hun- ter River are essentially gone due to industrial SOUTHEASTERN AUSTRALIAN SPIONIDAE 237 Figure 32 — Boccardia polybranchia (Haswell) — a, anterior end, dorsal view; b, hooded hook; c, simple spine from setiger 5; d-e, bristle- topped spines from setiger 5; f, pygidium. [Scale 1 = 200 wn; 2 = 20 /*m; 3 = 20 ™] development and pollution. Carazzi (1893) re- described the species based on materials from Naples. Subsequent authors have adapted Carazzi's definition of B. polybranchia and the species as such has been reported from many regions of the world. New collections have been acquired in the benthos in the Hunter River area from the one remaining oyster farm, as well as from adja- cent areas. In no case was a specimen taken which agreed with the current definition of B. polybranchia, Boccardia chilensis (see below), however, was abundant and proved to be the only species of Boccardia. Collections from Coif's Harbour, 325 km north of the Hunter River entrance, contained 2 specimens which fit B. polybranchia. Another specimen from Kilcunda, Victoria also proved to be B. poly- branchia. Because these specimens agree so well with the more cosmopolitan concept of B. polybranchia, they are here accepted as 238 JAMES A. BLAKE and JERRY D. KUDENOV representing that species, even though not found at the type locality. It is possible that B. chilensis may actually be what Haswcll orig- inally described, but because of inadequate des- criptions, the lack of type material and altera- tion of the type locality, that can probably never be definitely ascertained. The synonyms listed above represent a few of the many citations of B. polybranchia in the literature. A more complete review of this lit- erature and a comparison of the Coff's Harbour material with B. polybranchia from South America and elsewhere is the subject of an- other paper (Blake, in preparation). Distribution; New South Wales; Victoria; Tas- mania, Macquarie Island; cosmopolitan. Boccardia proboscidea Hartman, 1940 (Figure 33a-c) Polydora califomica Treadwell, 1914, p. 203. HOMO- NYM. Boccardia proboscidea Hartman, 1940, p. 382; 1941, p. 299; 1944, p. 259; 1961, p. 28; 1969, p. 95; Hart- man and Reish, 1950, p. 27; Berkeley and Berkeley, 1950, p. 51; 1952, p. 17; Woodwick, 1963, p. 132; 1977, p. 347; Imajima and Hartman, 1964, p. 279; Fauchald, 1977b, p. 47; [Not Carrasco, 1974, p. 186; 1976, p. 8 = B. tricuspa (Hartman)]. Material examined: VICTORIA. Port Phillip Bay, Werribee, MSG Monitoring Program (P609), coll. J. D. Kudenov, 11 Jan. 1977, Station 9 (919, NMV G3012-3016), numerous additional specimens with egg capsules, used for live study, many not pre- served, coll. J. A. Blake and J. D. Kudenov, 17 Feb., 3 Mar. 1977 (NMV G2851, 2854). Description: Prostomium rounded to weakly incised on anterior margin; caruncle to end of setiger 3; 4 eyes. Notosetae present on setiger 1. Spines of setiger 5 of 2 types: (1) falcate, simple (Fig. 33b); (2) bristle- topped (Fig. 33a). Bidentate hooded hooks beginning setiger 7 (Fig. 33c). Branchiae occurring on setigers 2, 3, 4, -, 6 and succeeding segments, absent from posterior i of body. Pygidium with 4 lobes, dorsal pair smaller than ventral. Remarks: Some specimens tend to show a faint incision on the prostomium, but the spe- cies agrees in all other respects with B. pro- boscidea from California. The colouration in life is a pale green on the body with 2 long black lines along the sides of the prostomium, exactly the same as those we have observed on live specimens from California. The presence of B. proboscidea in Port Phil- lip Bay is the first record from the southern hemisphere. Carrasco (1974; 1976) reported it from Chile, but his descriptions match those of B. tricuspa (Hartman), and his specimens should probably be referred to the latter spe- cies. Ecology: At the sewage outfall drains at Wer- ribee, B. proboscidea is the numerically dom- inant polychaete at some stations (Kudenov, unpublished). Johnson (1970) determined that B. proboscidea was opportunistic on sand flats in Tomales Bay, California, where it often dominates. Distribution: Victoria, North America, Califor- nia, Oregon; Panama; Japan. Boccardia chilensis Blake and Woodwick, 1971 (Figure 33d-e) Polydora polybranchia: Fauvel, 1916, p. 441. [Not Haswell, 1885], Boccardia sp. Hartman, 1948, p. 109. Boccardia chilensis. Blake and Woodwick, 1971, p. 36; Read, 1975, p. 398; Carrasco, 1976, p. 11. Boccardia jubata Rainer, 1973, p. 547. Fide Read, 1975. Material examined: NEW SOUTH WALES. Hun- ter River, Fullerton Creek, dredged, shell bottom, 16 Feb. 1977, coll. M. Skeel (65+, NMV G3017): Camden Haven, Evans Oyster lease, bottom sample, 20 April 1977, coll. M. Skeel (1, NMV G3018); Same, Kennedy-Googley's lease, on oysters, 20 April 1977, coll. M. Skeel (6, NMV G3019); Nambucca, Diemars Lease, in, on oysters, 26 April 1977, coll. M. Skeel (3, NMV G3020); MacLeary River, Stew- arts Point, Sheppards Lease, from oysters, 5 May 1977, coll. M. Skeel (10, NMV G3021); Port Ste- phens, North Bay arm, Phillips Lease, mud, 5 May 1977, coll. M. Skeel, (4, NMV G3022); Port Phillips, Swan Bay, oysters, 28 April 1977, coll. M. Skeel (4, NMV G3023); Port Stephens, Tilligery Creek, 28 April 1977, coll. M. Skeel (10, NMV G3024); Syd- ney, Vault Point, Drumoyne, inside terminal spire whorls of Velacumantes australis, 7 May 1975, coll. J. Walker (1, AM W7133); Paramatta River, Road Point, rock pool covered with Ulva, 27 Jan. 1971, coll. R. McCIoskie (3, AM W4523); Minnie Waters, offshore reef, under boulders in gravel, crevices, sandy-silt, 25 Feb. 1971, J. Holloway and P. Hutch- ings (2, AM W13034). VICTORIA. Port Phillip Bay, St. Kilda Marina, amongst Galcolaria tubes, 6 April 1977, coll. J. A. Blake and J. D. Kudenov (18, NMV G2853); Werribee, Station 14, MPSG Monitoring SOUTHEASTERN AUSTRALIAN SPIONIDAE 239 Program (P609), coll. J. D. Kudenov, 14 Feb. 1977 (38, NMV G3025-3027), numerous additional speci- mens with egg capsules, used for live study, not preserved, coll. J. A. Blake and J. D. Kudenov, 17 Feb., 3 Mar. 1977. TASMANIA. Macquarie Island, among coralline algae, Green Gorge, in rock pool, Jan. 1964 (7, NMV G2851; several in coralline algae, NMV G3028). Description: Prostomium deeply incised on an- terior margin; occipital tentacle present; car- uncle to posterior end of setiger 2 with ad- ditional mid-dorsal swelling on setigers 5-8. Setiger 1 with long notosetae. Setiger 5 with about 4 simple falcate spines and 4 spines with distal concavity and central cone (Fig. 33d); bidentate hooded hooks beginning on setiger 7 (Fig. 33e). Branchiae occurring on setigers 2, 3, 4, -, 6 and succeeding setigers. Pygidium a fleshy pad, weakly divided. Remarks: Australian specimens agree well with descriptions from Chile (Blake and Woodwick, (1971) and New Zealand (Rainer, 1973 as B. jubata; Read, 1975). Minor differences be- tween Chilean specimens and those from New Zealand were pointed out by Read (1975), but are taxonomically insignificant. We note, how- ever, that both New Zealand and Australian specimens have long capillary notosetae in setiger 1 whereas the Chilean specimens were not observed to have such long setae. Length of the notosetae on some specimens is reminis- cent of B. columbiana from the Pacific coast of North America. The lack of such setae in the Chilean material may be actual or merely re- flect the treatment and age of material ex- amined by Blake and Woodwick (1971) which was part of the Lund University Chile Expedi- tion of 1948. Larvae of B. chilensis recently described by Carrasco (1976) from Chile are nearly identical to larvae we have cultured from Port Phillip Bay (Blake and Kudenov, un- published). In Australia, B. chilensis is readily differen- tiated from related species by the form of the bristle-topped spines of setiger 5 (see key). Figure 3 3 — Boccardia proboscidea Hartman — a, bris- tle-topped spine from setiger 5; b, simple spine from setiger 5; c, hooded hook — Boccardia chilensis Blake and Wood- wick — d, group of simple and cusp-topped spines from setiger 5; e, hooded hook. 240 JAMES A. BLAKE and JERRY D. KUDENOV Distribution: New South Wales; Victoria; Chile; New Zealand; Falkland Islands; Mac- quarie Island. Genus Carazziella gen. nov. Type-species: Polydora citrona Hartman, 1941. Gen- der, feminine. Diagnosis: Prostomium rounded or incised, ex- tending posteriorly as a caruncle; eyes present. Setiger 1 with or without notosetae. Setiger 5 modified with 2 types of heavy spines arranged in a double curved row: (1) first type with an expanded tip bearing cusps or bristles; (2) sec- ond type simple, falcate; both types usually bristle-topped; superior dorsal notosetae pre- sent or absent; neurosetae of setiger 5 forming a well-developed fascicle of capillaries. Biden- tate neuropodial hooded hooks beginning on setiger 7-14, with conspicuous angle between teeth, without constriction on shaft. Branchiae beginning posterior to setiger 5. Pygidium with 2-4 lobes or 4 finger-like cirri. Remarks: Species having 2 types of major spines in 2 rows in setiger 5 as adults and pre- viously referred to the genus Polydora are herein assigned to Carazziella. The species of Carazziella form a link between the genera Boccardia and Pseudopolydora. Since both types of major spines in most species of Caraz- ziella bear bristled tips, they are among the most modified in the entire Polydora-complex. They are most closely related to Boccardia spe- cies in setal morphology, but unlike the latter genus, all lack branchiae anterior to setiger 5. Seven species are herein assigned to Caraz- ziella. These are Polydora citrona Hartman, 1941 from California designated the type-spe- cies, Pseudopolydora reishi Woodwick, 1964 from Eniwetok, Polydora quadricirrata Rainer, 1973 from New Zealand and 4 newly des- cribed species below. Five additional new spe- cies, three from North America and two from South America, will be described in a subse- quent paper (Blake, in preparation). Some juvenile and post-larval Polydora are known to have 2 types of spines in setiger 5. This is considered a larval and not an adult characteristic (see Remarks under Polydora). The genus Carazziella is named in honour of Carazzi, an early Italian student of the family Spionidae. KEY TO SPECIES OF CARAZZIELLA FROM SOUTHEASTERN AUSTRALIA la. Hooded hooks beginning on setigers 7-8; branchiae and notopodia separ- ate 2 b. Hooded hooks beginning on setigers 1 1-14; branchiae and notopodia fused C. hymenobranchiata 2a. Hooded hooks beginning on setiger 7; superior dorsal fascicle with dis- tinctive fimbriated setae (Fig. 37c) C. hirsutiseta b. Hooded hooks beginning on setiger 8; superior dorsal fascicle with sim- ple, unilimbate setae (Fig. 34d) or such setae absent 3 3 a. Setiger 1 without notosetae, notopo- dium short, same length as neuropo- dium; prostomium rounded (Fig. 34a-b) C. victoriensis b. Setiger 1 with notosetae, notopodium longer than neuropodium; prostomi- um bifid (Fig. 35a) C. phillipensis Carazziella victoriensis sp. nov. (Figure 34) 'Material examined: VICTORIA. Port Phillip Bay, PPBES Stations 901/1 (3, NMV G3103); 901/3 (4, NMV G3104); 901/4 (25, NMV G3105); 904/1 (1, NMV G3106); 914/4 (3, NMV G3107); 924/3 (1, NMV G3108); 930/2 (8, NMV G3109); 930/3 (HO- LOTYPE, NMV G2855; 2 PARATYPES, NMV G2857); 940/2 (8, NMV G3110); 940/3 (14, NMV G3111); 941/1 (9, NMV G3112); 941/3 (2, NMV G3113); 943/3 (10 PARATYPES, NMV G2856); 943/5 (7, NMV G3114); Westernport, WPBES Sta- tions 1701 (8); 1703 (1); 1711 (4); 1712 (5); 1713 (7) (NMV G3 115-3 119); Paynesville, Gippsland Lakes, jetty, sand, 2 m, 7 August 1975, coll. J. D. Kudenov (350-b NMVG3120). Description: A small species, up to 5 mm long and 0-5 mm wide for 38 setigerous segments. Body generally opaque white, except for brown flecks around eyes. Prostomium flattened or rounded on an- terior margin (Fig. 34a-b); caruncle unique, extending to posterior margin of setiger 1 where it terminates, followed by a second ridge SOUTHEASTERN AUSTRALIAN SPIONIDAE 241 :.: |.A " Figure 34 — Carazziella victoriensis gen. et sp. nov. — ■ a, anterior end, dorsal view in contracted state; a', detail of caruncle [not to scale]; b, anterior end, dorsal view in relaxed state; c, hooded hook; d, setae of setiger 5, single superior dorsal seta on left, bristle- topped spines in centre, ventral fascicle to right; e, bristle-topped spine from seti- ger 5; f-g, falcate spines from setiger 5; h, posterior end in dorsal view; i, pygi- dium. on setiger 2 (Fig. 34a-b); anterior margin of ridge elevated, sometimes overlapping ter- minal end of caruncle (Fig. 34a0; no occipital tentacle, although anterior edge of ridge may approximate tentacle; 2 pairs of eyes: anterior pair cup-shaped; posterior pair oval Peristo- mium inflated extending anteriorly beyond prostomium; palps long, extending posteriorly 12-15 setigers. Setiger 1 with well-developed noto- and neu- ropodial lobes, but lacking notosetae; neuro- setae including simple fascicle of capillary setae. Setigers 2-4 with fascicles of capillary noto- and neurosetae. Parapodial lobes short, conical. Setiger 6 and succeeding segments with fascicles of long capillary notosetae. Neuro- podia of setigers 6-7 with capillary setae; bi- dentate hooded hooks from setiger 8 (Fig. 34c), these number 2-6 per fascicle and accompanied by 2-3 inferior capillary setae for most of body length. Setiger 5 distinctly modified, with heavy dor- 242 JAMES A. BLAKE and JERRY D. KUDENOV sal musculature and prominent heavy modified spines, these spines of 2 types, both with bris- tle-tops and arranged in a double curved row: (I) ventral row of spines large, distally ex- panded and bristle-topped (Fig. 34d-e); (2) dorsal row smaller, falcate, with short bristles enveloping fang distally (Fig. 34d, f-g); 1 or 2 small, unilimbate superior dorsal capillaries present above anteriormost spine (Fig. 34d), occasionally absent; with prominent fascicle of sheathed, bilimbatc capillaries ventral to major spine scries (Fig. 34d). Branchiae from setiger 7, continuing for only 7-10 setigers. Pygidium with 4 lobes, ventral pair larger than dorsal pair (Fig. 35h-i). Remarks: See comments for C. phillipensis. Distribution: Victoria. Carazziclla phillipensis sp. nov. (Figure 35) Material examined: VICTORIA. Port Phillip Bay, PPBES Stations 901/1 (2 PARATYPES, NMV G2861); 901/3 (14, NMV G3122); 901/4 (5, NMV G3123); 901/5 (PARATYPH, NMV G2859); 901/5 (2, NMV G3124); 904/2 (1, NMV G3125); 915/3 (1, NMV G3126); 926/2 (1, NMV G3127); 933/3 (2, NMV G3129); 936/5 (1, NMV G3130); 943/3 (1, NMV G3131); 972/1 (HOLOTYPE, NMV G2858); Hobsons Bay-Yarru River, MSG Station 134/4, 10 Feb. 1975, coll. G. Poore and J. D. Kudenov (3 PARATYPES, NMV G2860). Description: Similar to C. victoriensis. Body small, up to 5 mm long and 5 mm wide for 36 setigerous segments. Generally opaque white in alcohol with dusky brown on prostomium and near eyes. Prostomium anteriorly weakly incised, ap- pearing nearly round on contracted specimens (Fig. 35a); caruncle generally as in C. vic- toriensis, without occipital tentacle; 4 pairs of eyes; anterior pair distinctly cup-shaped; posterior pair oval to irregular in shape. Peris- tomium inflated as in C. victoriensis; palps ex- tending posteriorly for 8-9 setigers. Setiger 1 with well-developed fascicles of capillary noto- and neurosetae (Fig. 35a); noto- podial lobe unusually long, much longer than neuropodial lobe. Setigers 2-4 with fascicles of capillary notosetae and neurosetae; notopodial lobes broad and short, neuropodial lobes short and finger-like. Setiger 6 and succeeding seg- ments with spreading fascicles of capillary no- tosetae; neuropodia of setigers 6-7 with capil- lary setae. Bidentate hooded hooks from seti- ger 8 (Fig. 35b); these hooks numbering 5-7 per ncuropodium throughout, accompanied by several capillary setae at each end of row of hooded hooks, although in some cases only in- ferior group of capillaries present* Setiger 5 large (Fig. 35c), with 2-3 superior dorsal capillaries anterior and dorsal to major spines; spines of 2 types: (1) ventral row of large, bristle-topped spines with distal cavity (Fig. 35c); (2) dorsal spines smaller, falcate, with bristles forming prominent tuft on convex surface (ventral) of spine (Fig. 35c); setiger 5 with well-developed ventral fascicle of capillary setae. Branchiae from setiger 7, occurring for only 5-6 setigers; branchiae broad, meeting at mid- line, Pygidium with 4 subequal lobes (Fig. 35d). Remarks: Carazziella victoriensis and C. philli- pensis are closely related. C. phillipensis bears notosetae on setiger 1, the prostomium is an- teriorly incised and the ventral row of major spines on setiger 5 exhibits a prominent distal concavity. In contrast, C. victoriensis lacks no- tosetae on setiger 1, the prostomium is an- teriorly entire and the ventral row of major spines on setiger 1 are distally conical and domed, showing only a faint notch or none at all. Distribution: Victoria, Port Phillip Bay. Carazziclla hymenobranchiata sp. nov. (Figure 36) Material examined: VICTORIA. Port Phillip Bay, PPBES Stations 904 (1, NMV G3049); 906/1 (PARA- TYPE, NMV G2862); 921/3 (5 PARATYPES, NMV G2863); 944/4 (4 PARATYPES, NMV G2864); 126/6 (HOLOTYPE, NMV G2865). Description: Most specimens incomplete anter- ior fragments; holotype measuring 5 mm long for 20 setigers. A paratype measures 10 mm long and 1-0 mm wide for approximately 48 segments. Body light tan, no other body pig- ment. SOUTHEASTERN AUSTRALIAN SPIONIDAE 243 Figure 35 — Carazziella phillipensis sp. nov. — a, anterior end in dorsal view; b, hooded hook; c, group of spines from setiger 5; d, posterior end in dorsal view. [Scale 1 = 200 Aim; 2 = 20 Am] 244 JAMES A. BLAKE and JERRY D. KUDENOV Figure 36 — CarazzieUa hymenobranchiata sp. nov. — a, anterior end, dorsal view; b-d, large bris- tle-topped spines from setiger 5; e-f, curved spines from setiger 5; g, hooded hook; h, right parapodium, setiger 10, anterior view; i, pygidium. [Scale 1 = 500 Mm; 2 = 200 Mm; 3 rr 50 Mm; 4 - 20 Mm] Prostomium distinctly bifid on anterior mar- gin; caruncle not apparent, nuchal ridge merg- ing with setiger 1 immediately posterior to eyes (Fig. 36a); 2 pairs of eyes: first pair cup-shaped and widely spaced; posterior pair rounded and closely spaced. Peristomium slightly enlarged, but not inflated; palps reaching posteriorly 8-9 segments. Setiger 1 with long noto- and neuropodial lobes, but lacking notosetae (Fig. 36a); neuro- setae including spreading fascicles of capillary setae. Setigers 2-4 with spreading fascicles of unilimbate capillary noto- and neurosetae; digi- tiform notopodial lobes of setigers 2, 3, 4, - and 6 become expanded leaf-like lamellae on seti- ger 7; these extending ventrally down the seg- ment and fusing dorsally with branchiae, form- ing membranous wings (Fig. 36a, h); con- dition persisting through gill bearing middle body region, thereafter notopodial lobes re- duced to simple protuberances. Notosetae uni- limbate capillaries from setiger 6, posterior capillaries lacking sheaths. Neuropodia of seti- gers 2, 3, 4, - and 6 at first digitiform, becom- ing progressively flattened blade-like lobes; neuropodial lobe a thin lamella throughout branchial region. Neurosetae in setigers 6-10 unilimbate capillaries. First appearance of hooded hooks variable from setiger 1 1-14; hooks with reduced secondary tooth (Fig. 36g), numbering 5-6 initially, increasing to 7-8 per neuropodium in mid-body region, reduced to 5-6 in posterior region; hooks accompanied by SOUTHEASTERN AUSTRALIAN SPIONIDAE 245 equal number of interspersed slender capillary setae. Modified spines of setiger 5 arranged in semi-circle, with 7-8 large brush-like spines en- closing 3-4 falcate spines; bristle-topped spines with tall peak, seen in certain angles (Fig. 36b-d); falcate spines with fine bristles on sub- distal curved portion of shaft (Fig. 36e-f); without superior dorsal setae, but with spread- ing fascicles of ventral unilimbate capillaries. Branchiae from setiger 7, continuing through middle body segments, these fused with flat- tened notopodial lobes, as noted above (Fig. 36h)« Pygidium with 4 subequal lobes (Fig. 36i). Remarks: The fusion of branchiae and noto- podial lobes in C. hymenobranchiata are unique for this genus, as is the lack of a pos- teriorly prolonged caruncle, and the relative lack of peristomial inflation. Distribution: Victoria, Port Phillip Bay. Carazziella hirsutiseta sp. nov. (Figure 37) Material examined: NEW SOUTH WALES. Botany Bay, Towra Point, NSWSF Stations, 17 April 1973 (HOLOTYPE, NMV G2866; 6 PARATYPES, NMV G2867-2969). Description: A small species measuring 3*2 mm long and 0-5 mm wide for 34 setigerous segments. Body widest at setiger 5, tapering abruptly anteriorly and in posterior segments. Generally opaque white in alcohol with some brown pigment near eyes. Prostomium rounded on anterior margin, ap- pearing bulbous, continuing posteriorly as car- uncle to posterior border of setiger 2 (Fig. 37a); 2-3 pairs of eyes: anterior pair cup- shaped, further apart and occasionally divided into 2 pairs; posterior pair oval, closer to- gether; no occipital tentacle. Setiger 1 with long notopodial and neuro- podial lobes, but lacking notosetae (Fig. 37a); neurosetae slender, unilimbate capillaries. Setigers 2, 3, 4, -, 6 and succeeding segments with thickened notopodia and fascicles of uni- limbate capillaries arranged in 2 tiers: first tier with shorter, thicker setae, those of second tier being longer; notosetae of posterior seg- ments reduced to a few long slender capillaries. Neurosetae of setigers 2, 3, 4, - and 6 similar to notosetae, arranged in spreading 2-tiered fascicle. Bidentate hooded hooks with reduced angle between teeth (Fig. 37b) from setiger 7, with 3 hooks and 5 capillaries at first, there- after 7-9 hooks and 3-4 capillaries in middle segments, with 3 hooks and no capillaries in posterior segments. Setiger 5 setae include a dorsal group of 3-4 peculiar truncated, distally fimbriated setae (Fig. 37c), 2 types of bristle-topped spines and a ventral inferior fascicle of unilim- bate capillary neurosetae. Bristle-topped spines of 2 types: (1) a ventral row of spines distally expanded into 3 very pronounced bluntly rounded knobs enclosing a deep central con- cavity (Fig. 37f-g), 1 of these knobs greatly exceeding the other 2 in length; (2) 4-5 falcate spines in a dorsal row (Fig. 37d-e) bearing bristled tips (Fig. 37d-e). Branchiae from setiger 7, occurring only over 5-6 segments or setigers 12-13. Branchiae short, thickened, barely reaching to mid-line. Pygidium with 4 thin subequal lobes or cirri (Fig. 37h). Remarks: Carazziella hirsutiseta is the only species of the genus with hooded hooks be- ginning on setiger 7. The large tricuspid bristle-topped spines of setiger 5 are unusually pronounced at their apex and bear a large ter- minal notch or cavity. The dorsal superior fas- cicle also bears highly distinctive fimbriated setae which appear to be transitional between normal capillaries and bristle-topped major spines. Distribution: New South Wales, Botany Bay. Genus Polydora Bosc, 1802 emended Polydora Bosc, 1802. Type-species: Polydora comma Bosc, 1802. Diplotis Montagu, 1813. Type-species: Diplotis hya- lina Montagu, 1813. Leucodore Johnston, 1838. Type-species: Leucodore ciliatus Johnston, 1838. Leipoceras Mobius, 1874. Type-species: Leipoceras uvijerum Mobius, 1874. Dipolydora Verrill, 1881. Type-species: Polydora concharum Verrill, 1881. Protopolydora Czerniavsky, 1881. Type-species: Poly- dora hamata Langerhans, 1880. (HOMONYM of Polydora hamata Webster, 1879b) = [Polydora posthamata Jones, 1962]. 246 JAMES A. BLAKE and JERRY D. KUDENOV 1R WW. MmM Figure 37 — Carazziella hirsutiseta sp. nov. — a, an- terior end, dorsal view; b, hooded hook; c, superior dorsal setae, setiger 5; d-e, fal- cate bristle-topped spines, setiger 5; f-g, large bristle-topped spines, setiger 5; h, posterior end, dorsal view. [Scale 1 = 200 Mm; 2 = 100 Mm; 3 = 20 Mm; 4 = 100 Mm] Pseudoleucodore Czerniavsky, 1881. Type-species; Leucodorum caecum Oersted, 1843. Diagnosis: Prostomium anteriorly entire or in- cised, extending posteriorly as caruncle; eyes present or absent. Setiger 1 with or without notosetae. Setiger 5 greatly modified, with major spines of one type, usually accompanied by slender companion setae; spines arranged in single curved row. Posterior notopodial spines present or absent. Neuropodial hooded hooks bidentate with conspicuous angle be- tween teeth, with or without constriction on shaft and beginning on setigers 7-17. Branchiae beginning posterior to setiger 5. Pygidium re- duced or enlarged, cuff-like, saucer-like or divided into lobes. Remarks: With the erection of the genus Caraz- ziella, species with 2 types of major spines in setiger 5 as adults are removed from Polydora. This includes P. citrona Hartman, which was noted by Woodwick (1964) as one of the spe- cies not exactly conforming to the definition of Polydora. Restricting the genus Polydora to include only those species with 1 type of major spine on setiger 5 stabilizes the genus, al- though there remain a few problems with spe- cies described as juveniles. Blake (1969) pointed out that some species of larval or post- larval Polydora have an initial simple falcate SOUTHEASTERN AUSTRALIAN SPIONIDAE 247 spine among the main series of spines which develop on setiger 5. This spine eventually drops out and is replaced by the setae typical for that particular species. At least 3 species have been described from juveniles having such a setal configuration: P. heterochaeta Rioja, 1 939; P. laticephala Hartmann- Schroder, 1959 and P. punctata Hartmann- Schroder, 1959. This type of setal arrangement is considered to be a larval and not an adult characteristic (Blake, 1969); the validity of any species described from larval or post-larval forms should be seriously questioned. Polydora cornuta Bosc, 1 802 is the indeter- minable type-species of the genus (Blake, 1971; Foster, 1971). Comments concerning the identification of P. cornuta will be included in a forthcoming paper (Blake, in preparation). KEY TO SPECIES OF POLYDORA FROM SOUTHEASTERN AUSTRALIA la. Hooded hooks with constriction on shaft (Fig. 43j) 10 b. Hooded hooks without constriction on shaft (Fig. 38b) 2 2a. Major spines of setiger 5 bearing accessory structures as teeth, flanges, or bristles, or with distal cavity 6 b. Major spines of setiger 5 simple, fal- cate, without accessory structures, but may have subterminal enlarge- ment (Fig. 38a, d, e) 3 3a. Occipital tentacle present (Fig. 39a) P. tentaculata b. Occipital tentacle absent 4 4a. Posterior notopodial spines or needles present; gizzard absent, or if present, seen only upon dissection 5 b. Posterior notopodial spines absent; gizzard present internally in setigers 18-19, prominently seen externally as swelling P. socialis 5a. Posterior notopodial spines present as individual acicula among capil- laries (Fig. 38h); major spines of setiger 5 with large subterminal pro- tuberance (Fig. 38f). . . .P. protuberata R b. Posterior notopodial spines present as dense expanded packets of needles (Fig. 38c); major spines of setiger 5 with weak subterminal swelling (Fig. 38a) P. flava 6a. Major spines of setiger 5 with crest of bristles on convex side and with accessory flange, collar or cavity 8 b. Major spines of setiger 5 without crest of bristles, with 1 or 2 acces- sory teeth, with or without cowling or hood connecting teeth 7 7a. Major spines of setiger 5 with large accessory tooth (Fig. 38i); posterior notopodial spines absent; branchiae from setiger 9 P. giardi b. Major spines of setiger 5 bifid with hood or cowling connecting 2 teeth (Fig. 43a-c); posterior notopodial acicular spines present (Fig. 43g); branchiae from setiger 7 P. armata 8a. Branchiae from setiger 9; major spines of setiger 5 with lateral ac- cessory flange (Fig. 40g-h); pos- terior notopodial spines present P. aciculata b. Branchiae from setiger 7; major spines setiger 5 with subterminal concavity or collar; posterior noto- podial spines absent 9 9a. Prostomium entire, major spines of setiger 5 with subterminal concavity and crest of bristles on convex side (Fig. 41b) P. notialis b. Prostomium incised; major spines of setiger 5 with collar on convex side from which bristles emerge (Fig. 42b-c) P. pilocollaris 10a. Posterior notopodia with needle packets or large spines 11 b. Without posterior notopodial spines . . 12 11a. Packets of posterior notopodial needles (Fig. 45j-k); major spines of setiger 5 with small lateral flange (Fig. 45'c-f ) P. latispinosa b. Posterior notopodial spines indi- vidual, large, not needle-like, curved in towards mid-line (Fig. 47e); major spines of setiger 5 with large flange (Fig. 47b-c). . P. hoplura 248 JAMES A. BLAKE and JERRY D. KUDENOV 12a. Occipital tentacle present; com- hooks numbering 3-4 per ramus, and nor- panion setae of setiger 5, delicate, mally with wide angle between teeth (Fig. 38b); feathery in appearance (Fig. 43h) one specimen from Towra Beach, N.S.W. bear- \ -P. Ugni ing a reduced secondary tooth in posterior b. Occipital tentacle absent; compan- setigers. With capillary notosetae in posterior ion setae of setiger 5, bilimbate, not setigers accompanied by packets of fine needles feathery (Fig. 46g) 13 (Fig. 38c). 13a. Prostomium entire; caruncle short, Setiger 5 modified, bearing curved row of globular (Fig. 46g); pygidium smal- simple falcate spines, each with subterminal ler than posterior segments, cuff- swelling (Fig. 38a); with bilimbate companion shaped (Fig. 46k) P. woodwicki setae and 5-6 superior dorsal pointed setae; b. Prostomium incised; caruncle long, ventral fascicle of sheathed unilimbate capil- narrow (Fig. 44a); pygidium, larger laries present, than posterior segments, saucer- Branchiae from setiger 8. Pygidium a 3- like (Fig. 44g) 14 Iobed disc, with 2 small dorsal lobes and a 14a. Major spines of setiger 5 with 1 ac- larger ventral lobe. cessory flange (Fig. 43k, m) Remarks: This species agrees well with des- • p - websteri criptions from other seas. The posterior b. Major spines of setiger 5 with 1 ac- needles are arranged in a tight, flattened group cessory flange located just ahead of as in P. convexa Blake and Woodwick (1972) an accessory tooth (Fig. 44c-e) and P. latispinosa (see below). P. haswelli Distribution: New South Wales; Victoria; Europe; Ceylon; Sumatra; Japan. Polydora flava Claparede, 1870 ~ _ . Polydora socialis (Schmarda, 1861) (Figure 38a-c) . } Polydora flava Claparede, 1870, p. 487; Fauvel, 1927, (Figure 38d-e) p. 52; Day, 1967, p. 468; Hartmann-Schroder, 1971, Pol ydora socialis: Blake, 1971, p. 20 (Synonymy); p. 305. [major references only] Ll S nt > I977 < P- 71. Polydora pusilla Saint-Joseph, 1894, p. 65. Polydora plena: Foster, 1971, p. 24. Fide Light, 1977, Polydora dorsomaculata Rainer, 1973, p. 558. New p " 7I * synonymy. Material examined: NEW SOUTH WALES. Syd- Maierial examined: NEW SOUTH WALES. North ne y» Wy-ar-gine Point, Middle Harbour, in Galeo- Heads, Sydney, AMSBS, May 1972 (1, AM W6941); larta associations, 14 Dec. 1968 (1, AM W13036); Botany Bay, Towra Beach, NSWSF Station, April Malabar, Sydney, AMSBS Stations (1, AM W6496; 5, 1973 (1, AM W13035). VICTORIA. Port Phillip AM W6497); Sydney Harbour, S.E. Chinamens Bay, PPBES Stations 907 (55), 913 (1), 915 (1), 919 Beach, Malacological club survey, 8 May 1971 (1, (56), 929 (318), 930 (4), 932 (4), 944 (2), 985 (1) AM W13037); Botany Bay, Towra Beach, NSWSF (NMV G3029-3037); Westernport, Crib Point, CPBS Station, 12 April 1973 (1, AM W13038). VICTORIA. Station 32A, April 1970 (5, NMV G3048). Port Phillip Bay, PPBES Stations 907 (56); 919 (144); ~ . . , . „ 929 (318); 932 (4); 944 (2); 953 (7); 965 (4); 978 (9); Description: A moderate-sized species averag- 982 (3) (widespread in PPB) (NMV G3158-3166). ing about 8 mm long and 0-5 mm wide for Description: A moderate-sized species, up to 80 setigerous segments. 9 ^ i ong and . 75 mm wide for 66 setigers Prostomium strongly bifid on anterior mar- Prostomium strongly incised on anterior mar- gin, caruncle extending to setiger 3; no occi- gin, caruncle extending to setiger 4-5; no occi- pital tentacle; no eyes. Setiger 1 with both pital tentacle; 2 pairs of eyes. Setiger 1 with noto- and neuropodial setal fascicles; noto- notosetae. Major spines of setiger 5 with sub- podial lobes of setiger 1 long. Setigers 2, 3, 4, terminal boss (Fig. 38d). Companion setae bi- - and 6 with well-developed fascicles of limbate (Fig. 38d). No posterior notopodial sheathed, unilimbated capillaries in both rami, spines. Bidentate hooded hooks from setiger 7 Bidentate neuropodial hooded hooks com- (Fig. 38e) accompanied by capillary setae pletely replacing capillaries from setiger 7, Branchiae from setiger 8. Pygidium disc-like SOUTHEASTERN AUSTRALIAN SPIONIDAE 249 Figure 38 — Polydora ftava Claparede — a, 3 major spines and 2 companion setae, setiger 5; b, hooded hook; c, posterior notopodial needles — Polydora socialis (Schmarda) — d, 4 major spines and 2 companion setae, setiger 5; e, hooded hook — Polydora pro- tuberata nov. sp. — f, 3 major spines and 2 companion setae, setiger 5; g, hooded hook; h, posterior notopodial spine — Polydora giardi Mesnil — i, 3 major spines, setiger 5; j, companion seta, setiger 5; k, hooded hook. Gizzard present internally in setigers 18-19. Remarks: Specimens agree well with collections from other geographic areas, although they seem generally smaller. Reduction of the sec- ondary tooth in posterior hooded hooks was observed by Blake (1971), for specimens from New England; this feature was not found in the present collections. Variation in that character has been observed in other popula- tions from North America and will be dis- cussed in a subsequent paper (Blake, in pre- paration). Some specimens exhibit dorsal pig- ment flecks, which are believed to be remnants of larval pigment (Blake, 1971). The gizzard is a highly diagnostic feature of P. socialis. Lo- cated internally at about setigers 18-19, it ap- pears as a conspicuous external bulge. A simi- 250 JAMES A. BLAKE and JERRY D. KUDENOV lar structure is also present in P. protuberata and P. tentaculata (see below), but is not as conspicuous as in P. socialis and best seen upon dissection. As shown by Blake (1971), P. socialis plena Berkeley and Berkeley, 1936 is an invalid subspecies because the character on which it is based, namely the presence of noto- setae on setiger 1, occurs on all specimens of P. socialis (sensu lato). Hence, elevation of P. plena to full species rank by Foster (1971) is not supported (see also Light, 1977). Distribution: New South Wales; Victoria; New Zealand; North America; South America. Polydora protuberata sp. nov. (Figure 38f-h) Material examined: VICTORIA. Port Phillip Bay, PPBES Stations 904 (1, NMV G3049) 907 (20 PARA- TYPES, NMV G2871); 908 (2, NMV G3050); 913 (1, NMV G3051); 922 (2, NMV G3052); 928 (3, NMV G3053); 930 (1, NMV G3054); 946 (HOLO- TYPE, NMV G2870). Description: Holotype the largest specimen, broken in 2 parts, measuring 1 6 mm long and 1 mm wide for 115 segments. Body colourless. Prostomium distinctly incised on anterior margin, caruncle extending to setigers 3-5, no occipital tentacle or eyes. Setiger 1 with capillary notosetae. Setigers 2, 3, 4, -, 6 and succeeding segments with uni- limbate capillary notosetae in 3 tiers; posterior notopodia with several short acicular spines in addition to long capillaries (Fig. 38h). Neuro- setae of setigers 2, 3, 4, - and 6 with unilim- bate capillaries in 2 tiers. Four to 5 bidentate neuropodial hooded hooks from setiger 7, ac- companied by capillary setae for only a few segments; hooks with wide angle between teeth; main fang forming oblique angle with shaft; constriction lacking on shaft (Fig. 38g). Setiger 5 modified, with superior dorsal fas- cicle of geniculate setae lacking sheaths, a curved row of major spines and sheathed bi- Iimbate companion setae and a ventral fascicle of unilimbate capillaries; major spines simple, falcate, with a large subterminal protuberance (Fig. 38f), this much more pronounced than in P. socialis (Fig. 38d). Branchiae from setiger 8, continuing to near posterior end. Pygidium disc-like, with dorsal notch. Weakly-developed gizzard, best seen in dissection, observed on some specimens at about setigers 17-18. Remarks: Polydora protuberata is very similar to P. caeca (Oersted, 1843) from Europe (Fau- vel, 1927; Hartmann-Schroder, 1971) and South Africa (Day, 1967). The 2 species differ in the structure of the major spines of setiger 5. In P. caeca, the spines have a slight swelling whereas P. protuberata has a large expanded protuberance. The 2 species also differ in habi- tat. P. caeca is a borer in calcareous habitats (Fauvel, 1927) while P. protuberata lives in sediment. Distribution: Victoria, Port Phillip Bay. Polydora tentaculata sp. nov. (Figure 39) Materia! examined: QUEENSLAND. Moreton Bay, Middle Banks, Sta. 34, March 1974, coll. W. Stephen- son (2 PARATYPES, QM G11597); Sta. 56, June 1974 (PARATYPE, QM G11600). NEW SOUTH WALES. Botany Bay, Towra Beach, NSWSF Station, April 1973 (HOLOTYPE, NMV G2885; 1 PARA- TYPE, NMV G2886). Description: A moderate-sized species up to 15 mm long and 10 mm wide for 88 setiger- ous segments. Body generally opaque white with some anterior brown pigment bars located dorsally on anterior setigers. Prostomium deeply bifurcate on anterior margin, forming 2 divergent lobes (Fig. 39a). Caruncle extending to middle of setiger 5; with triangular shaped occipital tentacle present at level of setiger 1 ; eyes 0-2. Setiger 1 with notosetae and long notopo- dial lobes (Fig. 39a). Setigers 2, 3, 4, -, 6 and succeeding setigers with fascicles of notosetae arranged in 2 tiers; posterior notosetae in- cluding long slender capillaries (Fig. 39b) and a stouter spine (Fig. 39c). Neurosetae of seti- gers 2, 3, 4, - and 6 with capillaries. Bidentate hooded hooks from setiger 7, hooks numbering 5-8 throughout; without constriction on shaft; angle between teeth variable within a single fascicle, being widest in superiormost position (Fig. 39d) and less in more inferior position (Fig. 39e); hooks accompanied by inferior capillaries in first few segments. Setiger 5 modified, larger than setigers 4 or SOUTHEASTERN AUSTRALIAN SPIONIDAE 251 Figure 39 — Poly dor a tentaculata sp. nov. — a, anterior end, dorsal view; b, long posterior capil- lary notoseta; c, short acicular-like pos- terior notoseta; d, hooded hook from su- perior neuropodial position; e, hooded hook from inferior neuropodial position; f, group of 4 major spines and 4 com- panion setae, setiger 5. [Scale 1 = 500 Mm; 2 = 200 m; 3 = 20 wn] 6; setae including a superior dorsal fascicle of geniculate setae lacking sheaths, a curved row of major spines accompanied by simple bi- limbate companion setae (Fig. 39f) and a ven- tral fascicle of unilimbate capillaries. Major spines simple, falcate, lacking accessory struc- tures (Fig. 39f). Branchiae from setiger 8, small at first, reaching full-size by setiger 10, present over most of body. Pygidium unknown. Gizzard observed at level of setigers 19-20 as bulge in dorsum; appears as muscular en- largement of intestine upon dissection; not as well-developed as in P. socialis. Remarks: Polydora tentaculata is most closely related to P. socialis, P. flava, P. protuberata and P. caeca. It differs from all of these in possessing an occipital tentacle and in bearing hooded hooks in which the angle changes with- in individual fascicles from the superior to in- ferior position. In the combined feature of bearing an occipital tentacle and simple major spines on setiger 5, P. tentaculata also re- sembles P. nuchalis Woodwick (1953) from California. The latter species, however, ex- hibits a constriction on the shafts of the hooded hooks, and belongs to a group of spe- 252 JAMES A. BLAKE and JERRY D. KUDENOV cies related to P. ligni Webster (sec below). Distribution: Queensland; New South Wales. Polydora giardi Mesnil, 1896 (Figure 3 8i-k) Polydora giardi Mesnil, 1896, p. 195; Fauvel, 1927, p. 50; Hartman, 1941, p. 309; 1969, p. 135; Rainer, 1973, p. 560; Read, 1975, p. 413. Material examined: NEW SOUTH WALES. Syd- ney, Wy-ar-gine Point, Middle Harbour, from Galea- laria association, 19 Dec. 1968 (10+, AM W13039). VICTORIA. Kilcunda, Bourne Creek, in shell (Halio- tus ruber, 11 April 1977, coll. J. A. Blake and J. D. Kudenov (6, NMV G2949); Bass Strait, 20 m, 38° 28"S-144° 48'E, dredged, from bryozoans, coralline algae on sandstone reef, coll. J. A. Blake and J, D. Kudenov, May 1977 (2, NMV G3055). Description: A small species measuring up to 7 mm long and 03 mm wide for 75 setigerous segments, but reported larger in other areas (Rainer, 1973). Prostomium incised, caruncle extending to anterior margin of setiger 3; no occipital tentacle or eyes. Setiger 1 with both noto- and neurosetae, and well-developed para- podial lobes. Notopodia of setigers 2, 3, 4, -, 6 and succeeding segments with fascicles of unilimbate capillaries; with long capillaries in posterior setigers, but spines lacking. Neuro- setae of setigers 2, 3, 4, - and 6 with unilim- bate capillaries. Bidentate hooded hooks from setiger 7, 3-4 hooks at first accompanied by 1-2 capillaries; hooks with acute angle between secondary tooth and main fang and wide angle between main fang and shaft (Fig. 38k), no constriction on shaft. Setiger 5 modified with dorsal fascicle of prominent geniculate setae, lacking sheaths, a curved row of heavy spines and bilimbate companion setae (Fig. 38j) and a ventral fas- cicle of winged capillary neurosetae; major spines falcate with large accessory tooth (Fig. 38i), some specimens appearing to have an ad- ditional small spur on convex side of shaft. Branchiae from setiger 9, absent from pos- terior i of body. Pygidium a small disc. Distribution: New South Wales; Victoria; Europe; California; New Zealand. Polydora aciculata sp. nov. (Figure 40) Material examined: VICTORIA. Kilcunda, Bourne Creek, in shell Haliotus ruber, 11 April 1977, coll. J. A. Blake and J. D. Kudenov (HOLOTYPE, NMV G2872. 8 PARATYPES, NMV G2873). Description: A moderate-sized species up to 10 mm long and 3 mm wide for 85 setigers. Body generally opaque white in alcohol, with faint brown lines along margin of caruncle. Prostomium entire and narrow on anterior margin (Fig. 40a), caruncle continuing as low ridge to posterior margin of setiger 3; no occi- pital tentacle or eyes. Peristomium and pro- stomium depressed and lower in lateral view than first setiger and succeeding segments. Setiger 1 with capillary setae in both rami (Fig. 40a). Setigers 2, 3, 4, -, 6 and succeeding setigers containing spreading fascicles of uni- limbate capillary notosetae arranged in 2 tiers, first tier with shorter setae and longer setae in second; in mid-body region, shorter capillary setae replaced by 2-3 stout acicular spines (Fig. 40b-c) accompanied by 3-4 long, thin capillary setae. Notosetae maintaining this ar- rangement to end of body (Fig. 40k-l). Three to 4 bidentate hooded hooks per neuropodium from setiger 7 accompanied by a single capil- lary for a few segments; without constriction on shaft (Fig. 40d); hooks with wide angle between secondary tooth and main fang; this angle gradually diminishing along body, sec- ondary tooth and hood gradually becoming smaller and more closely pressed to main fang (Fig. 40e), both eventually disappearing in far posterior segments (Fig. 40f), leaving neuro- podia with acicular spines. Setiger 5 modified, overlapping setiger 6 (Fig. 40a); setae including 5-6 superior dorsal geniculate setae (Fig. 40j) lying anterior to a semi-circular row of heavy spines alternating with smaller hastate companion setae (Fig. 40i) and small fascicle of unilimbate capillary neuro- setae located ventral to major spines; major spines falcate, with distinct lateral flange and crest of fine bristles on convex side of curved end (Fig. 40g-h), Branchiae from setiger 9, at first short reaching full-size on setigers 11-12, present only over first J of body, absent thereafter. Pygidium with 4 lobes, the dorsal pair smaller than ventral pair (Fig. 40k). Remarks: Polydora aciculata has several unique characteristics; the modified spines of SOUTHEASTERN AUSTRALIAN SPIONIDAE 253 Figure 40 — Polydora aciculata sp. nov. — a, anterior end, dorsal view; b-c, posterior notopodial spines; d, hooded hook from anterior setiger; e, hooded hook from medial body setiger; f, posterior neuropodial spine; g-h, major spines from setiger 5; i, hastate companion seta, setiger 5; j, su- perior dorsal seta from setiger 5; k, pos- terior end, dorsal view; 1, posterior seg- ment, dorsal view. [Scale 1 = 200 Mm; 2 = 20 wn; 3 = 20 Mm] setiger 5 are similar to those of P. caulleryi Mesnil and P. notialis sp. nov. (see below). Those of P. caulleryi are sharply falcate, with a crest of bristles, but no accessory structures, those of P. notialis bear a crest of bristles and a subterminal concavity. P. aciculata has a crest and a pronounced lateral flange. The presence of posterior acicular spines in both notopodia and neuropodia is rare in poly- dorids. P. convexa Blake and Woodwick (1972) has a similar arrangement with unidentate hooded hooks and specialized posterior noto- setae. The notosetae, however, are not acicu- la, but packets of fine needles. Boccardia basilaria Hartman (1961) has posterior noto- and neuropodial acicular spines. This situation was not noted by Hartman (1961) and was only determined during a recent examination of the types (Blake, unpublished). Ecology: Occupies twisting galleries in shells 254 JAMES A. BLAKE and JERRY D. KUDENOV Figure 41 — Polydora notidlis sp. nov. — a, anterior end, dorsal view; b, major spines and companion setae, setiger 5 showing dif- ferent angles and degrees of wear; c, hooded hook, setiger 8; d, hooded hook, setiger 35. [Scale 1 = 200 ^m; 2 = 20 Mm] of Haliotis ruber Leach, 1814. Associated with Polydora giardi Mesnil in the shells. Distribution: Victoria. Polydora notialis sp. nov. (Figure 41) Material examined: SOUTH AUSTRALIA. Tipara, from shell of Haliotus roei, 7 April 1971, coll. S. Shepherd (HOLOTYPE, NMV G2877). Description: A single, posteriorly incomplete specimen measures 3 5 mm long and 4 mm wide for 38 setigerous segments. Body gene- rally opaque white in alcohol, with lateral in- tersegmental brown pigment on setigers 1-3 and along margins of prostomium. Prostomium rounded on anterior margin, caruncle extending to posterior margin of setiger 3 (Fig. 41a); no occipital tentacle. Two pairs of subcutaneous eyes. Palps short, ex- tending posteriorly for 6 setigers. Setiger 1 with capillary notosetae in both noto- and neuropodia (Fig. 41a). Setigers 2, 3, 4, -, 6 and succeeding segments with 2 tiers of unilimbate capillary notosetae; first tier with shorter setae; number of capillary notosetae gradually diminishing in posterior segments; nature of posterior end unknown. Neurosetae of setigers 2, 3, 4, - and 6 as spreading fas- cicles of winged capillaries. Bidentate hooded hooks beginning on setiger 7, with 3 hooks in anterior setigers, accompanied by capillary setae for a few segments, number of hooks re- duced to 2 per neuropodium posteriorly; angle between main fang and secondary tooth acute, wider in anterior and becoming much reduced in posterior setigers (Fig. 41c-d); without con- striction on shaft. Setiger 5 modified, with heavy dorsal mus- culature, overlapping setiger 6; setae include dorsal fascicle of geniculate setae, curved row of 4-5 heavy spines, thin hastate companion setae (Fig. 41b) and ventral fascicle of uni- limbate capillary neurosetae; 4-5 major spines SOUTHEASTERN AUSTRALIAN SPIONIDAE 255 with expanded, slightly curved end, with subter- minal cavity (Fig. 41b), convex side of inflated end bearing bristled crest. Branchiae from setiger 7, reaching full-size by setiger 10, continuing for \ body length. Pygidium unknown, Remarks: Polydora notialis bears similarities to 2 groups of species. With a subdistal con- cavity on the major spines of setiger 5, P. no- tialis is similar to P. hornelli Willey (1905) from India, P. cavitensis Pillai (1965) from the Philippines, P. alloporis Light (1970a) from California and P. wobberi Light (1970b) from the Gulf of California. P. notialis differs from each of the foregoing species in possessing a crest of fine bristles on the convex surface of those same spines and in the lack of a con- striction on the shaft of the hooded hooks. P. notialis also resembles those species having a crest on the major spines: P. caulleryi Mesnil and P. aciculata (see remarks for preceding species). Distribution: South Australia. Polydora pilocollaris sp. nov. (Figure 42) Material examined: VICTORIA. Port Phillip Bay, PPBES Station 907 (HOLOTYPE, NMV G2878; 6 PARATYPES, NMV G2879-2880). Description: A small species, measuring up to 6-5 mm long and 5 mm wide for 70 setiger- ous segments. Holotype 3-5 mm long and 0-3 mm wide for 55 setigers. Opaque white in al- cohol. Prostomium distinctly incised anteriorly with 2 rounded lobes; caruncle thickened, ex- tending to posterior margin of setiger 2-3 (Fig. 42a); no occipital tentacle; eyes present as only 1 subcuticular pair, or absent. Setiger 1 with long finger-like notopodial lobe, notosetae lacking, with capillary neuro- setae. Setigers 2, 3, 4, -, 6 and succeeding setigers with fascicles of unilimbate capillary notosetae arranged in 2 tiers, setae of first tier the shorter and thicker; shorter setae gradually lost in posterior setigers. No specialized pos- terior notosetae, but with short pointed capil- laries accompanying longer ones. Neuropodia of setigers 2, 3, 4, - and 6 with spreading fascicles of unilimbate capillaries. Three to 5 bidentate hooded hooks from seti- ger 7, accompanied by a single capillary for first few setigers; hooks with relatively narrow angle between secondary tooth and main fang (Fig. 42d); no constriction on shaft. Setiger 5 modified, overlapping setiger 6; with dorsal fascicle of heavy geniculate setae, curved row of major spines, hastate companion setae (Fig. 42b) and a ventral fascicle of uni- limbate capillaries; major spines falcate, with collar on convex side bearing prominent tuft of bristles (Fig. 42c); unworn spines sometimes with a thin distal mucron. Branchiae from setiger 7, at first small, reaching full-size on setiger 10-11; absent from posterior i of body. Pygidium with 4 lobes, dorsal pair slightly smaller than ventral pair (Fig. 42e). Remarks: Polydora pilocollaris resembles P. convexa Blake and Woodwick (1972) from California in having a collar on the convex side of the major spines of setiger 5. P. con- vexa, however, lacks the tuft of bristles asso- ciated with the collar, has a posterior armature which P. pilocollaris lacks and has unidentate hooded hooks in posterior segments, rather than bidentate ones, as in P. pilocollaris. Distribution: Victoria, Port Phillip Bay. Polydora armata Langerhans, 1880 (Figure 43a-g) Polydora armata Langerhans, 1880, p. 93; Fauvel, 1927, p. 55; Hartman, 1941, p. 306; 1969, p. 127; Woodwick, 1964, p. 14; Day, 1967, p. 466; Rainer, 1973, p. 58; Read, 1975, p. 412. Polydora monilaris Ehlers, 1905, p. 43. Fide Day, 1954. Material examined: VICTORIA. Port Phillip Bay, Point Gellibrand, Williamstown, in shell of Haliotus ruber, 5 April 1977, coll. J. A. Blake and J. D. Kude- nov (70+, NMV G2881). SOUTH AUSTRALIA. Ti- para, shell of Haliotus roei, 19 May 1971, coll. S. Shepherd (4, NMV G3056). Description: A small species, up to 3-4 mm long and 0-5 mm wide for 30 segments. Prosto- mium bilobed anteriorly; caruncle extending to posterior margin of setiger 2; no occipital ten- tacle or eyes. Setiger 1 with 2-3 capillary no- tosetae; neurosetae more numerous. Setigers 2, 3, 4, -, 6 and succeeding setigers with unilim- bate capillary notosetae; posterior setigers with 256 JAMES A. BLAKE and JERRY D. KUDENOV Figure 42 — Polydora pilocollaris sp. nov. — a, anterior end, dorsal view; b, fascicle of 3 major spines and 2 companion setae, setiger 5; c, individual major spine, setiger 5; d, hooded hook; e, pygidium. [Scale 1 = 300 Mm; 2 — 20 Mm] cone-shaped bundles of acicular spines (Fig. 43g), these usually not emergent in small speci- mens. Neurosetae of setigers 2, 3, 4, - and 6 unilimbate capillaries. Bidentate hooded hooks beginning on setiger 7 (Fig. 43f), without con- striction on shaft, with acute, but fairly wide angle (about 50°) between main fang and secondary tooth. Setiger 5 highly modified, with dorsal fas- cicle of 2-3 geniculate setae, a row of heavy spines and ventral fascicle of unilimbate capil- lary neurosetae; major spines bidentate, teeth connected by lateral hood or cowling on one side (Fig. 43a-c) and thin shelf on opposite side (Fig. 43d-e). Cowling bears fine bristles seen only under oil immersion on new or un- worn spines. No companion setae of any sort. Branchiae from setiger 7, reaching full-size by setiger 8, continuing to setigers 11-12. Py- gidium small, cuff-shaped, with dorsal and ventral gaps. Remarks: We have found P. armata to have bidentate spines on setiger 5 as Lan- gerhans (1880) originally reported. The pres- SOUTHEASTERN AUSTRALIAN SPIONIDAE 257 Figure 43 — Polydora armata Langerhans — a-e, spines, setiger 5; f, hooded hook; g, posterior notopodial spine— Polydora ligni Webster— h, major spine, setiger 5; i, com- panion seta, setiger 5; j, hooded hook — Polydora websteri Hartman — k, major spine, setiger 5; 1, companion seta, setiger 5; m, major spine, setiger 5; n, hooded hook. 258 JAMES A. BLAKE and JERRY D. KUDENOV ence of minute bristles on the hood or cowl- ing is a new finding, although Figure 2(2) of Woodwick (1964) suggests that similar bris- tles are present in his Eniwetok specimens. P. armata, P. caulleryi Mesnil, P. quadrilo- bata Jacobi, 1883 and P. aciculata belong to a group of species having acicular spines in the posterior notopodia, hooded hooks lacking a constriction on the shaft and some similarity in design of the major spines of setiger 5. The major spines in P. armata are unique, but have been interpreted in various ways by dif- ferent authors, apparently because the spines look different in different views. Woodwick (1964) reviewed the literature on P. armata spines and presented illustrations based on Eniwetok specimens which compared favour- ably with the various past interpretations of other authors. In this manner he was able to support the synonymy of P. monilaris Ehlers, 1905 with P. armata, first proposed by Day (1954). Rainer (1973) did not disagree with this synonymy, but suggested that the types of P. monilaris should be examined. We support the synonymy of P. monilaris and P. armata, but also feel that type material should be lo- cated and examined. Distribution: Victoria; South Australia; Japan; South Africa; Europe; New Zealand; North Eastern Pacific. Polydora ligni Webster, 1879 (Figure 43h-j) Polydora ligni Webster, 1879b, p. 119; Blake, 1971, p. 5 (Synonymy); Foster, 1971, p. 22 (Synonymy); Light, 1977, p. 70. Material examined: VICTORIA. Port Phillip Bay, Hobsons Bay-Yarra River, MSG Stations, Feb.- Aug. 1975, coll. G. Poore and J. D. Kudenov, Sta. 128 (1144); 130 (58); 133 (7); 136 (28); 138 (1) (NMV G2882, 3 1 67-3 171); Werribee, MSG Monitoring Programme (P609), coll. J. D. Kudenov, Stations 10-11, 11 Jan. 1977 (3, NMV G3 172-3 174). Description: A medium to large species, often measuring up to 32 mm long for 80 setigerous segments (Blake, 1971), Australian specimens generally smaller. Colour in alcohol: opaque white, Prostomium incised with 2 widely separated Jobes; caruncle to setiger 2-3. Two pairs of eyes, widely spaced, forming a square; occipi- tal tentacle on caruncle behind eyes. No notosetae on setiger 1, but with well- developed finger-like notopodial lobe; neuro- setae with fascicle of delicate capillaries. Seti- gers 2, 3, 4, - and 6 with typical fascicles of unilimbate capillary noto- and neurosetae. No posterior spines. Bidentate neuropodial hooded hooks from setiger 7 (Fig. 43j), main fang forming approximately right angles with shaft; shaft with constriction. Setiger 5 with curved row of modified spines and closely adhering feather-like companion setae (Fig. 43i), spines weakly falcate with small secondary tooth (Fig. 43h); with ventral fascicle of unilimbate capillaries. Branchiae from setiger 7, continuing to pos- terior end. Pygidium a large flaring disc. Distribution: Victoria; North America; north- ern Europe. Polydora websteri Hartman, 1943 (Figure 43k-n) Polydora ciliata: Haswell, 1885, p. 272 [Not John- ston, 1838]. Polydora websteri Hartman, 1943, p. 70; Blake, 1971, p. 6 (Synonymy); Foster, 1971, p. 26 (Synonymy). Material examined: NEW SOUTH WALES. New- castle, ca. 1885 (3, AM G11390); Tuross Lake, from Crassostrea commercials, April 1977, coll. M. Skeel (4, NMV G3057). Description: A moderately-sized species measuring up to 10 mm long and 5 mm wide for 100 setigerous segments. Prostomium weakly bifid on anterior margin, caruncle extending to middle setiger 3; 4 eyes; no occipital tentacle. Setiger 1 lacking notosetae. Setigers 2, 3, 4, -, 6 and succeeding setigers with unilimbate capillary notosetae; posterior notopodial spines absent. Setigers 2, 3, 4, - and 6 with unilimbate capillary neurosetae. Six to 7 bidentate hooded hooks per neuropodium from setiger 7 having a wide angle between teeth (Fig. 43n); main fang forming right angle with shaft; with con- striction. Setiger 5 with dorsal and ventral fascicles of unilimbate capillaries and a row of major spines and bilimbate companion setae (Fig. 431), major spines falcate with lateral flange (Fig. 43k, m). SOUTHEASTERN AUSTRALIAN SPIONIDAE 259 Branchiae from setiger 7, absent from pos- terior $ of body. Pygidium disc-like with dor- sal notch. Remarks: Specimens agree well with P. web- steri from North American waters. The spe- cies may be widespread in oyster growing cen- tres where it forms mud-blisters in shells. Speci- mens of P. ciliata reported by Haswell (1885) most certainly should be referred to P. web- steri, since specimens in the Australian Mu- seum appear to be from Haswell's collection (E. Pope, communication to K. H. Woodwick, 1964). P. websteri is related to P. ciliata (John- ston, 1838) and has probably been mistaken for it on numerous occasions (see Hartman, 1943; Blake, 1971). Such references can only be verified by direct examination of the speci- mens in question. This uncertainty applies to the reports of P. ciliata in oysters from New South Wales by Whitelegge (1890) and Rough- ley (1922). Their specimens were possibly P. websteri, but in the absence of specimens, such suspicions cannot be verified. The presence in Australia of another closely related and newly described species, P. has- welli (see below), adds to the confusion. P. websteri and P. haswelli are separated with dif- ficulty (see key) and since both occur with oysters, careful attention to detail is required to separate reliably. Ecology: Drills into oyster shells, forming mud- blisters (for review, see Blake and Evans, 1973). Distribution: New South Wales; North Ameri- ca. Polydora haswelli sp. nov. (Figure 44) Material examined: NEW SOUTH WALES. Syd- ney, Wy-ar-gine Point, Middle Harbour, Galeolaria association, 19 Dec. 1968 (5, AM W13040); Scraping from huU of BEN SHORTRIDGE, tug from New- castle, coll. Sydney Harbour, 29 Feb. 1972 where tug was cleaned, P. Hutchings (1, AM W13041); Sydney Harbour, 8-10 m, near N. Chinamens Beach, sand, 8 May 1971, coll. P. Hutchings (HOLOTYPE, AM W 7283; 12 PARATYPES, AM W13042; 5 PARA- TYPES, NMV G2883); Botany Bay, Towra Beach, NSWSF Station, April 1973 (1, AM W13043); Cam- den Haven, Kennedy oyster lease, in mud blisters, 20 April 1977, coll. M. Skeel (3, NMV G3058). Description: A moderate-sized species, up to 13 mm long and 075 mm wide for 72 seg- ments. Body light tan with considerable black pigment, including a line along each side of prostomium, an area on peristomium anterior to palps and paired dorsolateral spots located on setigers 1-3; additional pigment on palps. Prostomium distinctly incised on anterior margin, caruncle extending posteriorly to middle of setiger 3 (Fig. 44a); occipital ten- tacle; 4 small, oval-shaped, subcuticular eyes, or eyes absent. Setiger 1 with short conical notopodial lobes lacking notosetae; neuropodial lobes finger- like, with fascicle of thin capillaries. Setigers 2, 3, 4, -, 6 and succeeding segments with spreading fascicles of unilimbate capillary noto- setae; posterior notosetae gradually reduced to few long slender erect capillaries; no pos- terior spines. Neurosetae of setigers 2, 3, 4, - and 6 unilimbate capillaries. Six to 8 bi- dentate hooded hooks in neuropodia from setiger 7; these with strong angle between sec- ondary tooth and main fang, main fang form- ing right angle with shaft; with prominent constriction on shaft (Fig. 44f); not accom- panied by capillaries. Setiger 5 strongly modified, musculature overlapping setiger 6; setae including dorsal fascicle of bilimbate setae, curved row of heavy spines (Fig. 44c-e) alternating with bilimbate companion setae (Fig. 44b) and a ventral in- ferior fascicle of capillaries; heavy spines slightly falcate with 2 accessory structures (Fig. 44c-e); one a prominent accessory tooth; the second a low flange located just distal to tooth. Branchiae from setiger 7, small at first, reaching full-size by setiger 9 and overlapping at mid-line; branchiae absent from posterior i of body. Pygidium a flaring disc with narrow dorsal notch (Fig. 44g). Remarks: Polydora haswelli resembles P. web- steri in having a flange on the major spines of setiger 5; it also resembles P. ciliata in having a sharp accessory tooth on those same spines. It differs from both species in bearing both a flange and an accessory tooth. The species is named for Professor W. F. Haswell, a pioneer- ing Australian polychaete taxonomist who 260 JAMES A. BLAKE and JERRY D. KUDENOV Figure 44-Polydora hasweW sp. nov.-a, anterior Distribution: New South Wales end, dorsal view; b, companion seta, seti- ger 5; c-e, major spines, setiger 5; f, hooded hook; g, posterior end, dorsal view. [Scale 1 = 200 ^m; 2 = 20 ™] Polydora latispinosa sp. nov. (Figure 45) published an early paper on Polydora in Aus- p ¥?™ a * €X f m ; ne * VICTORIA. Port Phillip Bay, tralia fHaswell 1SJ& ? omt C °° k > dred S ed ' °y sters and scallops (Pectin aU traua (Haswell, 1885). ba) shellS; 14 A n 1977 u Bla v ke v and L D SOUTHEASTERN AUSTRALIAN SPIONIDAE 261 Figure 45—Polydora latispinosa sp. nov.— a, anterior Kudenov (HOLOTYPE, NMV G2874; 50+ PARA- end, lateral view; b, anterior end, dorsal TYPES, NMV G2875-2876). view; c-f, major spines, setiger 5, various views; g-h, companion setae, setiger 5 (worn and unworn); i, hooded hook; j, posterior end, dorsal view; k, posterior bundle of needles. [Scale 1 = 500 ^m; 2 = 50 ^m; 3 = 20 /*m] Description: A large species, measuring up to 33 mm long and 1 2 mm wide for 210 setiger- ous segments. Body light tan in life with bright red blood vessels and branchiae; some dark ventral patterns caused by capillary blood 262 JAMES A. BLAKE and JERRY D. KUDENOV vessels; brown pigment along edges of car- uncle. Prostomium distinctly bilobcd anteriorly (Fig. 45a); caruncle extending posteriorly to end of setiger 2; with short triangular occipital tentacle formed by fold in caruncle at anterior margin of setiger 1 (Fig. 45b); 1-2 pairs of ob- scure, subcutaneous eyes present or eyes ab- sent. Palps short, thick, extending posteriorly only 5-6 setigers. Setiger 1 with well-developed finger-like notopodia, lacking notosetae; with fascicle of delicate capillary neurosetae. Setigers 2, 3, 4, -, 6 and succeeding segments with capillary noto- setae arranged in 2 tiers, first tier with shorter thicker setae; posterior notopodia with fewer, pointed thinner capillaries and flattened packets of fine needles, these sometimes pro- jecting through cuticle (Fig. 45j-k). Neuro- setae of setigers 2, 3, 4, - and 6 unilimbate ca- pillaries, not as prominent as notosetae. Bi- dentate hooded hooks from setiger 7, about 6 hooks per neuropodium at first, completely replacing capillary setae, increasing to 11-12 per neuropodium in middle body segments and diminishing, to 5-6 per ramus, posteriorly; hooks with strong angle between teeth through- out, main fang forming right angle with shaft; with distinct constriction on shaft (Fig. 45i). Setiger 5 large, overlapping 6 with heavy dorsal musculature; no dorsal fascicle; with a row of major spines alternating with narrow hastate setae (Fig. 45g-h), and ventral fascicle of unilimbate capillary setae; major spines fal- cate with lateral sheath or flange (Fig, 45c-f). Branchiae from setiger 7, at first small, reaching full-size on setigers 9-10, continuing to posterior end of body. Pygidium a large flaring disc, with wide dor- sal gap (Fig. 45j). Remarks: Polydora latispinosa belongs to the P. ciliata-websteri group; it is most closely re- lated to P. vulgaris Mohammad (1972) in hav- ing an occipital tentacle and posterior notopo- dial spines and in lacking superior notosetae on setiger 5. However, in P. latispinosa the pos- terior spines are packets of fine needles, while P. vulgaris bears individual notopodial spines. Ecology: Polydora latispinosa forms complex burrows and mud-blisters in shells of scallops and oysters in Port Phillip Bay. Distribution: Victoria, Port Phillip Bay. Polydora woodwicki sp. nov. (Figure 46) Material examined: VICTORIA. Port Phillip Bay, Point Gellibrand, Williamstown, shell of Haliotus ruber, 5 April 1977, coll. J. A. Blake and J. D. Ku- denov (HOLOTYPE, NMV G2873). Description: A moderate-sized species. The holotype is in 3 parts, measuring in total 15 mm long and 075 mm wide for 105 segments. Colour light tan with red-brown intersegmental pigment in posterior segments. Prostomium anteriorly entire; caruncle short, thick and bulbous, extending posteriorly to end of setiger 2 (Fig. 46a); no occipital ten- tacle; 2 pairs of oval eyes; palps regenerating on holotype (Fig. 46a) and normal length or form unknown. Setiger 1 with rounded notopodial lobes, lacking notosetae; neuropodial lobe ill-defined, but with spreading fascicle of delicate capil- laries. Setigers 2, 3, 4, -, 6 and succeeding setigers with short, thick notopodial lobes bearing unilimbate capillary notosetae arranged in 2 tiers: first tier with short setae, long, thin setae in second tier. In posterior setigers noto- podia reduced, setae fewer, more delicate. Pos- terior notosetae including both long and short stiff, erect capillary setae, imparting spinous appearance to posterior end (Fig. 46k). Neuro- setae of setigers 2, 3, 4, - and 6 in spreading fascicles of unilimbate capillaries. Five to 6 bi- dentate hooded hooks in anterior segments from setiger 7, completely replacing capillary setae, with hooks increasing to 8 per ramus in middle segments; hooks with wide angle be- tween teeth, main fang forming right angles or less with shaft; with prominent constriction on shaft (Fig. 46b). Setiger 5 modified, somewhat contracted and larger than setigers 4 and 6; with superior dor- sal fascicle of geniculate setae (Fig. 46j), a group of major spines alternating with bilim- bate companion setae (Fig. 46e, g) and a ven- tral inferior fascicle of unilimbated setae (Fig. 46i). Major spines slightly falcate with prom- SOUTHEASTERN AUSTRALIAN SPIONIDAE 263 Figure 46 — Polydora woodwicki sp. nov. — a, anterior end of holotype in dorsal view; b, hooded hook; c-h, 4 major spines and 2 com- panion setae, setiger 5; i, neuroseta, setiger 5; j, superior dorsal notoseta, setiger 5; k, posterior end, lateral view. inent accessory flange (Fig. 46c-d, f, h); un- worn spines with terminal mucron (Fig. 46h). Short branchiae from setiger 7, never ex- tending more than £ of distance across dorsum (Fig. 46a), continuing for only 21-23 seg- ments, absent from posterior I of body. Pygidium a small cuff-shaped disc, with mid- dorsal gap, somewhat telescoped into terminal segments (Fig. 46k). Remarks: Polydora woodwicki belongs to the P. websteri-ciliata group in having an accessory flange on the major spines of setiger 5. The species is most closely related to P. spondylana Mohammad (1973) in having a rounded pro- stomium. P. spondylana differs in having an enlarged, oval prostomium, a long narrow car- uncle reaching to setiger 5, no eyes and a large disc-like pygidium, while P. woodwicki has a typical narrow Prostomium, short bulbous caruncle reaching only to setiger 3, 4 eyes and a smaller cuff-shaped pygidium. P. woodwicki also resembles P. rickettsi Woodwick (1961) from Mexico in having a rounded prostomium. P. rickettsi, however, has a long caruncle reaching to setiger 5, the major spines of seti- ger 5 bear a distinct accessory tooth rather than a flange and the pygidium is large and disc-like, being larger than prepygidial seg- ments. This species is named in honour of Dr. Keith H. Woodwick, in recognition of his many studies on spionid polychaetes. Ecology: Bores into abalone shells. At Point Gellibrand it was associated with P. armata. Distribution: Victoria, Port Phillip Bay. 264 JAMES A. BLAKE and JERRY D. KUDENOV 50jjm Figure 47 — Polydora hoplura Claparede — a, com- panion seta, setiger 5; b-c, major spines, setiger 5; d, hooded hook; e, posterior notopodial spine. Polydora hoplura Claparede, 1870 (Figure 47) Polydora hoplura Claparede, 1870, p. 58; Carazzi, 1893, p. 20; Fauvel, 1927, p. 50; Day, 1967, p. 468; Read, 1975, p. 411. Material examined: NEW SOUTH WALES. Point Henry Pier, in oysters, 22 March 1975, coll. C. Pre- genzer (1, NMV G3059). TASMANIA. Simmons Beach, shells Crassostrea gigas, 17 June 1977, ship- ment specimens arranged M. Skeel (10-f, NMV G3060). Description: A large species measuring up to 40 mm long and 2 mm wide for over 160 seg- ments. Body colourless. Prostomium weakly incised, caruncle extend- ing to end of setiger 3; with low occipital ten- tacle. Notosetae lacking on setiger 1, neuro- setal capillaries present. Setigers 2, 3, 4, -, 6 and succeeding segments with fascicles of uni- limbate capillary notosetae; posterior segments with 1-2 large, medially directed falcate spines in each fascicle (Fig. 47e). Neurosetae of seti- gers 2, 3, 4, - and 6, short, unilimbate capil- laries; 8-10 bidentate hooded hooks from seti- ger 7, exhibiting wide angle between teeth; main fang and shaft forming oblique angle; with constriction on shaft (Fig. 47d). Major spines of setiger 5 falcate, with large subtcrminal flange, appearing tooth-like in certain views and depending upon wear (Fig. 47b-c); companion setae bilimbate (Fig. 47a); with dorsal and ventral fascicles of unilimbate capillary setae. Branchiae from setiger 7 along most of body, terminating just anterior to appearance of pos- terior spines. Pygidium a large disc with wide dorsal notch. Remarks: Australian specimens agree with des- criptions of specimens from New Zealand by Read (1975); the occipital tentacle and long caruncle, seen in these examples has not been reported in earlier accounts from Europe. Ecology: Forms mud-blisters in oysters. Distribution: New South Wales; Tasmania; Europe; South Africa; New Zealand. Genus Boccardiella gen. nov. Type-species: Polydora hamata Webster, 1879a. Gen- der feminine. Diagnosis: Prostomium anteriorly entire or in- cised, extending posteriorly as caruncle. Seti- ger 1 with or without notosetae. Setiger 5 strongly modified with only 1 type of major spine in a single row, with smaller companion setae; capillary notosetae present. Bidentate hooded hooks from setiger 7, main fang and shaft forming oblique angle with reduced (acute) angle between main fang and secondary tooth; main fang and secondary tooth more or less subequal without constriction on shaft; posterior notopodial spines or 'boat hooks' present or absent. Branchiae from setiger 2, present or absent on setiger 5, continuing for variable number of segments. Pygidium disc-like or reduced to small lobes or lappets, with or without small cirri or papillae. Remarks: Species with branchiae anterior to setiger 5 and with only 1 type of major spine on setiger 5 are herein removed from Boccar- dia to Boccardiella, which contains the follow- ing species: 1. B. bihamata sp. nov. (see below) SOUTHEASTERN AUSTRALIAN SPIONIDAE 265 2. B. hamata (Webster, 1879a) n. comb. Boccardia uncata Berkeley, 1927. Fide Blake (1966). 3. B. ligerica (Ferroniere, 1898) comb. nov. Polydora redeki Horst, 1920. Fide Blake and Woodwick (1971). Polydora uncatiformis Monro, 1938. New synonymy. 4. B. limnicola (Blake and Woodwick, 1976) comb. nov. 5. B. magniovata (Read, 1975) comb. nov. 6. B. truncata (Hartman, 1936) comb. nov. Boccardiella species are a distinct group of polydorids, sharing setal characteristics of seti- ger 5 with Polydora on the one hand, and branchial characteristics with Boccardia on the other. Distinctions between this species group and other Boccardia species were ob- served by Blake and Woodwick (1971). The synonymy of Polydora uncatiformis Monro, 1938 with Boccardia hamata (now Boccardiel- la) proposed by Blake (1966) was in error. Monro's species should instead be referred to synonymy with Boccardiella ligerica. Further comments on that situation will be published elsewhere (Blake, in preparation). For further information on the Boccardiella species see Blake and Woodwick (1971; 1976) and Read (1975). See Table 3 for generic relationships of Boccardiella to the other polydorid genera and remarks elsewhere. Boccardiella bihamata sp. nov. (Figure 48) Material examined: NEW SOUTH WALES. Botany Bay, Towra Beach, NSWSF Stations, 12-17 April 1973 (HOLOTYPE, NMV G2887; 4 PARATYPES NMV G2888-2889; 5 PARATYPES, AM W13044). Description: A moderate-sized species measur- ing up to 12 mm long and 0-75 mm wide for 75 setigers. Specimens mostly opaque white in alcohol; colour on some specimens indeter- minable due to staining by Rose Bengal. Prostomium distinctly bifid on anterior mar- gin, forming 2 rounded lobes (Fig. 48a), car- uncle extending posteriorly to near posterior border setiger 3; without occipital tentacle; 2 pairs of eyes, or eyes absent; when present anterior pair widely spaced and oval, posterior pair widely spaced and cup-shaped. Palps long, extending posteriorly for 20-22 setigers. Setiger 1 with well-developed noto- and neuropodia, but lacking notosetae (Fig. 48a); neurosetae including a small fascicle of capil- laries. Setigers 2, 3, 4, -, 6 and succeeding segments with spreading fascicles of unilimbate capillary notosetae arranged in 2 tiers, the more anterior setae shortest and thickest; num- ber of capillary setae gradually reduced through middle body segments. Posterior end highly modified, with 2 types of acicular noto- podial spines; each notopodium with (1) a large, stout recurved hook (Fig. 48k), (2) accom- panied by 3-4 smaller straight acicular spines (Fig. 481-n), and 2-3 long capillary setae (Fig. 48n); parapodia elevated on each side, forming middorsal channel into which acicular spines project (Fig. 48m-n). Neuropodia of setigers 2, 3, 4, - and 6 with fascicles of unilimbate capil- laries. Bidentate hooded hooks from setiger 7, numbering 6-8 through middle segments ac- companied by 2-3 inferior capillary setae, re- duced to 3-4 in posterior segments, with capil- laries becoming lost; structure of hooks chang- ing from anterior to posterior setigers: hooks initially bidentate with an obtuse angle be- tween nearly subequal teeth (Fig. 48g-h), sec- ondary tooth becoming lost in posterior seti- gers, resulting in unidentate hooded hooks (Fig. 48i), with superior hooks usually losing hood (Fig. 48j), with broken hood occasionally seen hanging from shaft. Setiger 5 with dorsal superior fascicle of 5-6 stout geniculate unilimbate setae (Fig. 48f), row of major spines (Fig. 48c-d) alternating with bilimbate companion setae (Fig. 48b) and a ventral inferior fascicle of 10-12 unilimbate capillaries (Fig. 48e); major spines simple, weakly falcate, without accessory structures (Fig. 48c-d). Branchiae occurring setigers 2, 3, -, -, 6 and subsequent segments for about f of body length. Pygidium a flattened plate from which 2 short lateral cirri arise (Fig. 48m), closely re- sembling that of B. hamata. The anus opens dorsally. Remarks: Boccardiella bihamata is closely re- lated to B. hamata and B. ligerica in the struc- 266 JAMES A. BLAKE and JERRY D. KUDENOV Figure 48 — Boccardiella bihamata gen. et sp. nov. — a, anterior end, dorsal view; b, companion seta, setiger 5; c-d, major spines, setiger 5; e, neuroseta, setiger 5; f, superior dor- sal seta, setiger 5; g-j, progression of hooded hook structure from anterior (g), medial (h) and posterior (i, j) setigers; k-1, large hook and small spine from pos- terior notopodium; m, posterior end, dor- sal view; n, posterior segment, dorsal view. [Scale 1 = 200 ^m; 2 = 200 Mm; 3 = 50 ^m; 4 = 50 /*m; 5 = 20 /*m] ture of the posterior end and in bearing large recurved posterior notopodial hooks. It differs significantly from both in the possession of an additional type of acicular notopodial spine in posterior segments and in the presence of uni- dentate neuropodial hooks in the same region. See Blake and Woodwick (1971; 1976) for further information on related species. Distribution: New South Wales, Botany Bay. Boccardiella limnicola (Blake and Woodwick, 1976) comb. nov. Boccardia limnicola Blake and Woodwick, 1976, p. 123. Material examined; QUEENSLAND. Brisbane, Centenary Bridge, Feb. 1972 (10+, AM W7491); De- ception Bay, Burpengary Creek, Jan. 1975, coll. J. Davie (70+, AM W7126). VICTORIA. Port Phillip Bay, Werribee, MSG Monitoring Programme (P609), coll. J. D. Kudenov, Station 10, 11 Jan. 1977 (16, NMV G3061-3063); Lake Bong Bong, 17 Nov. 1970, coll. B. V. Timms (HOLOTYPE AM W7033; 19 PARATYPES AM W7034); Glenelg River, July 1975, coll. J. D. Kudenov and P. A. Hutchings (48, SOUTHEASTERN AUSTRALIAN SPIONIDAE 267 NMV G3064-3065); Fitzroy River, July 1975, coll. J. D. Kudenov and P. Hutchings (20, NMV G3066); Port Fairy, July 1975, coll. J. D. Kudenov and P. Hutchings (50+, NMV G3068); Warrnambool, Merri River, July 1975, coll. J. D. Kudenov and P. Hutch- ings (21, NMV G3067); Paynesville, Gippsland Lakes, jetty, 2 m, sand, Aug. 1975, coll. J. D. Kude- nov (9, NMV G3069-3070). Remarks: The reader is referred to Blake and Woodwick (1976) for a description of the species. We note in the new material, that the length of the branchiae on setiger 5 is highly variable, being rudimentary on some specimens to as long as those on setigers 4 and 6 in others. Boccardiella limnicola is similar to B. mag- niovata (Read, 1975) from New Zealand. The latter species differs significantly, however, in lacking branchiae on setiger 5 and form of the pygidium. The two species are very closely related, however, and share a common an- cestry. Ecology: B. limnicola was described from freshwater Victorian lakes. Although favouring areas of reduced salinities, we have now found the species to occur in more saline environ- ments such as tidal rivers and near sewage out- falls. One record of a freshwater spionid over- looked by Blake and Woodwick (1976) was that of Marenzellaria wireni reported by Holm- quist (1967) from Alaska. Distribution: Queensland; Victoria. Genus Pseudopolydora Czerniavsky, 1881 emended Pseudopolydora Czerniavsky, 1881. Type-species: P. antennata (Claparede, 1870), by monotypy. Carazzia Mesnil, 1896. Type-species: C. antennata (Claparede, 1870), by monotypy. Polydorella Augener, 1914. Type-species: P. prolifera Augener, 1914, by monotypy. New synonymy. Diagnosis: Prostomium entire or incised, ex- tending posteriorly as a caruncle; occipital ten- tacle present or absent; eyes present or absent. Setiger 1 generally reduced, with or without notosetae (and in some species without neuro- setae if animals are reproducing asexually and regenerating). Setiger 5 generally not distinctly enlarged, with noto- and neuropodia often well-developed with postsetal lobes and bear- ing spreading fascicles of capillaries; with curved row of heavy spines of 2 types, or single type with companion setae; modified setae often forming J- or U-shaped setal group. Posterior notopodial spines or 'boat hooks' pre- sent or absent. Neuropodial hooded hooks from setiger 8, these hooks bidentate, with secondary tooth closely applied to main fang, with con- striction on shaft; accompanying capillaries present or absent. Branchiae first appearing posterior to setiger 5. Pygidium enlarged or reduced, collar-like or divided into lobes or small lappets. Remarks: Species of the genus Pseudopolydora are among the least modified of the Polydora- complex. Read (1975) pointed out that the low degree of modification to setiger 5 and the structure of the hooded hooks are more charac- teristic of Pseudopolydora than the first ap- pearance of hooded hooks on setiger 8. One species, P. reishi Woodwick (1964) W'as noted by Read (1975) to be doubtfully placed in Pseudopolydora. We agree, and herein trans- fer it to Carazziella. Polydorella, with modified spines on setiger 4, rather than 5, was noted by Read (1975) to be closely related to Pseudopolydora. In this study, we have been able to examine the type series of Polydorella prolifera, the type-species of the genus, and to demonstrate that setae may be present on the generally achaetous segment 1 on some specimens. This is a function of the state of regeneration, since the species is cap- able of asexual reproduction by paratomy. The discovery of a closely related and previously undescribed species, has confirmed this point of view. We therefore place Polydorella into synonymy with Pseudopolydora. The status of Polydorella novaegeorgiae Gibbs, 1971 re- mains uncertain, until type material is ex- amined. Its overall morphology is, however, very similar to species of Pseudopolydora. Pseudopolydora is herein defined to include species in which setiger 1 is completely reduced to an achaetous condition. KEY TO SPECIES OF PSEUDOPOLYDORA FROM AUSTRALIA la. Maximal number of segments 15-16; small, up to 3 mm long 2 b. More than 20 segments; large, over 6 mm long 3 2a. Pennoned spines of setiger 5 with fe- 268 JAMES A. BLAKE and JERRY D. KUDENOV nestrated edge surrounding cavity, and long bristled apex (Fig. 50n); prostomium entire P. prolifera b. Pennoned spines of setiger 5 with ex- panded bristle-topped ends (Fig. 50 f-h); prostomium incised. , P. stolonifera 3a. Prostomium entire. . P. paucibranchiata b. Prostomium incised 4 4a. Prostomium with 2 widely flaring an- terior lobes; occipital tentacle just behind eyes; caruncle continuing posteriorly to setigers 3-4; branchiae without glands P. kempi b. Prostomium with 2 rounded anterior lobes (Fig. 49a); nuchal tentacle at posterior end of caruncle on setiger 2; branchiae with glands (Fig. 49a) P. glandulosa Pseudopolydora paucibranchiata (Okuda, 1937) Polydora (Carazzia) paucibranchiata Okuda, 1937, p. 231. Pseudopolydora paucibranchiata: Imajima and Hart- man, 1964, p. 288; Blake, 1975, p. 215; Blake and Woodwick, 1975, p. 110; Read, 1975, p. 414; Light, 1977, p. 71 (Synonymy). Material examined: NEW SOUTH WALES. Botany Bay, Towra Beach, NSWSF Stations, 12 April 1973 (1, AM W 13045); Jervis Bay, Currubene Creek. 23 Jan. 1977 (1, AM W5223). VICTORIA. Port Phillip Bay, PPBES Stations 904 (2); 919 (15); 924 (2); 926 (10); 939 (4); 942 (47): 953 (36); 984 (2) (NMV G3 175-3 182); Hobsons Bay-Yarra River, MSG Sta- tions, Feb., May 1975, coll. G. Poore and J. D. Kude- nov, 128 (637); 130 (3); 131 (4); 132 (15); 138 (21) (NMV G2890, 3183-3187); Westernport, WPBES Stations 1702 (27, NMV G3133); 1716 (64, NMV G3134). Description: Prostomium entire on anterior margin, caruncle extending to posterior mar- gin of setiger 3; occipital tentacle present. Palps long, extending i length of worm. Setiger 1 without notosetae. Major spines of setiger 5 arranged in a U-shaped line, consisting of 12- 14 pennoned, curved spines and an equal num- ber of simple falcate spines; setiger 5 rela- tively unmodified, same size as setigers 4 and 6. Hooded hooks from setiger 8, completely re- placing neuropodial capillaries. Branchiae from setiger 7, continuing to about setiger 35. Pygidium with small, ventral, collar-like disc with large dorsal gap. Remarks: Specimens agree well with others ob- served by us in California. P. paucibranchiata is widely distributed in bays throughout the Pacific and is possibly introduced into Aus- tralia. It is similar in some respects to P. kempi, but differs in having an entire, rounded, rather than incised prostomium, and a ventral collar- like rather than a dorsally flaring pygidium. Distribution: New South Wales; Victoria; Jap- an; California; New Zealand. Pseudopolydora kempi (Southern, 1921) Polvdora (Carazzia) kempi Southern, 1921, p. 636; Okuda, 1937, p. 233. Neopvgospio laminifera Berkeley and Berkeley, 1954, p. 462. Fide Banse 1972, p. 219. Pseudopolydora kempi japonica Imajima and Hart- man, 1964, p. 287. Pseudopolydora kempi califomica Light, 1969, p. 542. Pseudopolydora kempi: Hartman, 1969, p. 167; Blake, 1975, p. 215; Blake and Woodwick, 1975, p. 126; Light, 1977, p. 71. Pohdora (Pseudopolydora) kempi japonica: Banse, 1972, p. 219. Material examined: QUEENSLAND. Moreton Bay, Jackson Creek, 12 July 1973, coll. C. Wallace and P. Hutchings (1, AM W6042); Same, Serpentine Creek, sand bank, coll. C. Wallace (1, AM W6043); Brisbane River, Quarantine Station May 1972 (6, AM W7474). NEW SOUTH WALES. Merimbula, Sta. 197H, 200J, 6 Oct. 1975, sand, coll. by J. H. Day, et al. (1, AM W11703, 1, AM W11704). Description: A moderately large species, up to 25 mm long for 45 setigers in some areas, generally smaller in Australia. Body opaque white, otherwise colourless, but with black an- terior intersegmental pigment in other locali- ties (Light, 1969). Anterior end expanded. Prostomium with 2 widely flared lobes, continuing posteriorly as a caruncle to setiger 3-4. Occipital tentacle pre- sent, just posterior to eyes; 2 pairs of eyes. Setiger 1 without notosetae, postsetal lobes erect, short. Setigers 2 and succeeding with well-developed notopodial fascicles of unilim- bate setae arranged in 3 tiers: shortest thickest setae in first tier; becoming progressively longer and thinner in tiers 2 and 3; setigers 4 and 6 with anterior tier developed ventrally, curving posteriorly and approaching the condition found in modified setae of setiger 5 (see Southern, 1921, p. 636, pi. 28, fig. 30; Okuda, SOUTHEASTERN AUSTRALIAN SPIONIDAE 269 1937, p. 235, fig. 13b). Neurosetae setigers 2-7 with well-developed fascicles of unilim- bate capillary setae; 25-30 bidentate hooded hooks from setiger 8. Setiger 5 with about 23-25 pairs of each of 2 types of specialized spines arranged in a J-shaped row: outer row consisting of short pennoned setae, inner row consisting of simple spines, slightly curved apically. Branchiae from setiger 7, continuing pos- teriorly for about i body length. Pygidium missing in Australian specimens, but elsewhere reported disc-like, dorsally notched and bear- ing 2 erect processes on each dorsolateral side (Okuda, 1937; Blake, 1975; Blake and Wood- wick, 1975). Remarks; Pseudopolydora kempi is one of the least modified members of the genus or the Polydora-complex for that matter. In over- all view, it is difficult to see the modification of setiger 5. Specimens reported here agree well with descriptions from other areas. Lack of pigmentation may be actual or be an artifact of preservation. Distribution: Queensland; New South Wales; Japan; India; British Columbia, California. Pseudopolydora glandulosa sp. nov. (Figure 49) Material examined; QUEENSLAND. Moreton Bay, Middle Banks, Stas. 43, 54, Sept. 1972, coll. W. Stephenson (2, QM Gl 1598-1 1599). NEW SOUTH WALES. Botany Bay, Towra Beach, NSWSF Sta- tions, April 1973, (2, AM W13046). VICTORIA. Westernport, Crib Point, CPBS Station 300 (HOLO TYPE, NMV G2899; 5 PARATYPES, NMV G2900). Description: A small species, measuring up to 11 mm long and 11 mm wide for 37 setiger- ous segments on an incomplete specimen. Ani- mals heavily pigmented, with brown pigment on prostomium, peristomium and anterior mar- gins of setigers 1-6; brown pigment on pos- terior margins of most segments ventrally. Anterior end broad, head and first 2 setigers small and contracted relative to setiger 3 and following segments. Prostomium distinctly in- cised on anterior margin (Fig. 49a), but not formed into flaring lobes; caruncle terminating at middle to end of setiger 2; occipital tentacle inserted at posterior end of caruncle (Fig. 49a). Peristomium well-developed, folded into 2 lobes; palps missing. Setiger 1 with short notopodial lobes, lack- ing notosetae; neuropodia well-developed with delicate capillary setae (Fig. 49a). Setiger 2 and succeeding setigers with rounded notopo- dial lobes and long, thin capillary notosetae, ar- ranged in 2 tiers, with shorter, thicker setae in first tier and longer, thinner setae in second; posterior spines absent; neurosetae of setigers 2-7 with dense fascicles of broad bilimbate capillary setae having fine granulations on the shaft (Fig. 49d). Bidentate hooded hooks num- bering 18-21 per fascicle without accompany- ing capillary setae beginning in setiger 8; hooks with reduced angle between teeth, a striated hood with serrated hood opening and distinct constriction on shaft (Fig. 49e). Setiger 5 only slightly modified, not much different from 4 and 6, with J-shaped row of modified spines located between the noto- and neuropodial fascicles. Outer setae pennoned (Fig. 49c), inner setae as simple spines with inflated slightly curved apex (Fig. 49b). Branchiae from setiger 7, continuing for only about 1 1 setigers. Branchiae grandular, with additional line of glands extending across dorsum between each gill. Pygidium unknown. Remarks; Pseudopolydora glandulosa has a relatively unmodified fifth setiger and ap- proaches P. kempi (Southern) in certain re- spects. The 2 species differ strikingly in overall appearance, especially the larger more erect notopodial lobes and wider flaring prostomium of P. kempi. The pigmentation of P. glandulo- sa contrasts with that of P. kempi in Australia, where it is generally lacking. In California, however, P. kempi is heavily pigmented in the intersegmental regions of the anterior setigers (Light, 1969, fig. 9), P. glandulosa on the other hand, has prominent dorsal and ventral pig- mentation. The striated hood of the hooded hooks of P. glandulosa is unique for the genus. Distribution: Queensland; New South Wales; Victoria. Pseudopolydora stolonifera sp. nov. (Figure 50a-j) Material examined: VICTORIA. Westernport, Crawfish Rock, 5 m, associated with various sponge 270 JAMES A. BLAKE and JERRY D. KUDENOV Figure 49 — Pseudopolydora glandulosa sp. nov. — a, anterior end, dorsal view; b, simple spine, setiger 5; c, pennoned spine, setiger 5; d, neuroseta, setiger 5; e, hooded hook. [Scale 1 = 200 m; 2 = 30 Mm; 3 = 20 species, and pycnogonid Nymphon mulleri, 3 Jan. and 13 May 1977, coll. D. Staples (HOLOTYPE, NMV G2897; 60+ PARATYPES, NMV G2898; 50+, NMV, G3071). Description: A small species measuring 2-8 mm long and 0-4 mm wide for a maximum of 16 setigerous segments. First 8 setigers narrow, last 8 greatly enlarged. Body with reticulated brown pigment on gills, peristomium and later- ally on anterior body segments; reflective white pigment on prostomium observed in living ani- mals. Some specimens undergoing active asexual reproduction with regeneration of a new head beginning between setigers 10-11. Following description refers to entire animals only, not regenerating ones. Prostomium incised on anterior margin (Fig. SOUTHEASTERN AUSTRALIAN SPIONIDAE 271 50a-b), caruncle terminating at posterior edge of setiger 2, occipital tentacle and eyes absent. Setiger 1 reduced, without notopodium or notosetae; neuropodium ventrally displaced, with fascicles of winged capillaries (Fig. 50b-c). Setigers 2, 3, 4, -, 6 and succeeding setigers with fascicles of unilimbate capillary notosetae, these becoming longer and fewer in posterior setigers. Neurosetae of setigers 2, 3, 4, -, 6 and 7 with unilimbate capillaries. Bidentate hooded hooks beginning on setiger 8, numbering 6-S throughout, lacking accompanying capillaries; hood with serrated opening (Fig. 50d). Setiger 5 modified, with both types of major spines arranged in a slightly curved line (Fig. 50b, e); major spines numbering 3-5 of each type, including: (1) pennoned, densely bristled spines with expanded ends (Fig. 50f-h) in ventral row, and (2) simple acicular spines (Fig. 50e, i) in dorsal row; no superior dorsal fas- cicle; with inferior ventral fascicle of 3-5 bilim- bate setae (Fig. 50j). Two pairs of branchiae on setigers 7-8. Py- gidium with 3 small cirri developed above smooth terminal ending (Fig. 50b). Asexual Reproduction: The collection includes animals in various stages of regeneration and budding. A new anterior end develops in a growth zone between setigers 10-11. Small palp buds appear with morphology of the head developing prior to budding. This type of asexual reproduction where development precedes budding is called paratomy (Schroe- der and Hermans, 1975). Different stages of re- generation are seen in the collections. Some late stages of regeneration appear complete, but setiger 1 lacks notosetae, a situation similar to that seen in P. prolifera (see below). Remarks: Pseudopolydora stolonifera shows more modification to setiger 5 than most spe- cies of the genus, in having poorly developed parapodia and in lacking notosetae. The struc- ture of the spines on setiger 5 and hooded hooks, however, clearly ally this species to Pseudopolydora and not Carazziella. The spe- cies is most closely related to P. prolifera, but differs in having a bilobed instead of an entire prostomium, having serrated openings on the hooded hooks, lacking superior dorsal noto- setae on setiger 5 and by having the pennoned major spines of setiger 5 with expanded den- sely bristled tips instead of a fenestrated spine with apical cavity. Distribution: Victoria, Westernport Bay, on sponges. Pseudopolydora prolifera (Augener, 1914) comb. nov. (Figure 50k-n) Polydorella prolifera Augener, 1914, p. 16. [Not Fau- vel, 1930, p. 36; 1953, p. 322]. Material examined: WEST AUSTRALIA. Sharks Bay, South Passage, 9 m, 16 April 1905, S.W. Aus- tralia Exped. 1905 (2 SYNTYPES, ZMH V 10 106). Description: A small species, capable of asex- ual reproduction by paratomy. Both syntypes with 16 setigers measuring up to 2*8 mm long and 0-6 mm wide. Both specimens slightly damaged, showing evidence of deterioration. Prostomium entire on anterior margin, no apparent caruncle or occipital tentacle; eyes absent. Setiger 1 reduced, without notopodium or notosetae; neuropodial lobe small, dorsally elevated with 3-4 very fine capillaries on one specimen and absent on second leaving an en- tirely achaetous segment. Setigers 2, 3, 4, -, 6 and succeeding setigers with fascicles of uni- limbate capillary notosetae; neurosetae fewer in number. Neuropodial hooded hooks from setiger 8 (Fig. 50k), 4-5 hooks at first increas- ing to 6 in middle body segments, reduced to 2-3 in posterior segments, without accompany- ing capillaries. Setiger 5 only slightly modified, with well- developed dorsal and ventral fascicles of uni- limbate capillary setae (Fig. 501) and 2 types of major spines: (1) 1 type distally enlarged, with one edge fenestrated and surrounding cavity and other bearing elongated bristle- topped projection (Fig. 50n); (2) second type a simple acicular spine (Fig. 50m). Branchiae missing on types, but according to Augener (1914), a single pair occur on what would be setiger 7. With a dark- brown achae- tous segment bearing 2 palp buds lying between setigers 10 and 11. denoting area where a new anterior end is regenerating. Pygidium consist- ing of 2 thickened lateral lobes, with wide me- dial gap (Augener, 1914). Remarks: Polydorella was described as having 272 JAMES A. BLAKE and JERRY D. KUDENOV Figure 50—Pseudopolydora stolonifera sp. nov.— a-b, entire animal in dorsal view, drawn in 2 parts (a, is the first 13 setigers; b, the last 3 setigers); c, anterior end, lateral view; d, hooded hook; e, group of spines, setiger 5, lateral view; f-h, pennoned spines^ setiger 5; i, simple spine, setiger 5; j, neuroseta, setiger 5—Pseudopoivdora prolifera (Augener)— k, hooded hook; 1, superior notoseta, setiger 5; m, simple spine, setiger 5; n, pennoned-type spine, setiger 5. SOUTHEASTERN AUSTRALIAN SPIONIDAE 273 the fourth setigerous segment modified instead of the fifth. An examination of the 2 syntypes has shown that one bears an achaetous first segment with a conspicuous neuropodial lobe, 3 normal setigers and then the modified seg- ment. The second specimen has 3-4 very fine capillary setae associated with the neuropodial lobe on the first segment. Thus Augener's spe- cies should be referred to Pseudopolydora since 4 setigers precede the modified one, in- stead of 3. Since 1 of the 2 specimens did ac- tually lack setae on the first segment, the presence and absence of setae in setiger 1 in different specimens of the species emphasize the tendency for reduction in setiger 1 among species of Pseudopolydora. In P. prolifera, however, the reduction of setiger 1 is in part explained by its mode of asexual reproduction. Specimens undergoing anterior regeneration would probably demonstrate various stages of setal development. If the regenerating region involves a specific characteristic such as an already reduced first setiger, then the total ab- sence of setae on setiger 1 would not be un- expected. We have this very situation in an- other, closely related species, P. stolonijera (see above). Since the type-species of the genus Poly- dorella is now known to lie well within the definition of the genus Pseudopolydora, the former genus must be sunk into synonymy in its entirety. Still another species may exist with a re- duced first setiger and asexual reproduction. Fauvel (1930; 1953) and Gravely (1927) re- ported a species from India which Fauvel called Polydorella prolifera and Gravely re- ferred to as 'post-larval chaetopterid'. This species, however, clearly has a different type of acicular spine on the modified segment. Gravely (1927) indicated it lived on the surface of sponges, as does Pseudopolydora stolonijera (see above). Distribution^ 'est Australia, Sharks Bay. Discussion Genera of the Spionidae: Apart from disagree- ments relating to the status of particular taxa at subgeneric or generic levels, the spionid genera have been relatively well understood due to recent efforts by Hartmann (1941; 1959), Pettibone (1963), Day (1967) and Foster (1971). Australian spionids, however, have presented so many new taxa that a re-examination of the status of certain genera and generic groups must be undertaken. Throughout this paper we have reviewed certain genera or generic com- plexes where appropriate, and either expanded their definitions to accommodate new species or have named new genera to accommodate taxa otherwise too far removed from existing taxa. We have also taken the opportunity to partly alter the generic arrangement of the Polydora-complzx because of the new species groups in our collections. At least 3 genera: Polybranchia Potts, 1928; Anaspio Chamberlin, 1920; and Morants Chamberlin, 1919 are not included, although Fauchald (1977a) diagnosed all 3 in his book on polychaete families and genera. We con- sider that the former may be validly placed among genera with frontal horns, but prefer to wait, pending a re-examination of the type- species. The 2 latter genera are highly doubt- ful. Both were possibly described from dam- aged material and neither has been reported since the original description. Type material is apparently lost. Both Anaspio and Morants are best considered nomina dubia. Generic Characteristics: Different taxonomic characters are important at the generic level in different generic groups. This is expected since different groups of species have specialized along their own lines following an early dicho- tomy from ancestral spionids. Spionids pos- sibly date from Devonian time (Cameron, 1967; 1969), and one should not expect to use the same characteristics with equal weight in genera such as Scolelepis, Prionospio and Polydora. Species of these genera and others have different ecological requirements and modes of life. Structures important in the adap- tive radiation of polydorids are certainly not the same as those in genera such as Scolelepis or Prionospio. Each genus has evolved inde- pendently and its species may not necessarily utilize the same structures as species belonging to other genera. With spionids, then, we can- not organize a simple system of uniform 274 JAMES A. BLAKE and JERRY D. KUDENOV characters to diagnose genera as is possible for certain other families (Kudenov and Blake, 1978). Instead, we must consider a wide range of features inherent in a given generic group; it is to be expected that the reasons for separating genera with frontal horns are not necessarily the same as those distinguishing groups with pointed prostomia. Some characteristics do remain conservative throughout the family: these include the shape of the prostomium and first appearance of branchiae. These characters have been and are still important in establishing the current generic system for spionids. For example, Sco- lelepis-Dispio; Malacoceros-Rhynchospio\ Po- ly dora-Boccardia ; Spio-Microspio ; Paraprio- nospio-Prionospio are some examples of gene- ric groups where the first appearance of bran- chiae are important. Other characters import- ant to some genera are the degree of fusion of branchiae and dorsal lamellae, the presence or absence of notopodial hooded hooks, the structure of the hooded hooks and their initial occurrence in some genera (in other genera, the occurrence of hooded hooks is age depend- ent [Hannerz, 1956] and is not always reli- able). Some genera have their own peculiar mor- phology: Dispio has accessory branchiae on the postsetal notopodial lobes, the polydorids have various degrees of modifications to seti- ger 5, and species of the Prionospio-complex have various peristomial and branchial modi- fications not found in other groups. Zoogeographic Considerations: Three species included in this report are only known from West Australia. Of the 63 species reported herein for the Southeast and 2 from the Great Barrier Reef (Queensland), 44 are endemic to Australia. The other species generally have more cosmopolitan distributions. Such species include: Aonides oxycephala, Malacoceros in- dicus, Paraprionospio pinnata, Prionospio steen- strupi, P. ehlersi, P. cirrijera, Spiophanes bom- byx, S. kroeyeri, Polydora flava, P. socialis, P. giardi, P. armata, P. ligni, P. websteri, P. hoplura and Boccardia polybranchia. Two species, Pseudopolydora tempi and P. Pauci- branchiata have wide distributions in the Pacific basin, while Boccardia chilensis, and Priono- spio aucklandica are limited to the Southern Hemisphere. Boccardia proboscidea is known from Western North America and Japan and has apparently been introduced to Port Phillip Bay. Spiophanes wigleyi . is known from continental shelf margins of eastern North America and the Gulf of Mexico. However, this apparent disjunct range may represent in- adequate sampling on continental shelf mar- gins. Several bays have been found to harbour large numbers of endemic species. For ex- ample, Port Phillip Bay is the only known lo- cality for 10 species. Westernport Bay has 2 endemic species and 1 species occurs in both bays. It is likely that several of these species will be found to be more widely distributed in Victoria once careful investigation of the Gippsland Lakes is completed. Two other areas containing large numbers of endemic forms are Botany Bay in New South Wales and Moreton Bay in Queensland; four new taxa come from the former and 5 from the latter location. Other areas in New South Wales and Queensland contain still other en- demic species. While many endemic species occur in several localities, the high percentage of species known from just a single locality is surprising. We suspect that some of these iso- lated records may reflect lack of adequate col- lecting, but it does appear that certain bodies of water, such as Port Phillip Bay, Victoria are important areas for adaptive radiation. We anticipate the discovery of still more spe- cies of Spionidae from this region. 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Polychaetous annelids of the Pacific coast in the collection of the Zoo- logical Museum of the University of California. Univ. Calif. Publ ZooL 13: 175-234. Verrill, A. E., 1873. Report upon the invertebrate animals of Vineyard Sound and the adjacent waters with an account of the physical charac- ters of the region. Rep. U.S. Fish. Comm. Wash. 1871-1872: 295-778. f 1879. Notice of recent additions to the marine Invertebrata, of the north-eastern coast of America, with descriptions of new genera and species and critical remarks on others. Pt. 1. Annelida, Gephyrea, Nemertina, Nematoda, Poly- zoa Tunicata, Mollusca, Anthozoa, Echinoder- mata, Porifera. Proc. U.S, Nat. Mus. 2: 165-205. 280 JAMES A. BLAKE and JERRY D. KUDENOV -, 1881. New England Annelida. Pt. 1. His- torical sketch, with annotated list of the species hitherto recorded. Trans. Conn. Acad. Arts Sci. 4: 285-324. Webster, H. E., 1879a. Annelida Chaetopoda of the Virginian coast. Trans. Albany Inst. N.Y. 9: 202- 269, 1 1 pis. ■, 1879b. The Annelida Chaetopoda of New Jersey. Ann. Rep. N.Y. State Mus. Nat. Hist. 32: 101-128. [Plates cited were not published until 1886]. and J. E. Benedict, 1884. The Annelida Chaetopoda from Provincetown and Welhleet, Mass. Rep. U.S. Comm. Fish. Wash., vol. for 1881, pp. 699-747, 8 pis. — and — , 1887. The Annelida Chaeto- poda from Eastport, Maine. Rep. U.S. Comm. Fish. Wash., vol. for 1885, pp. 707-755, 8 pis. Whitelegge, T., 1890. Report on the worm disease affecting the oysters on the coast of New South Wales. Rec. Aust. Mus. 1 : 41-53. Willey, A., 1905. Report on the Polychaeta collec- ted by Professor Herdman, at Ceylon in 1902. Suppl. Rep. Ceylon Pearl Oyster Fish., pt. 4: 243-324, pis. 1-8. Wiren, A., 1883. Chaetopoder fran Sibiriska Ishaf- vet och Berings Haf insamlade under Vega-Ex- peditionen 1878-79. V ega-Exped.-V ctenskapliga lakttagelser 2: 383-428, pis. 27-32. Woodwick, K. H., 1953. Polydora nuchalis, a new species of polychaetous annelid from California. Jour. Wash. Acad. Sci. 43: 281-283. , 1961. Polydora rickettsi, a new species of spionid polychaete from lower California. Pa- cific Sci. 15: 78-81. 1963. Comparison of Boccardia Columbi- ana Berkeley and Boccardia proboscidea Hart- man (Annelida Polychaeta). Bull. So. Calif. Acad. Sci. 62: 132-139. 1964. Polydora and related genera (Anne- lida, Polychaeta) from Eniwetok, Majuro and Bikini Atolls, Marshall Islands. Pacific Sci. 18: 146-159, 1977. Lecithotrophic larval development in Boccardia proboscidea Hartman. In Essays on Polychaetous Annelids in Memory of Dr. Olga Hartman, Edited by Reish, D. J. and Fauchald, K., Allan Hancock Found. Univ. South. Calif. Press, Los Angeles, pp. 347-371. Wu, B. L. and M. Chen, 1964. A new species of polychaete worm of the family Spionidae from Chushan Archipelago, East China Sea. Acta Zoo- taxonomica Sinica 1(1): 195-198 [In Chinese and English]. SURVEY OF THE VERTEBRATE FAUNA IN THE GRAMPIANS-EDENHOPE AREA OF SOUTHWESTERN VICTORIA W. B. Emison, J. W. Porter, K. C. Norris and G. J. Apps Fisheries and Wildlife Division, Ministry for Conservation, Arthur Rylah Institute for Environmental Research 123 Brown Street, Heidelberg, Victoria 3084 Abstract The Grampians-Edenhope Area of southwestern Victoria, Australia, contains four major land forms: woodland on sedimentary plains and sand dunes; heath, woodland and forest on sandstone ranges; woodland on plains and low hills of gravel and metamorphic rock; and woodland and forest, on basaltic plains. On the plains much of the land has been cleared for crops and pasture but the remnants of native vegetation support valuable wildlife communities and are especially important for nomadic birds. The Grampians ranges were least afTected by settlement and are the most important areas for wildlife in western Victoria. More than 50 plant communities and aquatic habitats are described and are used to discuss the distribution of 233 bird species, 54 mammal species, 40 reptile species, 14 amphibian species and 26 fish species. Most of these species are discussed individually: the birds in terms of abundance, distribution, habitat preference, breeding, feeding and conservation aspects; and the mammals and reptiles in terms of abundance, distribution, habitat preference, breeding and morphology. Introduction The Fisheries and Wildlife Division, Vic- toria, conducted wildlife surveys in the Gram- pians-Edenhope Area of southwestern Victoria (Fig. 1) between August 1974 and April 1975. Data collected during these surveys formed the basis of the fauna chapter in the Land Conservation Council's (LCC) report on their Southwest 2 Study Area (in prep.)- Because the report by the LCC simply docu- ments the existing fauna and is necessarily general, we publish this paper to provide a per- manent record of: (1) our assessments of areas important for fauna conservation; (2) data collected during the surveys on animal habitats and on individual vertebrate species; and (3) detailed faunal information generously contributed by naturalists residing in the survey area (see Acknowledgements). The following two sections of this paper describe the survey area, methods used and results obtained during the survey. Our assess- ments of areas important for fauna conserva- tion (by region) appear in the main text and are followed by appendices which contain the detailed information on flora and fauna. Survey Area Landform. The surveys were conducted over an area of approximately 16,000 km 2 (Fig. 1), of which about 20 per cent is still in public ownership. The survey area has been divided into four main regions based on Sibley's (1967) geomorphological descriptions: (1) the Eden- hope Region where plains of iluviatile and es- tuarine sediments are covered in many places by aeolian dunes and sheets of siliceous sands; (2) the Grampians Region (including the Black Range and Dundas Range) which consists of sandstone ranges uplifted into a cuesta forma- tion and rising to 800 m; (3) the Ararat Re- gion consisting mainly of undulating plains and low hills of slates and sandstones; and (4) the Hamilton Region where the plains and stony rises at Mt. Eccles and Mt. Napier have been formed by basalt lavas. Climate. The climate of the Grampians- Edenhope Region is characterized by warm temperatures, winter rains, summer droughts and prevailing northwesterly summer winds (Walter 1971). The average annual rainfall over the plains (i.e. Edenhope, Ararat and Hamilton Regions) 281 282 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS Figure 1— Place names and major blocks of Crown Land in the Grampians-Edenhope Area The survey area is divided into four regions: 1 = Edenhope Region; 2 = Grampians Region; 3 = Ararat Region; and 4 = Hamilton Region. GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 283 ranges between 450 and 700 mm, whereas that at Halls Gap in the mountains of the Grampi- ans Region is more than 900 mm. The wettest months (June, July and August) have a monthly average rainfall of 50-80 mm on the plains and more than 100 mm in the mountains. In sum- mer monthly averages are 20-30 mm on the plains and 35 mm in the mountains. Large, irregular fluctuations in annual rainfall oc- curred between 1940 and 1965 (Gibbs and Maher 1967). During this period droughts oc- curred over much of the survey area in 10 of the 25 years, with one drought lasting for three years (1943-1945). At the other extreme, wet years occurred during 8 of the 25 years. The winter during which our fauna surveys were conducted (1974) was relatively wet. Temperatures in towns on the plains aver- age 25-30°C (maxima) to 11-13°C (minima) during January and 11-13°C (maxima) to 3-5 °C (minima) during July. The mountainous area of the Grampians Region has lower average temperatures than those of the plains. Methods and Results Wildlife Habitats. The survey of the terres- trial wildlife habitats in the Grampians-Eden- hope Area consisted of one or more visits to 180 different sites to gather information on physical habitat, structure of the vegetation, relative numbers of each tree species, cover abundance and growth-form of most shrub species, and listing of common herbaceous plants. Data collected at each site are pre- sented as an annotated list (Appendix 1) and a table showing structure (Appendix 2) of 50 plant communities. A classification of terres- trial habitats (Table 1) was based on plant formations defined by Specht et al. (1974), and a simplified distribution of the major plant for- mations in the survey area is shown in Fig. 2. Nomenclature of plant species follows that of Willis (1962, 1972) and Appendix 3 lists scien- tific and common names of most plants col- lected during the survey and now held by the Fisheries and Wildlife Division, Victoria. Twenty-two species of Eucalyptus occur in large numbers in the survey area, and are the main plant species used in defining the plant communities in Table 1 and Appendix 1. Re- lationships between each pair of Eucalyptus species (whether in association or not in asso- ciation but in contact or not in contact) and also the relationship of each species or asso- ciation to each of the major regions of the sur- vey area are shown in Appendix 4. The survey of the aquatic habitats was more opportunistic than that of the terrestrial habi- tats and was mainly concerned with recording the flow, seasonality, depth and occurrence of wetland types in each survey region (Table 2) at the same time as surveying the vertebrate animals (usually birds) utilizing them. The land in the Grampians-Edenhope Area has been described by Blackburn and Gibbons (1956), Gibbons and Downes (1964) and Sib- ley (1967), in terms of land systems and land units. Salinities of some of the lakes in the sur- vey area are given in Williams (1967) and in Chessman and Williams (1974). By combining these data and our survey data a simplified wetlands distribution map has been drawn up for the survey area (Fig. 3). The following brief discussion of wildlife habitats in the Grampians-Edenhope Area is mainly based on the detailed survey informa- tion presented in the various figures, tables and appendices referred to above. The most prevalent plant formations in the Edenhope Region are shrublands and wood- lands. The shrublands are dominated by Brown Stringybark, heaths, (e.g. Leptospermum, Banksia) and sedges. These shrublands occur on the sandsheets and many hectares remain on Crown Land. Woodlands of Yellow Gum and River Red Gum once occurred continuously across the plains, and surrounded the patches of shrubland. Today most of these woodlands have been cleared for pasture and only scat- tered trees remain. Intrusions of mallee, typical of northwestern Victoria, occur near Mt. Ara- piles and east of Noradjuha but are small in area and do not constitute a major habitat in the Edenhope Region. Wetlands occur through- out the plains in the Edenhope Region except in those areas covered by sandsheets. These wetlands range from temporarily inundated, shallow depressions, such as occur on the gil- gaied soils north of Apsley, though country containing shallow swamps with sedge, reed 284 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS Figure 2 — Major vegetation formations in the Grampians-Edenhope Area. GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 285 Formation/ Structurally dominant species TABLE 1 Vegetation formations in the Grampians-Edenhope Area Community numbers in Appendix 1 Edenhope Grampians Ararat Hamilton Tall open forest Messmate Mountain Grey Gum Open forest Messmate Mountain Grey Gum Brown Stringybark Manna Gum Swamp Gum Candlebark Red Stringybark 7,11 11, 12 5, 8, 10 8, 9, 10 6, 8, 9, 10, 13 23 30,31 39 28 22 30, 31 40 Low open forest Messmate Brown Stringybark Long-leaf Box 14 15, 16 15,21 21 Woodland River Red Gum Yellow Box Yellow Gum Grey Box Black Box Manna Gum Slender Cypress Pine 33, 34, 36, 37 42 42, 43, 44, 45 46 47 25 49 33, 34, 36, 37 42 42,43 24 37 41 24 37 Shrubland Brown Stringybark Mallee eucalypts Salt Paper-bark 17, 18, 19, 20 48 50 Heath White Sallee Grampians Gum Scent-bark Shining Peppermint Scented Paper-bark Cross-leaf Honey-myrtle Common Heath 1 2,3,4 26 29 32 35 27 Grassland Native Grasses Pastures, Crops Present Present Present 37,38 Present 286 Flow/Salinity Standing Fresh Water Flowing Fresh Water W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS TABLE 2 Occurrence of wetlands in the Grampians-Edenhope Area Seasonality / Depth Edenhope Grampians Ararat Hamilton Temporarily Inundated Depressions Seasonal Swamps X Permanent Swamps and Shallow (<2 m) Lakes Permanent Deep (>2 m) Lakes Temporarily Inundated River Plains Permanent Slow-Flowing Rivers Permanent Fast-Flowing Streams Standing Saline Water Temporarily Inundated Depressions Seasonal Swamps and Saltpans Permanent Deep (>2 m) Lakes X X X X X X X X = Present in substantial numbers or area or both. and River Red Gum margins, into country containing deep lakes, mostly of a lunette for- mation. A chain of large saline lakes extends north from White Lake past Mt. Arapiles. Fig. 4 shows the important wetlands in the Eden- hope Region and Table 3 lists some of the more notable wildlife inhabitants and uses de- pendent on the wildlife resources. The major plant formations in the Gram- pians Region are open forest, low open forest, heath and woodland. The open forests occur primarily on the ranges and surrounding slopes and are dominated by trees of Brown Stringy- bark, Messmate and Mountain Grey Gum. Be- neath the trees in these forests heaths may cover large areas. Other shrubs in the open forests include small communities of meso- morphic shrubs along watercourses which are often fringed by dry sclerophyll shrubs. The most common herbs throughout the open forests are mid-height sedges which form a sparse to mid-dense layer. Tall dense sedges and fern layers occur only along wet gullies. Heath communities are found on shallow in- fertile soils in both sub-alpine situations and valleys. In the valleys, heaths merge into Scent- bark forests on dry margins and into Swamp Gum forests on wet margins. Woodland com- munities composed of Yellow Gum, Yellow Box, and River Red Gum are extensive in the Black Range and intrude along valleys into the Grampian Ranges. Another common forma- tion in the Black Range is low open forest dominated by Brown Stringybark, Messmate, Long-leaf Box and a tall, mid-dense growth of heaths (e.g. Oyster Bay Pine, Dwarf She- oak and Silver Banksia). The major wetlands of the Grampians Region are permanent fast- flowing streams and canals and four deep freshwater reservoirs (viz. Rocklands, War- took, Moora Moora and Bellfield). Seasonally flooded deposition plains occur along the Glen- elg River and Tea-tree, Dwyer, Fyans and Mt. William Creeks, the banks of which are usu- CRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 287 TABLE 3 Important uses or inhabitants of some wetlands in the Edenhope Region (see Fig. 4). Wetland Use or inhabitant Boikerbert Swamp Booroopki Swamp Boundary Swamp Bringalbert (Lake) Carchap (Lake) Carpolac (Lake) Centre Lake Charlegrark (Lake) Clarke (Lake) Clear Lake Collins Lake Connagorach Swamp Darragan Swamp Dollanoke (Lake) Donald Swamp Dumbopperty Swamp Gymbowen Lake Jaka Lake Jarracteer (Lake) Kanagulk (Lake) Karnak (Lake) Koynock (Lake) Kingcourt Swamp Leak Swamp MIga Lake Mill Swamp Mockinya Swamp Mullancoree (Lake) Murranbool Swamp North Lake No-where-else Swamp Ratzcastle (Lake) St. Marys Lake Hunting Potential Murray Cod holding site Hunting Limited hunting; Redfin fish- ing Waterfowl sanctuary Redfin fishing Wading birds Murray Cod hatchery Hunting Limited hunting; Redfin and Yabbie fishing Redfin fishing Limited hunting; Yabbie fish- ing Limited hunting Hunting; Redfin fishing; few Brolgas Limited hunting Limited hunting; Redfin fish- ing Yabbie fishing Important for waterfowl; limited hunting Hunting; Redfin fishing; few Brolgas Important for waterfowl; hunting Hunting; Yabbie fishing Hunting; Yabbie fishing Limited hunting Redfin fishing Limited hunting; Redfin fish- ing Yabbie fishing Limited hunting Hunting; Redfin fishing; few Brolgas Yabbie fishing Wading birds Potential Murray Cod hold- ing site Redfin fishing Limited hunting; Redfin fish- ing Toolondo Reservoir Wallace (Lake) White Lake Redfin and trout fishing Important for waterfowl; Redfin and trout fishing Wading birds ally covered by dense communities of tea-tree and River Red Gum. The Ararat Region contains two floristic in- trusions which do not occur elsewhere in the survey area. One intrusion consists of species belonging to the Long-leaf Box and Red Stringybark open forests occurring mainly near Ararat; the other, which occurs near Horsham, is woodland of Grey Box from the Northern Plains. The Ararat Region contains relatively few wetlands, although there are a few im- poundments (e.g. Lakes Fyans, Lonsdale, Tay- lor and Pine) and slow-flowing rivers. A few Black Box and lignum swamps and gilgaied de- pressions occur east of Horsham but most are now under cultivation. The Hamilton Region contains mainly agri- cultural land and remnants of the original vegetation of the volcanic plains. Our survey in this region was concentrated on the Stones State Faunal Reserve near Mt. Eccles. The reserve is covered by Manna Gum open forest and is atypical in relation to the rest of the vol- canic plains which were once covered by River Red Gum grassy open woodlands and grasslands. This region contains a few large sa- line lakes (e.g. Lake Condah) which are now mostly drained and some areas of slightly sa- line depressions occurring on the basalt plains. Birds. Terrestrial field work consisted pri- marily of surveying birds in as many of the 180 vegetation survey sites (Appendix 1) as pos- sible. Some surveys were not standardized be- cause the time, duration and effort of observa- tions varied between different sites and be- tween visits to the same sites. However, dur- ing all surveys we recorded actual or relative numbers of all avian species present and their utilization of particular structures within the habitat. Other surveys consisted of recording birds along lines (transects) 0-5 km long established in a number of vegetation survey 288 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS ■ ■ "W» ^* " Y / " ■■■'■■ C^ / ^5i2>' , *7 ' : " :%Jfc-Swimp > /I Temporarily inundoled depreitioni iwompl ond ihaUo— permanent lotts (utuolly lolinl) — I Few wellondl p.aieM Figure 3 — Distribution of wetlands in the Grampians-Edenhope Area. GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 289 9^"' moiu "* ■HatloHrfi *(.*«( CUUl'EaKlU #H0 WMEM-Otf ttMWP - .' ■■■■','r.-- HMTZCABTlt* • »taw im _ »Ot)NDA*T S¥ ai-tfl tJt*e l«i/««»i»too' twin? l«« OOtlWIOM* L«I Ntnt^NCCMirl tlMI MftlMCrffA ^houth Life B |cf»rw U/M ^. V i**£ K^NtauiK r 1 uociiivt \ \ 1 KinaCOUPl SMH' Figure 4 — Major lakes, swamps and reservoirs in the Edenhope Region. sites. Birds were recorded along most of these transects during spring, summer and autumn. Spotlighting from vehicles (usually in conjunc- tion with the mammal survey) was used for surveying nocturnal species such as owls and frogmouths. The wetland birds were recorded whenever they were encountered during field activities. However, in the Edenhope, Grampi- ans and Ararat Regions we conducted periodic censuses (spring and autumn, occasionally also summer) of waterbirds on selected swamps, lakes and rivers. Another large portion of our information came from the literature and pre- vious work done by staff of the Fisheries and Wildlife Division, Victoria, and other interested individuals (see Acknowledgements). Records held by the National Museum of Victoria, and by the Royal Australasian Ornithological Union were also important sources of informa- tion., From the above sources we have docu- mented the occurrence of 233 avian species in the Grampians-Edenhope Area. This number excludes seabirds that occur along the southern boundary of the survey area. One hundred and twelve species of birds occur in the forest, woodland, shrubland and heath for- mations (Table 4); 52 avian species are in- habitants of the semi-cleared pasture, grassland and suburban areas (Table 5); and 69 species occur in aquatic habitats (Table 6). Detailed information, including distribution, abundance, habitat preference, feeding and breeding, for each species appears in an annotated list (Ap- pendix 5). Nomenclature follows Condon (1975) for the non-passerines and Schodde (1975) for the passerines. Mammals. The published information avail- able on the mammal fauna of the Grampians- Edenhope Area is meagre and consists mainly of old imprecise popular accounts and a more recent general discussion by Wakefield (1974). Some specific and up-to-date information has been derived from specimens in the collections of both the National Museum of Victoria, and the Fisheries and Wildlife Division, Victoria, and field data have recently been available from two mammal survey groups operating in the State. However, most of the information presented here is derived from the field work (about 140 man-days) of two members of the survey team. Mammal trapping was conducted in 31 of the 180 vegetation survey sites (Ap- pendix 1), usually when the vegetation was being analysed. One hundred wire mesh cage traps (360 x 200 x 165 mm), baited with a 290 W, B. EMISON, J. W. PORTER, K. C. NORRTS and G. J. APPS mixture of peanut butter, honey and rolled oats, were set at each locality for two nights and the intervening day. Traps were checked on the first morning and captured animals were either retained or marked and released. All traps were reset and checked again on the sec- ond morning, after which they were removed TABLE 4 Checklist of avian species occurring in forest, woodland, shrubland and heath areas in the Grampians-Edenhope Area. See Appendix 5 for scientific names Brown Goshawk Collared Sparrowhawk Painted Button-quail Bush Stone-curlew Japanese Snipe Peaceful Dove Diamond Dove Common Bronzewing Brush Bronzewing Red-tailed Black Cockatoo Yellow-tailed Black Cockatoo Gang-gang Cockatoo Rainbow Lorikeet Musk Lorikeet Purple-crowned Lorikeet Little Lorikeet Cockatiel Budgerigah Swift Parrot Crimson Rosella Blue-winged Parrot Elegant Parrot Pallid Cuckoo Fan-tailed Cuckoo Black-eared Ouckoo Rufous-tailed Bronze- cuckoo Shining Bronze-cuckoo Powerful Owl Spotted Owl Barking Owl Masked Owl Tawny Frogmouth Owlet-nightjar Willie Wastail Spotted Quail-thrush Grey-crowned Babbler White-browed Babbler Rufous Songlark Superb Blue Wren Variegated Wren Southern Emu-wren White-browed Scrubwren Chestnut-rumped Hylacola Fieldwren Speckled Warbler Weebill White-throated Warbler Brown Thornbill Chestnut-rumped Thornbill BufT-rumped Thornbill Yellow Thornbill Striated Thornbill Varied Sittella White-throated Treecreeper Brown Treecreeper Red Wattlebird Little Wattlebird Spiny-cheeked Honey- eater Regent Honeyeater Blue-faced Honeyeater Yellow-faced Honeyeater Singing Honeyeater White-eared Honeyeater Yellow-tufted Honeyeater Fuscous Honeyeater White-plumed Honeyeater Black-chinned Honeyeater Spotted Nightjar Sacred Kingfisher Rainbow Bee-eater White-bellied Cuckoo- shrike White-winged Triller Scaly Thrush Southern Scrub-robin Rose Robin Pink Robin Flame Robin Scarlet Robin Red-capped Robin Hooded Robin Eastern Yellow Robin Crested Shrike-tit Gilbert's Whistler Golden Whistler Rufous Whistler Grey Shrike-thrush Satin Flycatcher Restless Flycatcher Rufous Fantail Grey Fantail Brown-headed Honeyeater White-naped Honeyeater Painted Honeyeater Crescent Honeyeater New Holland Honeyeater White-fronted Honeyeater Tawny-crowned Honey- eater Eastern Spinebill Mistletoebird Spotted Pardalote Striated Pardalote Silvereye Red -browed Firetail Diamond Firetail Olive-backed Oriole White-winged Chough Masked Woodswallow White-browed Wood- swallow Dusky Woodswallow Grey Butcherbird Pied Currawong Grey Currawong TABLE 5 Checklist of avian species occurring in grasslands, semi-cleared pastures and suburban areas in the Grampians-Edenhope Area. See Appendix 5 for scientific names Fork-tailed Swiftt Kookaburra Singing Bushlark* Common Skylark* Welcome Swallow Tree Martin Fairy Martin Richard's Pipit Black-faced Cuckoo-shrike Common Blackbird* Jacky Winter Brown Songlark Yellow-rumped Thornbill Southern Whiteface Noisy Miner White-fronted Chat European Goldfinch* European Greenfinch* House Sparrow* Emu Black-shouldered Kite Black Kite Whistling Kite Wedge-tailed Eagle Little Eagle Spotted Harrier Swamp Harrier Black Falcon Peregrine Falcon Little Falcon Brown Falcon Nankeen Kestrel Stubble Quail Brown Quail Little Button-quail Plains-wanderer Domestic Pigeon* Crested Pigeon GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 291 Galah Long-billed Corella Sulphur-crested Cockatoo Eastern Rosella Red-rumped Parrot Barn Owl Spine-tailed Swif tt * Introduced t Aerial Eurasian Tree Sparrow* Common Starling* Australian Magpie Lark Little Woodswallow Australian Magpie Australian Raven Little Raven TABLE 6 Checklist of avian species occurring in aquatic habitats in the Grampians-Edenhope Area. See Appendix 5 for scientific names Great Crested Grebe Hoary-headed Grebe* Little Grebe* Australian Pelican Darter Pied Cormorant Little Pied Cormorant* Black Cormorant Little Black Cormorant White-necked Heron* White-faced Heron Cattle Egret Large Egret Nankeen Night-heron Brown Bittern Glossy Ibis White Ibis Straw-necked Ibis Royal Spoonbill Yellow-billed Spoonbill Plumed Tree-duck Black Swan* Freckled Duck Cape Barren Goose Mountain Duck* Black Duck* Grey Teal* Chestnut Teal Shoveller Pink-eared Duck* White-eyed Duck Wood Duck* Blue-billed Duck Land Rail Water Rail Marsh Crake Spotted Crake Spotless Crake Black-tailed Native-hen* Dusky Moorhen* Swamphen Coot Brolga Painted Snipe Pied Oystercatcher Masked Plover* Banded Plover Red-kneed Dotterel Hooded Dotterel Double-banded Dotterel Red-capped Dotterel* Black-fronted Dotterel* Pied Stilt Banded Stilt Red-necked Avocet Greenshank Sharp-tailed Sandpiper Red-necked Stint Curlew Sandpiper Sanded in g Silver Gull Whiskered Tern Gull-billed Tern Azure Kingfisher Clamorous Reed-warbler Little Grassbird Musk Duck Golden-headed Cisticola White-breasted Sea-eagle * Recorded breeding in the Edenhope Region during 1974. from the site and set in another locality later in the day. In Appendix 6, reference is made to a trapping rate which is a measure of trapping susceptibility rather than abundance; neverthe- less it gives some indication of relative num- bers of small ground-dwelling mammals. The rate is simply the percentage of traps occupied by individuals of a particular species where the unit duration of a trapping period is one night. Spotlighting from vehicles at night allowed long distances to be covered and was particu- larly useful for recording large macropod spe- cies and many of the possums. Hand-held spot- lights were used while walking through small areas of bush and allowed quiet observations to be made on the relatively small arboreal mam- mals which can be overlooked while spotlight- ing from vehicles. Daylight observations were restricted mainly to large species such as kangaroos and deer. Other information came from finding dead animals, skeletal remains, tracks and identifiable faecal deposits. Anecdo- tal and often valuable information was supplied by interested local naturalists and farmers. During the survey we documented the oc- currence of 54 mammal species in the Gram- pians-Edenhope Area (Table 7); about 20% were introduced species. It appears that the mammal fauna of the plains in western Vic- toria has been altered by some species of rodents, dasyurids, bandicoots and macropods disappearing either directly or indirectly as a result of European settlement (Wakefield 1974). Some species still extant in western Victoria are presently in danger of extinction; these will be considered in more detail later in the dis- cussion. Detailed information on abundance, distribution, habitat preference, breeding and recent records for each mammal species in the survey area appears in an annotated list (Ap- pendix 6). Scientific nomenclature for the na- tive mammals follows Ride (1970) except for Potorous apicalis which is replaced here by P. tridactylus following Johnston (1973). 292 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS TABLE 7 Checklist of mammals in the Grampians-Edenhope Area. See Appendix 6 for scientific names Echidna Platypus Tiger Cat Tuan Yellow-footed Antechinus Brown Antechinus Swainson's Antechinus Mouse Dunnart Fat-tailed Dunnart Short-nosed Bandicoot Gunn's Bandicoot Koala Brush-tailed Possum Ring-tailed Possum Sugar Glider Squirrel Glider Yellow-bellied Glider Feather-tailed Glider Eastern Pigmy Possum Western Pigmy Possum Potoroo Eastern Grey Kangaroo Western Grey Kangaroo Red-necked Wallaby Brush-tailed Rock Wallaby Hare* Rabbit* * Introduced Bush Rat Black Rat* Swamp Rat Sewer Rat* Silky Desert Mouse Smokey Mouse Heath Rat House Mouse* Eastern Water Rat Fox* Cat (feral)* Pig (feral)* Red Deer* Goat (feral)* Sheep (feral)* Greater Long-eared Bat Lesser Long-eared Bat Bent-winged Bat Gould's Wattled Bat Chocolate Bat Little Bat Tasmaman Pipistrelle Large-footed Bat Little Broad-nosed Bat White-striped Bat Little Flat Bat Yellow-bellied Bat Reptiles and amphibians. Field work on these groups was usually confined to oppor- tunistic collecting while we were surveying either birds or mammals. The annotated list of reptiles (Appendix 7) and the checklist (Table 8) are based primarily on specimens in the National Museum of Victoria. However, a few days during the survey period were devoted to searching for reptiles in suitable habitats and a representative collection of species was lodged with the National Museum of Victoria. Specimens were collected by searching in hol- low logs, around rocky out-crops, under logs and in litter. The distributions of the 40 spe- cies of reptiles occurring in the survey area are consistent with the assessment by Rawlin- son (1971) that the reptile fauna is composed of three groups which correspond to the three major zoogeographic zones (Eyrean, Warm Temperate Bassian and Cool Temperate Bas- sian) in the survey area. Nomenclature for the reptiles follows Cogger (1975) except for Ano- tis maccoyi, Lampropholis delicata and L. guichenoti which follow Greer (1974) and the addition of Leiolopisma coventryi following Rawlinson(1975). The distributions in the survey area of the 14 species of amphibians known to occur there (Table 9) are based on recently published data (Brook 1975) and are shown in Appendix 8. Nomenclature for the amphibians follows Cogger (1975) except for the genus Crinia which is replaced here by Ranidella following Blake (1973). Fishes. During the present survey our in- volvement with terrestrial vertebrates precluded the gathering of much information on fishes. However, detailed studies and surveys are presently being conducted on selected lakes and streams in the Grampians-Edenhope Area by the Freshwater Fisheries Section of the Fisher- ies and Wildlife Division, Victoria, and their results will be presented elsewhere. A list of fishes occurring in the survey area appears in Table 10. The list does not include all estuarine and marine fishes which may occur along the southern boundary of the study area. Nomen- clature follows Barnham (in prep.). TABLE 8 Checklist of reptiles in the Grampians-Edenhope Area Chelidae (Side-necked Tortoise) Chelodina longicollis (Long-necked Tortoise) Agamidae (Dragon Lizards) Amphibolous barbatus (Bearded Dragon) Amphibolous diemensis (Mountain Dragon) Amphibolous muricatus (Jacky Lizard) Amphibolous pictus (Painted Dragon) Gekkonidae (Geckos) Diplodactylus tessellatus (Tessellated Gecko) Phyllodactylus marmoratus (Marbled Gecko) Pygopodidae (Snake-lizards) Delma inornata Delma impar GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 293 Scincidae (Skinks) Anotis maccayi Cryptoblcpharus boutonii Ctenotus robustus Egernia luctuosa (Mourning Skink) Egernia saxatilis (Black Rock Skink) Egernia whitii (White's Skink) Hem iergis dtcresiensis Hetniergis peronii Lampropholis delicata Lampropholis guichenott Leiolopisma entrecasteauxii Leiolopisma trilineata Leiolopisma coventryi (Coventry's Skink) Lerista bougainvillii Morcihia adelaidens'ts Morethia boulengeri Morethia obscura (Ocellated Skink) Sphenomorphus tympanum Sphenomorplius sp. TUiqua nigrolutea (Blotched Hlue-tongued lizard) TUiqua scincoides (Eastern Blue-tongued Lizard ) Traehydosaurus rugosus (Shingle-back) Varanidae (Monitor Lizards) Varanus gouldii (Sand Monitor) Varanus varius (Lace Monitor) Elapidae (Elapid Snakes) Austrelaps superba (Lowlands Copperhead) Drysdalia coronoides (White-lipped Snake) Notechis scutatus (Eastern Tiger Snake) Pseudechis porphyriacus (Red-bellied Black Snake) Pseudonaja textilis (Eastern Brown Snake) Unechis flagellum (Little Whip Snake) Typhlopidae (Blind Snakes) Typhlina proxima TABLE 9 Checklist of amphibians in the Grampians-Edenhope Area Hylidae (Tree Frogs) Litoria ewingii (Brown Tree Frog) Litoria lesueurii (Lesucur's Frog) Litoria raniformis Leptodactylidae (Southern Frogs) Geocrinia laevis Geocrinia victoriana Limnodynastes dumerilii (Eastern Banjo Frog) Limnodynastes peronii (Brown-striped Frog) Limnodynastes tasmaniensis (Spotted Grass Frog) Neobatrachus centralis (Trilling Frog) Neobatrachus pictus (Meeowing Frog) Pseudophryne bibronii (Brown Toadlet) Pseudophryne semimarmorata (Southern Toad- let) Ranidella parinsignifera Ranidclla signifcra (Common Eastern Froglet) TABLE 10 Checklist (incomplete) of fishes in the Grampians-Edenhope Area Mordaciidae Geotria australis (Pouched Lamprey) Relropimiidae Retropinna semoni (Australian Smell) Anguillidae Anguilla australis (Short-finned Eel) Mugilidae Aldrivhctta forsteri (Yellow-eyed Mullet) Elcotridae Philypnodon grandtcepS (Hiy-IiL*;idcd Gudgeon) flypscleotris eonipressus (Carp Gudgeon) Ilypscleotris klunz'mgeri (Western Carp Gud- geon) Gadopsidac Gadopsis marmoratus ( Blac k fish ) Plotosidac Tandanus tandanus (Caifish) Bovichthyidao Pseudaphritis urviili (Tupong) Nannopercidae Nannoperca australis (Pigmy Perch) Plcclroplitidac Plectroplitss ambiguus (Golden Perch) Maccullochcllidae Maccullochella pccli (Murray Cod) Macquariidae Percolates colonorum (Estuary Perch) Macquaria australasica (Macquarie Perch) Sparidao Acanthopagrus butcheri (Black Bream) Galaxiidae Galaxias maculatUS (Common Galaxiid) GalaxiaS olidus (Ornate Mountain Galaxiid) Galaxiella pusillus (Dwarf Galaxiid) Percidae Perca jluviatilis (English Perch or Rcdfin)* Pocciliidae Gambusia afjinis (Mosquito Pish)* Salmonidao Salmo gairdneri (Rainbow Trout)* Salmo trutta (Brown Trout)* Cyprinidac Carassius auratus (Goldfish)* 294 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS Carassius carassius (Crucian Carp)* Tinea tinea (Tench)* * Introduced Discussion The first European to explore the Gram- pians-Edenhope Area was Major Thomas Mit- chell, who climbed Mt. William in the Gram- plans in July 1836 (Mitchell 1839). His party then traversed the northern portions of the Ararat and Edenhope Regions before striking southwest past White Lake to the Glenelg River and down the river to the ocean, which they reached on 20 August 1836. Later that year they passed through the Hamilton Region, climbing both Mt. Napier and Mt. Abrupt (Grampians Region) before striking northeast towards Sydney. Mitchell extolled the beauty and the richness of the soil of the plains coun- try, and his descriptions of the physical sur- roundings suggest that much of the plains was covered by relatively open, grassy woodlands. Today most of the areas through which Mit- chell travelled are crop or pasture lands with few of the woodland trees and native grasses remaining. The mountainous areas of the Grampians Region have been less affected by European settlement, and the species of birds inhabiting the Grampians Region in 1891 (Anon. 1892) were essentially the same as those recorded there during the present survey. Modification of wildlife habitats on the plains has undoubtedly benefited some ani- mals, particularly domestic mammals (cattle, sheep, horses) some introduced bird species (starling, pigeon, sparrow, goldfinch) and some native bird species (quail, some cocka- toos, pipit, magpie). However, the loss of such habitat is also detrimental to many species of native wildlife, and should most of the remain- ing native trees, shrubs and grasses be cleared, a large portion of the original complement of wildlife species would disappear. Most standing surface water in the survey area is in the Edenhope Region, although a few large freshwater and saline lakes and im- poundments occur in the other regions. The Grampians Region supports most of the fast- flowing streams in the survey area with a few slow-flowing rivers occurring in the other re- gions. Many of these bodies of water have been modified by draining, clearing of native vege- tation from the catchments, stream or lake modifications, diversions, building of dams, ir- rigation and dumping of waste. However, many of the wetlands still support large assemblages of vertebrates but the continued modification of such waters will probably result in a de- crease in the diversity of species inhabiting them. Despite extensive habitat modification throughout the survey area the terrestrial and aquatic areas of Crown Land support valuable vertebrate communities. The wildlife values of the four survey regions (Fig. 1) of the Gram- pians-Edenhope Area are discussed below. Wildlife values of the Edenhope Region. Remnants of the original vegetation indicate that most of the alienated land in the Edenhope Region (unshaded in Fig. 1) originally sup- ported woodlands of River Red Gum, Yellow Gum, Yellow Box and, possibly, Grey Box. Because these woodlands occurred on the deeper more fertile soils, they were cleared for pasture and crops. Consequently, most of these woodlands have now gone, and only roadside remnants and a few plantations of small River Red Gums (along the margins of some blocks of Crown Land) remain of what was once the major terrestrial habitat in the region. It is impossible to determine the number of animal species which were common in these wood- lands before European settlement, but there are indications that, at least, some mammal species have recently disappeared (Wake- field 1974). The results of our survey indi- cate that at present about 100 terrestrial verte- brate species occur in the woodland rem- nants of the Edenhope Region. These species include only those which are either common throughout the year or rely on the woodlands during some periods of the year (e.g. for feed- ing or nesting). Bird species make up about 60 per cent of the total number of common species in the woodlands, mammals about 25 per cent and reptiles and amphibians the re- maining 15 per cent. Approximately 5 per cent are species alien to Australia which, in gene- ral, exert a disproportionate influence on the GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 295 vegetation and on the well-being of many of the native animal species. The populations of many of the native animals will decrease and some will disappear if the woodland remnants continue to be modified or destroyed. We sug- gest that at least 15 per cent of the total num- ber of species are either already, or will soon be, experiencing difficulties in maintaining their population numbers in the Edenhope Region (see annotated lists for details). These include 13 species of birds (Bush Stone-curlew, Rain- bow Lorikeet, Musk Lorikeet, Little Lorikeet, Purple-crowned Lorikeet, Swift Parrot, Long- billed Corella, Barking Owl, Grey-crowned Babbler, Blue-faced Honeyeater, Painted Honeyeater, Regent Honeyeater and Grey Butcherbird), one mammal species (Tuan) and one reptile species (Lace Monitor). Most species whose populations may be in difficulties, as well as many of the other native woodland animals, fall into one or more of the following categories: (1) birds such as lorikeets and honeyeaters which move to the woodlands seasonally to feed on pollen, nectar and insects when the River Red Gums, Yellow Gums and Grey Box are flowering; (2) animals such as corella, cockatoos, owls, possums, tuans and Lace Monitor which require hollows in the trunks or limbs of large trees for breed- ing or sheltering; and (3) birds such as curlews and babblers which forage on the ground and appear to depend on woodland with relatively unchanged native undergrowth. Maintenance of woodland habitats in the Edenhope Region will require the cooperation of government agencies and private land holders. As a first step, a detailed botanical study of the woodlands of the entire Gram- pians-Edenhope Area should be conducted to ascertain: (1) where the various woodland com- munities occurred originally; (2) the species composition in each community at present; and (3) the best means of ensuring that adequate mature trees and associated plants are present and that regeneration is occurring. Those woodland remnants still on Crown Lands (e.g. margins of the larger Crown Land blocks, roadside reserves, etc.) should be carefully managed to ensure that enough large trees re- main to support populations of nectar-feeding v animals, that enough over-mature trees remain to provide holes for nests and shelter and, if possible, that the understoreys of some wood- lands be restored to their original composition. Ways of encouraging private landholders to plant appropriate woodland species when land- scaping, creating shelter belts or even speci- fically planting areas for native wildlife should also be explored. The vegetation on most of the large Crown Land blocks (shaded in Fig. 1) in the Eden- hope Region is mainly Brown Stringybark shrubland. These shrublands were not alien- ated during early settlement because the soils were too poor to support crops or pasture. Consequently, these Crown Land blocks now support a relatively unmodified habitat, al- though grazing and fires have probably some- what altered the ground-cover. The species composition of the vertebrate wildlife is dif- ferent from that occurring in the adjacent pas- ture/woodland areas and consists of about 65 commonly occurring species (50 per cent birds, 30 per cent mammals and 20 per cent reptiles/ amphibians). The relatively dense undergrowth on some of the blocks appears to be an import- ant structural feature influencing the distribu- tion of some of the wildlife species, particu- larly some thornbills, honeyeaters and the Western Pigmy Possum. The Red-tailed Black Cockatoo occurs in these blocks and we saw large numbers in the Jilpanger Block. The range of this cockatoo in Victoria has de- creased since European settlement, and the large Crown Land blocks in the Edenhope Region appear to be one of the last strong- holds of this species in the State. Also the Little Wattlcbird sometimes occurs in large numbers in dense stands of banksias, which provide an important food source. Occurring together within some of the Crown Land blocks in the Edenhope Region are typical inland (Eyrean) species of reptiles (Painted Dragon and Ocellated Skink) and temperate (Bassian) species of reptiles (Black Rock Skink). The shrublands are generally rich in reptiles which attain maximum diversity in blocks such as Mt. Arapiles where rocky outcrops adjoin woodland, shrubland and low open forest. The Silky Desert Mouse occurs 296 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS along the high sand ridges in areas which have not suffered heavy grazing or burning. Mouse Dunnarts are probably widespread in the shrub- lands but their numbers are difficult to assess because they are not usually captured in con- ventional traps. Other native mammals occur- ring in this habitat include the Echidna, Yellow- footed Antechinus, Western Grey Kangaroo and Red-necked Wallaby. To ensure perpetuation of the assemblage of wildlife species now present in these shrublands in the Edenhope Region we suggest that as many as possible of the large Crown Land blocks be retained intact and in public owner- ship. A more detailed study of the fauna of these areas is required to ascertain the import- ance of each Crown Land block, particularly in relation to similar vegetation further south. Several of the blocks have fringes of woodland which, if at all possible, should be retained to provide wildlife habitat. The intrusions of mallee vegetation into the Mt. Arapiles, Noradjuha and Wonwondah North areas are small but constitute some of the southern-most extensions of this habitat into western Victoria. Species of birds such as the Southern Scrub-robin, Variegated Wren, Gilbert's Whistler, White-fronted Honeyeater and Spiny-cheeked Honeyeater are at the southern limit of their range in western Vic- toria in these small intrusions of mallee. These wildlife species are more common further north but the Mt. Arapiles block, in particular, con- tains not only these animals, which have ad- apted to a mallee habitat, but also wildlife ad- apted to living in woodlands, low open forests, or on exposed rocky outcrops because of the diversity of habitats in this one block. The pastures and croplands in the Eden- hope Region support populations of most of the avian species listed in Table 5. The open terrain is suitable for hunting over by such predatory birds as eagles, kites, harriers and falcons. Some members of the parrot and cockatoo family (particularly Long-billed Corella, Galah and Sulphur-crested Cockatoo) forage in the pastures for roots and bulbs. Other wildlife species particularly well-ad- apted to survival in this open habitat include bird species (such as some quails and larks, Richard's Pipit, Yellow-rumped Thornbill, White-fronted Chat, Jacky Winter, House Sparrow, goldfinch, starling, magpie and ra- ven), mammal species (such as Fat-tailed Dun- nart, Rabbit, House Mouse, Fox and Cat) and reptile species (such as Eastern Blue-tongued Lizard, Shingle-back, and Eastern Brown Snake). During periods of high rainfall many of the pastures are covered either entirely or partially by standing water. These temporary wetlands are favoured foraging areas for some of the large waders (e.g. Large Egret, White-necked and White-faced Herons), the small grebes and some of the ducks (e.g. Black Duck and Grey Teal). The Brolga is also a conspicuous spe- cies along the margins of the open pasture and aquatic areas in the Edenhope Region. This species still breeds in the region, but its long- term survival in the area is not certain. A de- tailed study of the habitat requirements of the Brolga in western Victoria would be a first step towards a management plan needed to ensure their survival. The importance of the Edenhope Region in providing a large proportion of the aquatic habitat in western Victoria is apparent in Fig. 2. Although some avian species utilize im- poundments and temporarily flooded agricul- tural land, their main habitats are the large areas of natural fresh and saline wetlands, most of which are unprotected and unmanaged in the Edenhope Region. Our survey only touched on some of these important wetlands and a more detailed study is required to determine priorities for conservation and management. Requirements for breeding, feeding and shelter of a particular avian species may be fulfilled only by a wide range of aquatic conditions. Therefore, a regional management plan must aim at preserving maximum diversity of wet- land habitats. Hundreds of saline and freshwater lakes, swamps and depressions which hold water tem- porarily occur throughout the western two- thirds of the Edenhope Region. The spring and early summer of 1974 were unusually wet, and standing water was widespread throughout the region. Waterbirds took advantage of these con- ditions and nests and broods of 15 different GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 297 species were documented in the region (Table 6). However, the area of nesting habitat has been considerably reduced by the clearing of land down to the margins of some lakes and by the draining of many of the swamps and sea- sonal depressions. Many of these lakes and swamps provide public duck hunting areas and a few of the more important ones are listed in Table 3 and shown in Fig. 4. The permanent water bodies are probably important refuges for waterfowl during periods of low rainfall and after most temporary bodies of water have dried up, as occurred in 1972 (Mitchell 1973). During the non-breeding sea- son the mud flats of some of these waters also provide foraging areas for six species of inter- continental migrants, of which two breed in the Arctic (Red-necked Stint and Curlew Sand- piper), three breed in Eurasia (Japanese Snipe, Greenshank and Sharp-tailed Sandpiper) and one breeds in New Zealand (Double-banded Dotterel). The narrow strip of land containing saline lakes and swamps, which extends from Mitre Lake on the northern boundary of the Edenhope Region through White Lake near the southern boundary (Fig. 2), may be important to such wader species as Pied and Banded Stilts and Red-necked Avocets. Very large flocks of these waders, as well as Black Swans and various species of ducks, have been re- ported from some of these lakes (particularly North, Centre and White Lakes near Douglas). We suggest that detailed and regular observa- tions be made on the waders especially. The wide range of aquatic habitats is, in part, also responsible for the widespread occurrence in the Edenhope Region of at least five species of amphibians (and four other less widely distribu- ted amphibians) as well as supporting other vertebrate species such as the Eastern Water Rat and the Long-necked Tortoise. Some of the permanent lakes in the Eden- hope Region (Table 3 and Fig. 4) support an important amateur freshwater fishery based mainly on Redfin (a few lakes also support trout populations). Amateur fishing for yab- bies is popular, but at present they are not be- ing commercially exploited in the region al- though the potential for such an industry exists. The Freshwater Fisheries Section of the Fisher- ies and Wildlife Division, Victoria, has re- cently established a Murray Cod hatchery on Lake Charlegrark. Two nearby bodies of per- manent water, Booroopki Swamp and No- where-else Swamp (Morea Block), may be important for use as alternate Murray Cod holding sites as the project develops. Wildlife values of the Grampians Region. The area of Crown Land remaining in this region exceeds 200,000 ha, most of it being continuous through the Grampians Ranges (e.g. Victoria, Scrra and Mt. Difficult Ranges) and the Black Range (Fig. 1). The diverse habitats contained within this large Crown Land area support a correspondingly large and diverse assemblage of native vertebrate species. To maintain this diversity it is important that the continuous area of public land be managed as a single functional unit rather than arbit- rarily dividing it into a series of small discrete units and managing each as an entity. Manage- ment of this Crown Land area as a single unit will enhance the chances of survival of those species of wildlife whose life cycles require the utilization of two or more distinct habitats as well as those species which attain their highest densities in ecotones. The Crown Land west of the Victoria Range probably contains what is now the largest re- maining area of woodland (River Red Gum, Yellow Gum and Yellow Box) in southwestern Victoria (Plate 19, Fig. 1). Although this plant formation was one of the major habitats in the survey area before European settlement, it has now nearly gone and the woodland area west of the Victoria Range covers only slightly more than 20,000 ha. These woodlands are the re- mains of a vast area of woodland which originally also covered most of the Edenhope Region to the west. Consequently, remarks made previously on the fauna of the woodlands in the Edenhope Region apply equally to the fauna of woodlands in the Grampians Region. However the woodlands in the Grampians re- gion represent the most important area of this habitat remaining in southwestern Victoria and one major aim of their management should be to provide food, breeding sites and shelter for the large assemblage of vertebrate animals (some 100 species) which inhabit them. We 298 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS strongly suggest that an assessment of some of the structural features known to be important to vertebrate species (e.g. mature trees with large hollows, dense undergrowth, decaying logs) be made to ascertain how these woodlands can be managed to provide a more suitable en- vironment (i.e. similar to pre-European wood- lands) for many of the native species of wild- life. Heath communities, widespread throughout the Crown Lands in the Grampians Region (Fig. 2), are inhabited by about 80 species of commonly occurring vertebrate animals (55 per cent birds, 30 per cent mammals and 15 per cent reptiles/amphibians). Lowland heaths, including the ecotone of heath under open forest, and sub-alpine heaths are the most ex- tensive heath types occurring in the Gram- piaiis. The lowland heaths (particularly those in the Victoria Valley) appear to be the northern extremity in Victoria of a predominantly coastal heath type which extends inland along the Glenelg River to the Grampians (Plate 19, Fig. 2). In these heaths such species as the Southern Emu Wren and the Heath Rat reach the northern limits of their distributions in Victoria. Similarly, the most northerly popula- tions of the Short-nosed Bandicoot and the Potoroo in Victoria occur in heaths in the Grampians Region, but these heaths are usually under large trees and generally adjacent to thick gully vegetation. The importance to na- tive mammals of this ecotonal zone in the Grampians has been discussed by Seebeck (1976). A single specimen of the Mourning Skink, a rare species whose centre of dis- tribution appears to be coastal Victoria, has been collected from a heathy swamp in the Victoria Range. Large numbers of three spe- cies of honeyeaters (New Holland, Tawny- crowned and Eastern Spinebill) move into the lowland heaths in the spring to feed on the nectar from the flowers of the Common Heath. The fauna of the heath communities in the Vic- toria Valley is of special interest to biologists because of its similarities to the vertebrate fauna in the heaths occurring along the coast of Victoria. The perpetuation of these plant communities (and their associated fauna) in the Grampians should be a major aim in the management of this region. The management of these heaths may require periodic burning to maintain numbers and diversity of the verte- brate species inhabiting them (Cockburn 1975). However, this should be the subject of careful study and experimentation before wildlife man- agement based mainly on controlled burning is introduced. The number of the Victorian endemic Smokey Mouse recorded during our 1974-75 survey of the sub-alpine heaths in the Gram- pians Region exceeded the total number re- corded in Victoria prior to that time and illus- trates the likely importance of the Grampians area for this species' conservation. Most of the bird species which occur in the lowland heath and five species of reptiles also inhabit the sub- alpine heaths. The heaths are one of the nest- ing areas of the Flame Robin in western Vic- toria. Crescent Honeyeaters were common in the heaths near the summit of Major Mitchell Plateau in August 1974. The open and low open forests, including the vegetation mapped as scrub by the Forests Commission of Victoria (undated), form the most extensive vegetation formation occurring in the Grampians Region (Fig. 1). At least 100 species of wildlife (55 per cent birds, 25' per cent mammals and 20 per cent reptiles/amphi- bians) commonly occur in these formations. Most of these species occur in other forma- tions in the survey area, but the open forests are particularly important because of the rela- tively large area they cover. The chances of maintaining viable populations of many of the vertebrate species now occurring in southwest- ern Victoria will be enhanced if the areas of low open and open forest in the Grampians remain large. Uncommon species, such as Powerful and Barking Owls, have recently been recorded in the open forests of the Gram- pians. The forests also support the largest western Victorian populations of such species as Gang-gang Cockatoo and Koala and the low open forests contain large numbers of Scaly Thrushes and New Holland Honeyeaters. At present this region is a popular recrea- tion and tourist area which will probably be- come even more important in the future. GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 299 Recreation and tourism presently is com- patible with flora and fauna conservation. However, as the number of people involved in such activities increases, conflicts between re- creation and conservation will appear, par- ticularly in local areas where concentrations of people occur because of easy access, scenic qualities, picnic facilities and presence of wildlife. We also suggest that the formulation of wildlife management policies for this Crown Land should take into account the importance of mature or overmature trees for those species requiring nest hollows and dens. Their precise requirements should be the subject of more de- tailed studies. The Grampians Region contains the most ex- tensive and scenic sandstone mountain ranges, including numerous cliffs and caves, in western Victoria. Thousands of tourists visit the Gram- pians each year partly because of the spectacu- lar scenery and panoramic views provided by these mountain ranges. However, these ranges are also the habitat of the only known Brush- tailed Rock Wallaby colony remaining in western Victoria and they support a large breeding population of Peregrine Falcons. Both these species have been proposed for proclama- tion as Notable Species under the Wildlife Act 1976; the wallaby because of its serious de- cline in this State since 1900 and the falcon because of its decline in the northern hemi- sphere and fears that the local population may suffer a similar decline. Management authori- ties should avoid constructing roads, carparks, camp sites, picnic areas, nature walks etc. in areas which are important to these two species. Other species which occur in these rock com- munities include Nankeen Kestrel, Welcome Swallow, Eastern Pigmy Possum, some bats, Marbled Gecko and Black Rock Skink. The few small scattered areas of tall open forest in the Grampians Region are important as the western limit of this habitat type in Vic- toria. The largest area of tall open forest in the Grampians Region occurs along Dairy Creek near Silverband Falls. Although we recorded only about 30 species of wildlife occurring in this habitat, it is important in supporting either migrants or relatively large populations of at least five of these species (Rufous Fantail, Satin Flycatcher, Crescent Honeyeater, Smokey Mouse and Coventry's Skink). Two pine plantations (east of Wartook Re- servoir and at Billywing) have been established in the Grampians Region by the Forests Com- mission of Victoria. Our survey did not include a census of vertebrate animals in these planta- tions, so we cannot compare their density or diversity with those of the adjacent native forests. However, a recent survey of birds in pine and native forests in New South Wales showed 'that the loss of diversity of species and actual numbers of individuals is very great when native forest is removed for pines' (Dis- ney and Stokes 1976). We strongly recom- mend that careful consideration be given to the effects on native fauna before further ex- pansion of existing pine plantations or estab- lishing new plantations in the Grampians Re- gion. The main aquatic habitats in the Grampians Region are man-made reservoirs (e.g. Rock- lands Reservoir, Moora Moora Reservoir, War- took Reservoir, Lake Bellfield) and permanent fast-flowing streams and canals. There are also a few permanent swamps in the plains adjacent to the Grampians Ranges (e.g. Bryans, Mar- neys, Brady s) as well as some short portions of slow-flowing creeks and rivers (e.g. Green Creek and the upper reaches of the Wannon and Glenelg Rivers) (Plate 20, Fig. 3). The reservoirs and slow-flowing rivers pro- vide a large area of habitat for fishes. During our recent surveys seven Dwarf Galaxiids were found in Green Creek, a slow-flowing stream with a silt or mud bottom at an altitude of about 200 m. The only other records of this species from eastern Australia come from a few other areas in southwestern Victoria (Franken- berg 1969; Chessman and Williams 1974) and from restricted localities around Western Port. Therefore, Green Creek presents an excellent opportunity for preservation of a population of this little known species. Also another native species, the Pigmy Perch, is common in slow- flowing streams around the Grampians. Other fish species include Redfin, Brown and Rain- bow Trout, Tench and Carp. The importance of these reservoirs and slow-flowing streams to the avifauna is not well known, but they prob- 300 W. B. EMISON, J. W. PORTER, K. C. NORRJS and G. J. APPS ably provide refuge for many waterbirds dur- ing droughts. Aquatic mammals, such as the Platypus and Eastern Water Rat, and reptiles, such as the Long-necked Tortoise, also occur in some of these waters. The permanent fast-flowing streams and canals in the Grampians Ranges support a fish fauna which is, as yet, relatively undisturbed. At present Blackfish and the Ornate Mountain Galaxiid inhabit some of these streams, but the effects of habitat alteration or competition with introduced species on populations of these two native species should be examined. Many species of mammals, birds, reptiles and amphi- bians depend on these permanent streams for water when other temporary sources have dried up. Therefore any water management plans for this part of Victoria should take into account the need to maintain adequate stream flows and standing water suitable for wildlife throughout the year. Bryans Swamp, a shallow but relatively per- sistent body of water, is a State Faunal Reserve in the Grampians Region. It was surveyed on three occasions during 1974 and 1975, and 19 species of birds were recorded. Changes in the species composition of the avifauna ap- peared to be related to seasonal variations in water level. The avifauna on this swamp also appears to be highly variable between years, and as many as 500 White-necked Herons and 1000 White-faced Herons as well as Royal Spoonbills, Glossy Ibis, and Whiskered Terns have been reported (Mitchell 1973). Wildlife values of the Ararat Region. Most of the land in this region has been cleared and is in private ownership. The few small remain- ing areas of Crown Land are either woodland (Plate 20, Fig. 4) or open forest. All these areas are valuable because, as Parsons et al. (1972) point out, 'very little is known of the ecology of the Grampians Plains and the destruction of the last remnants of native vegetation would make future study impossible'. The Crown Land blocks around Lake Lons- dale (e.g. Illawarra, Mokepilly and Lonsdale Blocks in Fig. 1) are predominantly woodlands and are inhabited by a vertebrate fauna which once occurred throughout the plains surround- ing the Grampians but which is now restricted to a few isolated Crown Land areas. These woodlands contain many of the 100 (approxi- mately) vertebrate species that occur in similar woodland remnants in the Edenhope and Grampians Regions. Additionally, the Moke- pilly Block contains a mixture of open forest and woodland vegetation and consequently has a more diverse fauna than either the wood- land areas to the east or the open forest areas to the west. The woodlands of Yellow and Long-leaf Box and Red Stringybark in the Jallukar Block also contain some avian species (e.g. Speckled Warbler, Fuscous Honeyeater, Yellow- tufted Honeyeater, etc.) which attain their largest Victorian populations in the box- ironbark vegetation further east. However, their populations in the Jallukar Block are large relative to the other Crown Land blocks in the Grampians-Edenhope survey area. The woodlands in the Ararat Region orig- inally supported populations of the Squirrel Glider (Plate 21, Fig. 5), a species uncommon in Victoria. Two records exist for the Ararat Region: 5 km northwest of Stawell in 1970; and at Dadswell Bridge in 1968. This species may still be extant in the Ararat Region, but will require research to determine its status and to enable recommendations to be made on ap- propriate actions needed to maintain popula- tions of this species in the survey area. The Lace Monitor has also been recorded from the woodlands of the Illawarra Block. The open forest of Red Stringybark and Long-leaf Box which originally extended into the eastern portion of the survey area is now mainly confined to One Tree Block near Ararat. Most wildlife species which commonly occur in this block are found in the open forests on other blocks of Crown Land in the survey area. However, the floristics of this area differ from those of most other blocks in the survey area and a more detailed fauna survey might reveal subtle differences in species com- position of the wildlife. We endorse the recommendations implicit in the statement by Parsons et ah (1972) that 'it is necessary to reserve as many of the surviving remnants of the plains vegetation as possible, to ensure adequate preservation of the unique vegetation of the whole Grampians area'. We GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 301 point out that because the vegetation on these areas of Crown Land is different from that in the Grampians Ranges, the vertebrate fauna is also markedly different and further supports the suggestion that these woodland and open forest areas be preserved. Pastures in the Ararat Region support most of the bird species listed in Table 5. Introduced vertebrate species such as House Sparrow, European Goldfinch, Common Starling, Do- mestic Pigeon, Rabbit, House Mouse, Fox and Cat, compete with or prey upon many of the native vertebrate species and are undoubtedly responsible for reductions of some populations. The impact that these introduced species have on native vertebrate populations is little known and a study on the interactions between some of these species is required. In the Ararat Region, the lakes and reser- voirs containing permanent or relatively per- sistent standing water include Alexandra Lake, Dock Lake, Lake Fyans, Green Lake, Lake Lonsdale, Norval Dam, Pine Lake and Taylor Lake. These waters support populations of Blackfish, Redfin, Carp, Tench, Brown and Rainbow Trout and, rarely, Murray Cod (Tun- bridge and Rogan 1976). Golden Perch have recently been introduced into Green Lake (Barnham, Fisheries and Wildlife Division, pers. comm.). The few permanent streams (e.g. Fyans Creek, MacKenzie River and Mt William Creek) in this region support a similar fish fauna but probably have few Tench or Murray Cod. The permanent waters of the reservoirs, streams and some swamps (e.g. Mt. William Swamp) probably provide refuge for many of the species of birds listed in Table 6. Some of the reservoirs still have many dead trees standing around their margins. These trees are important roosting and nesting structures for some parrots and cockatoos (e.g. Musk Lori- keet, Sulphur-crested Cockatoo, Long-billed Corella, Galah, Eastern Rosella, Red-rumped Parrot) and some birds of prey. However, once the trees decay and fall they will not be re- placed. Platypus, Eastern Water Rat, Long- necked Tortoise and several species of amphi- bians are present in some of the permanent waters in the region. Wildlife values of the Hamilton Region. Be- cause most of this region is now in private ownership, the few remaining areas of Crown Land supporting native vegetation are very important. Before European settlement this region was covered by two main plant forma- tions: woodlands of large trees with grassy undergrowth; and treeless grasslands of the volcanic plains (Fig. 2). The Stones and Mt. Napier Blocks (Fig. 1) were originally rela- tively small rocky areas of volcanic craters and associated lava flows which supported Manna Gum open forest; these areas were surrounded by deeper soils on which the woodlands grew. The woodlands were replaced by exotic grasses and other crops soon after settlement, and the land was alienated, leaving only the rela- tively infertile areas (Stones and Mt Napier) in public ownership. Today, although modified by fires and grazing, the vegetation on these two Crown Land areas is still composed of many of the original plant species. The Stones Block includes both the Stones State Faunal Reserve (5241 ha) and Mt. Eccles National Park (395 ha). The wildlife reserve supports a large assemblage of bird species, most of which occur commonly in other open forest habitats. However, the mam- malian fauna is distinctive and few areas, if any, support a ground fauna composed of the Tiger Cat, Brown Antechinus, Swainson's An- techinus, Eastern Grey Kangaroo, and Bush Rat, and an arboreal fauna which probably includes the Koala, Yellow-bellied Glider, Brush-tailed Possum, Sugar Glider and Feather-tailed Glider. The wildlife reserve also supports a large reptile population, and large numbers of Eastern Tiger Snakes have been reported by local residents. The Stones State Wildlife Reserve provides one of the best op- portunities to protect and study species such as the Tiger Cat, the Yellow-bellied Glider, and the Eastern Tiger Snake within a small well-defined area of Victoria. Lake Surprise, in the adjacent national park, provides a per- manent source of water for the vertebrate ani- mals inhabiting the block, and the cliffs around the lake also provide habitat for some cliff-nest- ing birds. The Stones Block is important both for fauna conservation and for public recrea- 302 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS tion. However, the recreation consists primarily of sight-seeing and picnicking around Lake Surprise and it is doubtful whether the Manna Gum forests would support intensive recrea- tional use without disturbance to some of the more important vertebrate populations. Around the Stones Block there are a few small isolated blocks of Crown Land which should be man- aged in conjunction with that block. The small blocks contain habitats (e.g. grasslands, mar- shes) which are not well represented in the present wildlife reserve/national park complex in the Hamilton Region. The vegetation in the Mt. Napier Block appears to have been modified to a greater extent than that in the Stones Block and may not support all the species known from the Stones. The woodlands, grasslands and heaths which originally covered most of the Hamilton Re- gion have been cleared mainly for pasture and crops. We are uncertain what the vertebrate species compositions of these areas were be- fore settlement by Europeans, but a number of animal populations have probably declined as a result of habitat modification. A detailed his- torical study of the flora and fauna of this re- gion would be useful in determining future land-use priorities in relation to wildlife popu- lations. At least eight species of animals may be having difficulties in maintaining popula- tion numbers in the Hamilton Region; these in- clude four avian species (Brolga, Plains-wan- derer (Plate 21, Fig. 6), Long-billed Corella and Grey Butcherbird) and four mammal spe- cies (Gunn's Bandicoot, Tuan, Eastern Grey Kangaroo and Red-necked Wallaby). Habitat alteration (e.g. clearing of woodland trees or undergrowth, draining of swamps, grazing and other alterations to the grasslands, clearing of hcathlands) is probably responsible for the de- clining populations of most of these species. De- tailed biological and distributional studies, par- ticularly on the Brolga, Plains-wanderer, Gunn's Bandicoot and Tuan, are required to effectively protect the remaining small popu- lations of these species in western Victoria. The grasslands along the Wannon River and northeast of Hamilton (Fig. 2) were the western limits of an extensive grassland which originally extended eastward almost to Melbourne. In Victoria these grasslands represent the plant formation most drastically reduced as a result of European settlement. Conservation of the remnants of native woodlands which occur in roadside reserves, on margins of the few larger Crown Land blocks and on freehold land should be en- couraged. These small areas should be care- fully managed to ensure that enough large trees remain to support populations of nectar- feeding animals, to produce seeds for natural regeneration and to provide holes for nests and shelter. The narrow strip of heath along the coast was not the subject of a detailed survey. How- ever, the coastal heaths east of the survey area support species such as the Rufous Bristle Bird, Tawny-crowned Honeyeater, Beautiful Firetail, Swamp Antechinus and several species of skinks (Emison et al. 1975) and some of these species are probably present along the coastline of the Hamilton Region. We suggest that a specific effort be made to document the vertebrate fauna in this particular habitat. Many swamps and depressions in the Hamil- ton Region have been drained. A few of the larger water bodies (e.g. Lake Condah, Buck- ley Swamp) may occasionally fill to nearly their original capacity, but usually they are either dry or a series of drains and channels. Brolgas and waterfowl still nest in the region, but their wet- land habitat has been reduced to relatively small areas. Any Crown Lands with potential for supporting wetland habitat in the region should be retained and managed to perpetuate the presence of surface waters and associated vegetation. The Eumeralla River flows through pasture land and has both a mud and a gravel bottom. Its upper reaches have been modified into a steep-sided, deep channel by river improve- ment works. It contains Brown Trout, eels, Tench, Crucian Carp, Tupong and Blackfish. The Shaw River carries a small population of Brown Trout. Lake Yambuk, which is formed by the joining of the Eumeralla and Shaw Rivers, is surrounded by sand dunes and has a sand and mud bottom. Species of fish com- monly caught include bream, mullet, salmon and Estuary Perch. Brown Trout which enter GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 303 the lake from the Eumeralla and Shaw Rivers are sometimes taken (Tunbridge and Rogan 1976). Acknowledgements We are indebted to the following residents in or near the survey area who generously pro- vided fauna data which they had collected over many years of observations: C.N. Austin of Coleraine (now residing at New Gisborne); A. C. Isles of Warrnambool; I. R. McCann of Stawell; A. M. McGarvie of Cavendish (now living on King Island, Tasmania); J. McQueen of Natimuk; W. G. D. Middleton of Wail; J. H. Morris of Hamilton (Fisheries and Wildlife Officer); and O. J. Thomas of Horsham (Fisher- ies and Wildlife Officer). The support of this survey by the following organizations and personnel is gratefully ack- nowledged. National Museum of Victoria: J. M. Dixon for comments on mammals; A. R. McEvey for comments on birds; A. J. Coventry for comments on reptiles; and P. W. Menk- horst for references on the birds. La Trobe University: P. A. Rawlinson for comments on the reptiles. Monash University: H. Parnaby for comments on the bats. Fisheries and Wild- life Division: J. C. F. Wharton (Director), S. J. Cowling (Assistant Director, Wildlife) and J. K. Dempster (Officer-in-charge, Wildlife Re- search) for administrative and financial sup- port; W. M. Bren, D. B. Hespe and J. M. Mar- cius for field and laboratory support; A. Mc- Shane for draughting support; and J. B. Cooper for providing the photographs for all plates. Royal Australasian Ornithologists Union: vari- ous individuals who contributed nest record cards on birds in the survey area; and H. B. Young for her assistance in providing the nest record cards to us. References Anonymous, 1892. Field Naturalist Club's excur- sion to the Grampians, 21st November to 5th December, 1891. Victorian Nat. 8: 181-196. Ashby, E., 1927. The Grampians Range of Victoria and its bird life. Emu 26: 285-292. Attiwill, A. R., 1960. Red-tailed Black Cockatoo in south-east of South Australia. South Aust. Orn. 23: 37-38. Audas, J. W., 1920. Through the Murra Murra coun- try (western Grampians) . Victorian Nat. 37: 59-65. , 1925. One of nature's wonderlands. The Victorian Grampians, Ramsey, Melbourne. Austin, C. N., 1951. Birds of the southern end of Victoria-South Australian border. South Aust. Orn. 20: 14-15. , 1953a. The Black Falcon and some other raptores in south-west Victoria. Emu 53: 77-80. , 1953b. Swift Parrot in south-west Victoria. Emu 53: 260-261. •. 1954. White-throated Warbler in western Victoria. Emu 54: 28. - 1962a. Cattle Egret in south-west Vic- toria. Emu 62: 183. , 1962b. 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Rufous Fantail in western Victoria. Bird Obs. 400: 4. — , 1966. Birds of Portland (Victoria) district. Portland Field Naturalist Club. Maci>onai i), J. D., 1973. Birds of Australia. A. H. and A. W. Reed, Sydney. McCANN, I. R. ( 1957. Regent Honeyeaters. Victorian Nat. 74: 73. — , 1961. Major Mitchell Plateau, in the Vic- torian Grampians. Victorian Nut. 78: 4-7. — , 1963. /// Along the by-ways with the edi- tor. Victorian Nat. 80: 161. — , 1964. In Along the by-ways with the edi- tor. Victorian Nat. 80: 369. McCui loch, F. M., 1973. Birds breeding at Bal- moral, Victoria. Aust. Bird W. 5: 97-99. M( I'M v, A , 1958. Some notes on the Yellow-tufted Honeyeater, South Aust. Om. 22: 59. Mucin 11, A., t973. Christmas camp at Balmoral. Bird Obs. 495. Mitchell, t. l., 1839. Three expeditions into the interior of eastern Australia; with descriptions of the recently explored region of Australia Felix, and of the present colony of New South Wales. Vol. II. 2nd ed. carefully revised. T. and W. Boone, 1 ondon. Parsons, R. I .. N. H. Scarlett and N. J. Rosen- cjrln, 1972. Ecology of some Eucalyptus wood- lands near Halls Cap, Victoria. Victorian Nat 89: 41-49. POOLE, W. E H 1973. A study of breeding in grey kangaroos, Macropus gtganteus Shaw and M. fuliginosus (Desmarest), in Central New South Wales, dust. J. Zool. 21: 183-212. Rwvlinson, P. A., 1971. Reptiles. Victorian Year Book 85: 11-36. , 1975. Two new lizard species from the GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 305 genus Leiolopisma (Scincidae:Lygosominae) in southeastern Australia and Tasmania. Mem. Nat. Mus. Vict. 36: 1-16. Ride, W. D. L., 1970. A guide to the native mammals of Australia. Oxford University Press, Mel- bourne. Rowley, L, 1970. The genus Corvus (Aves:Corvi- dae) in Australia. CSIRO Wildl. Res. 15: 27-71. Schodde, R., 1975. Interim list of Australian song- birds. Passerines. RAOU, Melbourne. Seebeck, J. H., 1976. Mammals in the Pomonal area, The Grampians. Victorian Nat. 93: 138-147. Shanks, D., 1953. Random notes from south-eastern Victoria. Emu 53: 276-279. Sibley, G. T., 1967. A study of the land in the Grampians area. Soil Conservation Authority, Victoria. Slater, P., 1970. A field guide to Australian birds: non-passerines. Rigby, Adelaide. , 1974. A field guide to Australian birds; passerines. Rigby, Adelaide. Smith, M. D., 1962. Long-billed Corella. Aust. Bird W. 1: 201. Specht, R. L., E. M. Roe and V. H. Boughton, 1974. Conservation of major plant communities in Australia and Papua New Guinea. Aust. J. Bot. Supplement no. 7. Storr, G. M., A. H. Lendon and R. W. McKechnie, 1952. Some observations in south-eastern South Australia and adjacent parts of Victoria. South Aust. Or n. 20: 70-71. Sullivan, C, 1928. Notes from the western district, Victoria. South Aust. Orn. 9: 241-249. , 1929a. Arrivals and departures of birds in south-western Victoria and other bird notes. Emu 29: 39-44. , 1929b. The Whiskered Tern. Emu 29: 77. , 1929c. The corella in southern Victoria. Emu 29: 75. 1929d. Birds of the western district, Vic- Walter, H., 1971. Ecology of tropical and sub- tropical vegetation. Oliver and Boyd, Edinburgh. Williams, W. D., 1967. The chemical characteristics of lentic surface waters in Australia: a review. In Australian inland waters and their fauna: eleven studies. Australian National University Press, Canberra. Willis, J. H., 1962. A handbook to plants in Vic- toria. Volume I. Ferns, conifers and monocotyle- dons. Melbourne University Press, Melbourne. , 1972. A handbook to plants in Victoria. Volume II. Dicotyledons. Melbourne University Press, Melbourne. Wheeler, J. and T. Pescott, 1973. Birds of Rock- lands. GeelongNat. 9: 93-96. Wheeler, W. R., 1964. Easter camp at Edenhope, Vic- toria. Bird Obs. 392. , 1965. Bird notes, 1964-65. Bird Obs. 406: 2-4- , 1967a. A handlist of the birds of Victoria. Melbourne University Press, Melbourne. , 1967b. Easter, 1967. Bird Obs. 427: 4-6. toria. South Aust. Orn. 10: 139-141. Tunbridge, B. R. and P. L. Rogan, 1976. A guide to the inland angling waters of Victoria. Fisheries and Wildlife Division, Victoria. Wakefield, N. A., 1963a. Mammal remains from the Grampians, Victoria. Victorian Nat. 80: 130- 133- , 1963b. The Australian pigmy-possums. Victorian Nat. 80: 99-116. -, 1971. The Brush-tailed Rock-wallaby (Pet- rogale penicillata) in western Victoria. Victorian Nat. 88: 92-102. 1974. Mammals of Western Victoria. In The natural history of western Victoria. Aus- tralian Institute of Agricultural Science, Hamil- ton. Wakefield, N. A. and R. M. Warneke, 1967. Some revision in Antechinus (Marsupialia) — 2. Vic- torian Nat. U: 69-99. White, C. M., 1968. Is there a genetic continuity concerned in eyrie maintenance? In Peregrine Falcon populations: their biology and decline. University of Wisconsin Press, Milwaukee. Abbreviations used in the following appendices AR Ararat Region d.b.h. diameter at breast height E east or eastern ER Edenhope Region FWD Fisheries and Wildlife Division, Victoria GR Grampians Region Herb herb layer (Appendix 1) HR Hamilton Region km kilometre m metret mm millimetre N north or northern NMV National Museum of Victoria px. plant community pers. comm. personal communication RAOU Royal Australasian Ornithologists Union S south or southern Shrub shrub layer (Appendix 1) Tree tree layer (Appendix 1) Unpub. MS unpublished manuscript W west or western 306 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS Appendix 1 Annoted list of plant communities in the Grampians-Edcnhope Area of southwestern Victoria. See Appendix 3 for scientific names 1. Heath (White Sallee on sub-alpine sites). Oc- currence. Major Mitchell Plateau and Mt. William (rare), GR. Survey Sites. 12/8/74: S side of Major Mitchell Plateau at 1050 m. 20/3/75: SE side of Mt. William at 1120 m. Floristics. Tree: White Sallee. Shrub: Spike Wattle, Coast Banksia, Silver Banksia, Dwarf She-oak, Shiny Tea-tree and Prickly Tea-tree (also see McCann 1961). Herb: tussocks of sedge and grass (mat-rush, Tasman Flax-lily), several Compositae and other forbs. Moss mats are present. 2. Heath (Grampians Gum on moist sub-alpine sites). Occurrence. Mt. William, GR. Survey Sites. 5/11/74 and 22/3/75: summit of Mt. William at 1120 m. Floristics. Tree: Grampians Gum. Shrub: Spike Wattle, Silver Banksia, Coast Bank- sia, Dwarf She-oak, Prickly Tea-tree, Heath Tea- tree and Shiny Tea-tree. Herb: saw-sedges and many tussocks of short sedges. Moss is present. 3. Heath (Grampians Gum on dry sub-alpine sites). Occurrence. Serra Range, GR. Survey Sites. 21/3/75: W facing slope on Mirranatwa Gap at 460 m. 25/3/75: Serra Range E of Serra Park Homestead. Floristics. Tree: Grampians Gum. Shrub: Silver Banksia, Oyster Bay Pine, Dwarf She-oak, Prickly Tea-tree, Shiny Tea-tree, Common Correa, Common Heath, Pine Heath, Horny Cone-bush, Variable Prickly Grcvillea, Flame Grevillea, flat-pea and Leafless Currant- bush. Herb: mainly sedges. 4. Heath. Occurrence. Victoria, Mt. Difficult and Mt. William Ranges, GR. Survey Sites. 12/8/74: W slope of Major Mitchell Plateau at 1080 m. 19/3/75: Mt. Difficult Range at 800 m. 20/3/75: Carters Track, Mt. Difficult Range at 620 m. 22/4/75: Mt. Thackeray and near Castle Rock, Victoria Range at 850 m. Floristics. Tree: Gram- pians Gum and Brown Stringybark. Shrub: Mountain Hickory Wattle, Spike Wattle, Silver Banksia, Coast Banksia, Veined Beard-heath, Prickly Tea-tree, Grampians Fringe-myrtle, Notched Phebalium, Cross-leaf Honey Myrtle, Wedge-leaf Hop-bush, Rough Coprosma and Hairy Correa. Herb: prostrate sedges, grass tus- socks and some low ferns. Moss is present. 5. Open forest (Messmate and Brown Stringy- bark with a Mountain Hickory Wattle and Veined Beard-heath shrub layer). Occurrence. Uncommon and restricted to high altitudes in the ranges, GR. Survey Sites. 24/4/75: Victoria Range near Mt. Thackeray at 920 m. Floristics. Tree: Messmate and Brown Stringybark. Shrub: Mountain Hickory Wattle, Prickly Tea-tree, Orange Bell-climber, Common Heath, Pine Heath and Veined Beard-heath. Herb: tall tus- socks of saw-sedge with a lower layer of grass tussocks, ferns and other forbs. 6. Open forest (Brown Stringybark, Shining Peppermint and heaths). Occurrence. High alti- tudes in Victoria, Mt. William and Mt. Difficult Ranges, GR. Survey Sites. 14/8/74: Mt. William Range above Stockyard Creek at 600-800 m. 7/9/74: near Wallaby Rocks, head of Sheet of Water Creek, Mt. Difficult Range at 600 m. 9/9/74: head of Hut Creek, Victoria Range at 600 m. Floristics. Tree: Brown Stringybark, Shin- ing Peppermint and Messmate. Shrub: Myrtle Wattle, Mitcheirs Wattle, Spike Wattle, Silver Banksia, Coast Banksia, Oyster Bay Pine, Prickly Tea-tree, Heath Tea-tree, Dwarf Hakea, Bushy Needlcwood, Furze Hakea, Grampians Fringe- myrtle, Grampians Thryptomene, Common Heath and several other low heath species. Herb: sedge tussocks with an underlayer of moss. 7. Tall open forest (Messmate and Narrow-leaf Wattle). Occurrence. Uncommon and restricted to sheltered areas at high altitudes in the ranges, GR. Survey Sites. 9/9/74: near Hut Creek Junc- tion. Victoria Range at 600 m. 19/3/75: Long Gully, Mt. Difficult Road at 600 m. Floristics. Tree : Messmate. Shrub : Blackwood, Narrow- leaf Wattle, Prickly Moses, Prickly Tea-tree, vel- vet-bush, Rough Coprosma and a few low heaths. Herb: tussock and prostrate sedges, Austral Bracken, grass and moss. 8. Open forest (Brown Stringybark and stunted Mountain Grey Gum). Occurrence. Sandstone outcrops at high altitudes on Mt. Difficult and Victoria Ranges, GR. Survey Sites. 5/9/74: near Goat Track, Victoria Range at 640 m. 7/9/74: Wallaby Rocks Track, Mt. Difficult Range. Floris- tics. Tree: Brown Stringybark and Mountain Grey Gum. Shrub: Myrtle Wattle, Spike Wattle, Silver Banksia, Oyster Bay Pine, Prickly Tea-tree, Heath Tea-tree, Grampians Fringe-myrtle, Notched Phe- balium, Dwarf Hakea, Furze Hakea, Variable Prickly Grevillea, Pine Heath, Common Heath and several other low heaths. Herb: prostrate sedges, grass tussocks, Austral Bracken and lichens. GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 307 9. Open forest (Brown Stringybark and Moun- tain Grey Gum). Occurrence, High Altitudes in Victoria and Serra Ranges, GR. Survey Sites. 3 1 / 1 0/ 74 : Sawmill Track, Victoria Range at 600 m. Floristics. Tree: Brown Stringybark and Mountain Grey Gum. Shrub: Blackwood, Prickly Moses, Hedge Wattle, spyridium, Hairy Correa, Orange Bell-climber, Golden-tip, Rough Copros- ma, Rough Bush-pea, and Love Creeper. Herb: Forest Wire Grass, Austral Bracken and a few other ferns. 10. Open forest (Messmate, Brown Stringybark and Mountain Grey Gum). Occurrence. Uncom- mon in the ranges, usually on margins of tall open and open forests, GR. Survey Sites. 14/8/74: Stockyard Creek Track, Mt. William Range at 480 m. 9/9/74: head of Hut Creek, Victoria Range at 400 m. Floristics. Tree: Mess- mate, Brown Stringybark and Mountain Grey Gum. Shrub: Narrow-leaf Wattle, Prickly Moses, Hedge Wattle, Myrtle Wattle, Rough Wattle, Silver Banksia, Dwarf She-oak, Prickly Tea-tree, Yeiiow Hakea, Dwarf Hakea, Common Correa, Common Heath, Austral Grass-tree, spyridium, Hairy Correa, Rough Bush-pea, Rough Coprosma and Orange Bell-climber. Herb: tussocks of sedges, goodenia and lichens. 11. Tall open forest (Messmate and Mountain Grey Gum). Occurrence. Sheltered gullies in the ranges, GR. Survey Sites. 1/11/74: above Silver- band Falls, Serra Range at 450 m. Floristics. Tree: Messmate and Mountain Grey Gum. Shrub: Blackwood, Narrow-leaf Wattle, Hazel Poma- derris, Hairy Correa, Prickly Currant-bush, Rough Coprosma and Victorian Christmas-bush. Herb: saw-sedge tussocks, Austral Bracken, grass, flax-lily, prostrate sedges, moss and several ground ferns. 12. Tall open forest (Mountain Grey Gum). Occurrence. Uncommon and restricted to gullies, GR. Survey Sites. 9/9/74: Happy Track, Victoria Range at 400 m. Floristics. Tree: Mountain Grey Gum. Shrub: Hazel Pomaderris, Blue Ho- wittia, Hairy Correa, Cherry Ballart, spyridium and Rough Coprosma. Herb: saw-sedge tussocks, grass and flax-lily. 13. Open forest (Brown Stringybark). Occur- rence. Uncommon and mainly restricted to shel- tered scree deposits below Victoria Range, GR. Survey Sites. Happy Track, Victoria Range at 400 m Floristics. Tree: Brown Stringybark. Shrub: Myrtle Wattle, Common Correa, velvet- bush, Love Creeper, Rough Bush-pea and Slender Dodder-laurel. Herb: goodenia, flax-lily, tussocks of grass and saw-sedge. 14. Low open forest (Brown Stringybark and Messmate). Occurrence. Common and widespread in the ranges, GR. Survey Sites. 3/11/74: near Camp of Emus Foot, Victoria Range at 400 m. 29/10/74: Muirfoot Track, Black Range at 400 m. 19/3/75: 6 km N of Halls Gap on Mt. Zero Road at 400 m. 20/3/75: shores of Lake Wartook at 400 m. 10/9/74: junction of Wallaby Rocks and Roses Creek Roads at 440 m. Floristics. Tree: Brown Stringybark and Messmate. Shrub: Myrtle Wattle, Mitchell's Wattle, Spike Wattle, Sallow Wattle, Silver Banksia, Oyster Bay Pine, Dwarf She-oak, Winged Spyridium, Prickly Tea-tree, Woolly Tea-tree, Heath Tea-tree, Fringe-myrtle, three species of hakea, Common Heath, Variable Prickly Grevillea, bitter-pea and Cherry Ballart. Herb: sedge tussocks, Austral Bracken, prostrate sedges and flax-lily. 15. Low open forest (Brown Stringybark and Long-leaf Box). Occurrence. Common around low rock and scree slopes in the Grampians and Black Ranges, GR. Survey Sites. 6/9/74: N end of Black Range at 300 m. 11/9/74: below Asses Ears, Mt. Difficult Range at 380 m. Floristics. Tree: Brown Stringybark and Long-leaf Box. Shrub: Myrtle Wattle, Spreading Wattle, Var- nish Wattle, Juniper Wattle, Silver Banksia, Oy- ster Bay Pine, Dwarf She-oak, Winged Spyridium, Heath Tea-tree, grcvilleas. Common Correa, Com- mon Heath, hakeas, Grampians Thryptomene, Cross-leaf Honey-myrtle and several bush-peas. Herb: sedge tussocks, mosses and lichens. 16. Low open forest (Brown Stringybark). Oc- currence. Common on lower slopes of outwash sand in and around the mountains and ranges, GR and ER. Survey Sites. 10/9/74: Hines Track S of Mt. Difficult Range at 240 m. 2/11/74: be- low Asses Ears, Mt. Difficult Range at 300 m. 29/10/74: SW of Mt. Zero at 250 m. 19/3/75: Halls Gap on Mt. Zero Road below Mt. Difficult Range at 260 m. 24/3/75: Mt. Zero at 250 m. 26/3/75: below Red Rock, Victoria Range. 17/4/75: Mt. Arapiles at 250 m. Floristics. Tree: Brown Stringybark. Shrub: 30 species recorded in this association and listed in Appendix 3. Herb: prostrate sedges and annual grasses. 17. Shrubland (Brown Stringybark). Occurrence. Uncommon and restricted to ER where it oc- curs on the crests of a few high sand ridges. Sur- 308 W. B. EMISON, J. W. PORTER, K. C. NORRTS and G. J. APPS vey Sites. 16/4/75 and 19/4/75: Jilpangcr Block. Floristics. Tree: Brown Stringybark. Shrub: Desert Banksia, Mitchell's Wattle, Oyster Bay Pine, Dwarf She-oak, Heath Tea-tree, Daphne Heath, Spreading Brachyloma, Rosy Bacckea, Fringe-myrtle and Beaked Hakca. Herb: mainly prostrate sedges. 18. Shrubland (Brown Stringybark). Occurrence. Common on sand sheets, ER. Survey Sites. 6/10/74: Yallakar Block at 200 m. 6/12/74: Kadnook Block. 16/4/75 and 19/4/75: Jilpanger Block. Floristics. Tree: Brown Stringybark. Shrub: Desert Banksia, Oyster Bay Pine, Dwarf She-oak, Common Heath, bush-pea, Fringe- myrtle, Austral Grass-tree and Spike Wattle. Herb: mainly prostrate sedges. 19. Shrubland (Brown Stringybark). Occur- rence. Common and restricted to sand sheets, ER. Survey Sites. 3/10/74, 4/10/74 and 5/10/74: Tooan Block at 160 m. 4/12/74: Tallageira Block at 160 m 5/12/74: Morca Block at 160 m 11/12/74: Yallakar Block at 200 m. 19/4/75: Jilpanger Block. Floristics. Tree: Brown Stringy- bark. Shrub: same as in 18 (above) plus Myrtle Wattle, Spike Wattle, Silver Banksia and Heath Tea-tree. Herb: prostrate sedges, flax-lily, Aus- tral Bracken, clover and mosses. 20. Shrubland (Brown Stringybark). Occurrence. Common and widespread on sand sheets, ER. Survey Sites. 6/10/74: Yallakar Block at 200 m. 8/12/74: Jilpanger Block. Floristics. Tree: Brown Stringybark with Yellow Gum and Manna Gum on some margins. Shrub: Black Wattle, Prickly Tea-tree, Heath Tea-tree, Fringe-myrtle, Silver Banksia, Desert Banksia, beard-heath, bush-pea, Dwarf She-oak and Oyster Bay Pine. Herb: pro- strate sedges, Austral Bracken and flax-lily. 21. Low open forest (Long-leaf Box). Occur- rence. Uncommon, scattered and restricted to AR and E side of GR except for an isolated oc- currence at Mt. Arapiles, ER. Survey Sites. 11/9/74: near Zumsteins below Mt. Difficult Range, GR, at 240 m. 17/4/75: near lookout on Mt. Arapiles, ER, at 300 m. 15/3/75: near Jallu- kar Block, AR, at 240 m. 15/3/75: One Tree Block, AR, at 400 m. 29/3/75: Black Range S of Stawell, AR, at 400 m. 20/3/75: Pohlners Track near Mt. Zero, GR. 21/4/75: Chinamans Track below Mt. Difficult Range, GR, at 300 m. Floristics. Tree: Long-leaf Box. Shrub: Black Wattle, Hedge Wattle, Wallowa Wattle, Varnish Wattle, Rough Wattle, Sallow Wattle, Juniper Wattle, Golden Wattle, Silver Banksia, Austral Grass-tree, Grampians Thryptomene, Beaked Hakea, Variable-Prickly Grevillea and Cherry Ballart. Herb: a mixture of sedges, forbs and grasses. 22. Open forest (Manna Gum). Occurrence. Re- stricted to stony rises in the Stones and Mt. Na- pier Blocks, HR. Survey Sites. 18/2/75: near the Natural Bridge, Stones Block at 240 m. 18/2/75: near Vaughan Buffer, Stones Block at 240 m. 22/2/75: Millard Track, Stones Block at 240 m. Floristics. Tree: Manna Gum. Shrub: Blackwood, Black Wattle, Cherry Ballart, Sweet Bursaria, Tree Lucerne and cassinia. Herb: Austral Bracken, grass tussocks, thistles, maiden-hair fern, clover and lichens. 23. Open forest (Manna Gum). Occurrence. Mainly on uncleared private lands near Halls Gap, GR. Survey Sites. 1/11/74: S of golf course, Halls Gap at 260 m. Floristics. Tree: Manna Gum. Shrub: Blackwood, Black Wattle and Prickly Tea-tree. Herb: Austral Bracken, grass tussocks, mosses and orchids. 24. Woodland (Manna Gum). Occurrence. Re- stricted to small areas in GR and AR. Survey Sites. 19/3/75: Black Range S of Stawell, AR, at 340 m. 21/4/75: Chinamans Track W of Mt. Difficult Range, GR. Floristics. Tree: Manna Gum (may be a hybrid with Scent-bark). Shrub: Black Wattle, Golden Wattle and Silver Banksia. Herb: a scattered cover of annual grasses. 25. Woodland (Manna Gum). Occurrence. Re- stricted to a narrow zone between Brown Stringy- bark and River Red Gum or Yellow Gum on sands, ER. Survey Sites. 2/10/74: W of Bartons Swamp, Toolondo Block at 160 m. 6/10/74: Haylocks Road, Yallakar Block at 1 80 m. 4/12/74: Tallageira Block at 180 m. 8/12/74: Apsley Common. 5/12/74: Dopewora Block. Floristics. Tree: Manna Gum (possibly a hybrid with Scent-bark). Shrub: Blackwood, Black Wat- tle, Silver Banksia, Prickly Tea-tree, Daphne Heath and Flame Heath. Herb: scattered annual grasses. 26. Heath (Scent-bark). Occurrence. Extensive on outwash sands in GR, particularly in Victoria Valley and Billywing area. Survey Sites. 19/3/75: Mt. Difficult Track at 500 m. 20/3/75: Copper- mine Track near Mt. Zero at 300 m. 22/3/75: Halls Gap rubbish dump near Mokepilly, AR, at 200 m and Cassidy Gap, Scrra Range at 300 m 10/9/74: Lodge and Siphon Roads junction, Vic- GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 309 toria Valley at 220 m. 2/9/74: Lodge Road, Vic- toria Valley. Floristics. Tree: Scent-bark (possibly a hybrid with Manna Gum). Shrub: Varnish Wattle, Myrtle Wattle, Wiriida, Hedge Wattle, Sallow Wattle, Rough Wattle, hakeas, Heath Tea- tree, Prickly Tea-tree, Silver Banksia, Flame Heath and Common Heath. Herb: scattered an- nual grasses and occasionally sedges. 27. Heath (Plate 19, Fig. 2). Occurrence. Com- mon in Victoria Valley and Billywing area, GR. Survey Sites. 3/9/74: Siphon Road, Victoria Val- ley. 4/9/74: Billywing area. Floristics. Tree: none. Shrub: Silver Banksia, Desert Banksia, Dwarf She-oak, Common Heath, beard-heath, Flame Heath, guinea-flower, Furze Hakea, Com- mon Correa and Broom Spurge. Herb: saw-sedge and rapier-sedge. 28. Open forest (Messmate). Occurrence. SW of Bessiebelle HR. Survey Sites. 20/2/75: SW of Bessiebelle at 100 m. Floristics. Tree: Messmate. Shrub: Blackwood, Black Wattle, Prickly Tea-tree and Sweet Bursaria. Herb: Austral Bracken and grass. 29. Heath (Shining Peppermint). Occurrence. Uncommon, mainly in valleys on SE side of GR. Survey Sites. 12/3/75: near head of Wannon River, GR, at 300-380 m. 22/3/75: rubbish dump E of Halls Gap, AR, at 200 m. Floristics. Tree: Shining Peppermint, Brown Stringybark, Mess- mate and Scent-bark. Shrub: Varnish Wattle, Silver Banksia, Oyster Bay Pine, Dwarf She-oak, Common Heath, Prickly Tea-tree, Heath Tea-tree, Variable-Prickly Grcvillea and Common Correa. Herb: mainly sedges (see Parsons et ah 1972). 30. Open forest (Swamp Gum) . Occurrence. Widely distributed along margins of gullies in the S portions of GR and HR. Survey Sites. 9/9/74: Hut Gully, Victoria Range, GR. 11/9/74: near Roses Gap below Mt. Difficult Range, GR. 1/11/74: Halls Gap, GR. 20/2/75: S of Bessie- belle, HR. Floristics. Tree: Swamp Gum; this is an ecotonal community which merges into nearly pure Swamp Gum on the wetter margins and into Manna Gum, Brown Stringybark, Scent-bark, Long-leaf Box, Yellow Box, Messmate, Mountain Grey Gum and Shining Peppermint on the drier margins. Shrub: similar to those listed for com- munity 31 (below) with a few additional species from the drier adjacent communities. Herb: Aus- tral Bracken, grass tussocks, mosses and orchids. 31. Open forest (Swamp Gum). Occurrence. Re- stricted to a few small wet gullies in GR and HR. Survey Sites. 20/2/75: SW of Bessiebelle, HR. 21/3/75: E side of Lake Wartook, GR. 21/3/75: near Cassidy Gap, Serra Range, GR. Floristics. Tree: Swamp Gum. Shrub: Blackwood, Black Wattle, Varnish Wattle, Wiriida, Prickly Moses, Silver Banksia, Dwarf She-oak, Flame Heath, Prickly Tea-tree, Woolly Tea-tree, Heath Tea-tree, Scented Paper-bark and baucra. Herb: Austral Bracken, saw-sedge, grass, moss, goodenia, lich- ens, prostrate sedges and rapier-sedge. 32. Heath (Scented Paper-bark). Occurrence. Uncommon and restricted to a few gullies in GR, becoming common SW of the Stones Block, HR. Survey Sites. 5/9/74: near Goat Track, Victoria Range, GR, and E side of Lake Wartook, Mt. Difficult Range, GR. Floristics. Tree: a few Scent- barks or Messmates may be present. Shrub: Scen- ted Paper-bark, Swamp Honey-myrtle, Prickly Moses, Silver Banksia, Dwarf She-oak and Prickly Tea-tree. Herb: saw-sedge, prostrate sedges, mosses and lichens. 33. Woodland (River Red Gum) (Plate 19, Fig. 1). Occurrence. Widespread in ER, GR and AR. Survey Sites. 9/9/74: Moora Moora Reservoir, Victoria Valley, GR. 12/9/74: Lodge Road, Vic- toria Valley, GR. 1/11/74: Halls Gap rubbish dump. Floristics. Tree: River Red Gum. Shrub: Blackwood, Black Wattle, Prickly Moses, Hedge Wattle, Spike Wattle, Varnish Wattle, Golden Wattle, Prickly Tea-tree, Cross-leaf Honey-myrtle, Cherry Ballart and Silver Banksia. Herb: short sedges, broad-leaved forbs, mosses and grasses. 34. Woodland (River Red Gum and myrtles). Occurrence. Mainly on swamp margins (usually freehold) in ER. Survey Sites. 3/10/74: Kallun- gar Block, ER. 24/3/75: Sheet of Water Creek, Victoria Valley, GR. Floristics. Tree: River Red Gum. Shrub: Prickly Tea-tree, Cross-leaf Honey- myrtle and Scarlet Bottlcbrush. Herb: tall sedges, rushes, water milfoil and spike-rush. 35. Heath (Cross-leaf Honey-myrtle) . Occur- rence. Areas of winter inundation, GR. Survey Sites. Sheet of Water Creek, Victoria Valley, GR. Floristics. Tree: a few River Red Gum. Shrub: Silver Banksia, Prickly Tea-tree, Manuka, Cross- leaf Honey-myrtle, Dwarf Hakea and Yellow Hakea. Herb: grasses, mosses and moisture toler- ant forbs. 36. Woodland (River Red Gum with open grass areas underneath) (Plate 20, Fig. 4). Occur- rence. Originally covered large areas on the 310 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS plains in ER and GR; now mainly cleared. Survey Sites. 2/9/74: near Halls Gap, GR. 8/9/74: near Glenisla Crossing, Victoria Valley, GR. 4/12/74: swamp margin N of Apsley, ER. Floristics. Tree: River Red Gum. Herb: mainly grasses, lilies and other forbs. 37. Woodland and pasture (River Red Gum and annual grasses). Occurrence. River Red Gum woodlands were once widespread and common, now mainly cleared to pastures with scattered trees; common throughout the plains of ER, GR and AR. Survey Sites. 4/9/74: Glcnisla-Billywing area, GR, and Mokepilly, AR. 6/12/74: Thomp- son Creek at Benayeo N of Apsley, ER. 5/12/74: Dopewora Block, ER. Floristics. Tree: River Red Gum. Shrub: a few Bull-oaks. Herb: mainly annual grasses except on gilgaied soils where lilies and sedges predominate. 38. Grassland with River Red Gum fringing the rivers. Occurrence. Natural grassland was not originally extensive in the survey area, occurring only in HR and AR. Grasslands have increased since European settlement. Survey Sites. 19/2/75: along Wannon River and in Wannon Valley near Coleraine, HR. Floristics. Tree: River Red Gum along the rivers. Herb: mainly annual grasses; along rivers Cane Grass and tall forbs are pre- sent. 39. Open forest (Candlebark). Occurrence. Rare, seen only in GR. Survey Sites. W of Lake Bellfield at 300 m. Floristics. Tree: Candlebark. Shrub: similar to the list for community 41 (be- low). 40. Open forest (Red Stringybark) . Occurrence. Uncommon in E portion of AR. Survey Sites. 15/3/75: One Tree Hill Block at 420 m. Floris- tics. Tree: Red Stringybark. Shrub: similar to the list for community 41 (below). Herb: tus- socks of grass. 41. Woodland (Yellow Box). Occurrence. Un- common and restricted to GR and AR. Survey Sites. 9/3/75: SW of Stawell, AR, at 400 m. 21/4/75: Chinamans Track, GR, at 320 m. 15/3/75: Jallukar Block, AR, at 240 m and W of Ararat, AR, at 400 m. Floristics. Tree: Yellow Box, Long-leaf Box and Red Stringybark. Shrub: Blackwood, Black Wattle, Narrow-leaf Wattle, Prickly Moses, Hedge Wattle, Sallow Wattle, Lightwood, Silver Banksia, Oyster Bay Pine, Drooping She-oak, Common Heath, Flame Heath, Cranberry Heath, Prickly Tea-tree and Grampians Thryptomene. Herb: grasses, forbs and mosses. 42. Woodland (Yellow Gum and Yellow Box). Occurrence. Widespread in ER, GR and AR where it fringes Crown Land; much has been cleared. Survey Sites. 6/9/74: 10 km E of Halls Gap in a Crown Land road reserve, AR, at 200 m; H.G.H. Corner, Black Range, GR, at 240 m; and Roses Gap, GR. 11/9/74: Mt. Victory and Asses Ears Roads junction, GR, at 240 m. 3/10/73: Toolondo Block, ER, at 200 m. 1/12/74: Mokepilly Block, AR. 15/3/75: Jallu- kar Block, AR. Floristics. Tree: Yellow Gum and Yellow Box. Shrub: wattles and a few heath-like species. Herb: short sedges or annual grasses, de- pending on soil moisture. 43. Woodland (Yellow Gum and wattles). Oc- currence. Outwash sands at Mt. Arapiles, ER, probably also present in GR. Survey Sites. 16/4/75: NW side of Mt. Arapiles, ER, at 240 m. Floristics. Tree: Yellow Gum and Yellow Box. Shrub: Golden Wattle, Juniper Wattle, Gold-dust Wattle, Wallowa, Pine Heath, Cranberry Heath, guinea-flower and Heath-myrtle. Herb: annual grasses, short sedges and other forbs. 44. Woodland (Yellow Gum and heaths). Occur- rence. Widespread in ER, occurring in small stands on winter-wet clay pans in and around sand sheets. Survey Sites. 5/12/74: Morea Block at 160 m. 7/12/74: Tallageira Block. 22/3/75: Yallakar Block. Floristics. Tree: Yellow Gum and Brown Stringybark. Shrub: Silver Banksia, Flame Heath, Cranberry Heath, guinea-flower, Prickly Tea-tree, Broom Baeckea, Fringe-myrtle, Mallee Honey-myrtle, Violet Honey-myrtle, Moonah, Scarlet Bottlebrush, Desert Hakea, Muntries, Yel- low Rice-flower, Cherry Baliart and several spe- cies of wattles. Herb: not assessed. 45. Woodland (Yellow Gum and Bull-oak). Oc- currence. Uncommon in ER and AR, mainly cleared and now only occurring as remnants on freehold land. Survey Sites. 7/12/74: Thompson Creek along Benayeo Road, ER. 22/3/75: near Mokepilly, AR. Floristics. Tree: Yellow Gum. Shrub: Bull-oak. Herb: annual grasses and a few sedges, 46. Woodland (Grey Box). Occurrence. Orig- inally widespread in the plains of ER and AR, now mainly cleared; a few small stands still fringe some Crown Lands. Survey Sites. 30/10/74: road- side near Lah-arum, AR. 7/12/74: Thompson GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 311 Creek at Benayeo N of Apsley, ER. 17/4/75: golf course at Mt. Arapiles, ER. 18/4/75: 2 km S of Horsham, ER. 15/4/75: near Wimmera River E of Horsham, AR. Floristics. Tree: Grey Box, Black Box and Bull-oak. Shrub: Bull-oak and a few species of wattles. Herb: see Connor (1966) for a list of both shrub and herb species. 47. Woodland (Black Box and Bull-oak). Occur- rence, Uncommon and restricted to flood plains of the Wimmera River, ER and AR. Survey Sites. 8/8/74: NW of Mt. Arapiles, ER. 15/4/75: Wimmera River E of Horsham, AR. Floristics. Tree: Black Box and Bull-oak. Shrub: Bull-oak. Herb: see Connor (1966) for list. 48. Shrubland (mallee eucalypts). Occurrence. A few small areas of mallee occur between Hor- sham and the Tooan Block, ER. They are outliers of mallee from the Little Desert and it is sugges- ted that 2 or 3 tongues of mallee once extended S across what is now farmland along ironstone ridges into the Natimuk area (Middleton pers. comm.). Survey Sites. 1/10/74: Tooan Block. 6/10/74: SW of Horsham and E of Noradjuha. 16/4/75: NW slopes of Mt. Arapiles. 17/4/75: flora reserve E of Noradjuha. Floristics. Tree: Kamarooka Mallee, Dumosa Mallee, Yellow Mal- lee and Peppermint Box. Shrub: Golden Wattle, Gold-dust Wattle, Spiny Wattle, Juniper Wattle, Grey Mulga, Oyster Bay Pine, Flame Heath, Cranberry Heath, hakeas, guinea-flower, Rosy Baeckea, Muntries, Fringe-myrtle, Leafless Bitter- pea, Thorny Bitter-pea, correa, Sweet Bursaria, Cross-leaf Honey-myrtle, Violet Honey-myrtle and Broom Honey-myrtle. Herb: annual grasses, mosses and a few sedges. 49. Woodland (Slender Cypress Pine). Occur- rence. Mainly restricted to old lunettes in ER. Survey Sites. 5/10/74: S edge of Crown Land in Kallungar Block. 18/4/75: 4 km S of Horsham. Floristics. Tree : Slender Cypress Pine. Shrub : Flame Heath and Daphne Heath. Herb: annual grasses. 50. Shrubland (Salt Paper-bark). Occurrence. Restricted to the shores of a chain of saline lakes occurring along the E boundaries of the Tooan, Jilpanger and Toolondo Blocks, ER. Survey Sites. 7/10/74: lakeside N of Douglas in Jilpanger Block. Floristics. Tree: Salt Paper- bark. Herb: mainly sedges. W 312 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS a a 9 a o 6 y a CT3 11 o 3 w o 5 1 « I p 1 1 ,1 1 p ■ — v >, ..-> * ■a p »o oo ° i-s o rn ~ © 4 °~ rj I I un 6 2-S g II # ^6 en O Tt O ^ O m -L m i— I »/~> ^t" IT) Tf >Ti rt OO ^H o ^ en (N eN © © n ^O ir» " en »— < 6 *" <^ o o A t- (N ■^ o o _ »n rl -t O O >o *^ 00 ^o Tt in n-( ° o S u §i p p ii ii s .2 '•£3 i u O c O o c I 8. o H S o SC e D C E a o < cj rl --h o O I i—i h m oo ,' 00 °° "O O J, ^ ^ ^ i »o io o 4 ^ 31 ^ »C >n O « n so oo il^OOO o ^ I 00 ^h vo m e^ ^ O O i Z: ^ M >n O in o o ^h en rs o 6 m m O ** I O ^j as oo io »n I O ^ 00 o m o *o I in I n © On rn »n © *n © ©°° ■^ © © © t-- ■«t © © <^© ^ »o © ^ © to "* © -t© f^ un «o © ^ © © m o ©^ oo o ^ ■e I §: K oq GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 313 00 00 o N in in vi r4 in n i— i § 1 so T— 1 m s 1 SO >t o s ^ 5 -a * u ^ S3, «> I o o X »n rt o Is 1 BO B £ o _5i aj ■ 4> o o a fe a j-i s »-* ^ .2 cd s a xl o o ON -L -L *—• © o ' 00 t^ <0 N 2 111" o 4-^ ^ Hio'OhO rl 6 <* o ^ 6 ' o o r* © ^6 (N ^6* m o ■* I -H moo -1 GO r-. (N O "OO'-lOO TfiriHO 3 . ^-s^ o h4 [ o ffi pq »n [ * O c3 f-j > 4H ■t -1 O q_, <^-( o o c 8-2 o o « 3 H) & Pi & ^ 314 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS Appendix 3. — Plant species list Common name Scientific name Plant community numbers (see Appendix 1 ) Annual Veldt Grass Ehrharta longiflora Austral Bracken Pteridhim esculentum Austral Grass-tree Xanthorrhoea australis 4, 6,7, 10, 14-19,21,26 Australian Clematis Clematis aristata 1,9, 11, 13 Australian Dusty Miller Spyridium pcirvifolium Australian Indigo Indigofera australis Beaked Hakea Hakea rostrata 14-18, 21 Bitter-pea Davlesia mimosoides Black-anther Flax-lily Dianella revoluta Black Box Eucalyptus largiflorens 47 Black Rapier-sedge Lepidosperma carphoides Black Wattle Acacia mearusii 14, 16, 20-26, 28, 30, 31, 33, 39-41 Blackwood Acacia melanoxylon 7, 9, 11, 22, 23, 25, 28, 30, 33, 39^1 Blue Boronia Boronia caerulescens Blue Howittia Howittia trilocularis 12 Blue Stars Chamaescilla corymbosa Blue Tinsel-lily Calectasia cyanea Boneseed Chrysanthemoides monilijera Broom Baeckea Baeckea behrii 21,44,48 Broom Honey-myrtle Melaleuca uncinata 48 Broom Spurge Amperea xiphoclada 1,21, 27, 29 Brown Stringybark Eucalyptus baxteri 4-5,7-10, 13-20 Bull-oak Casuarina luehmannii 16,37,38,42,45,46 Bundled Guinea-flower Hibbertia fasciculata 14-16, 18-20, 25, 26, 27, 29, 33, 43 Bush Heath Brachyloma ericoides 15, 18 Bushy Needlewood Hakea sericea Bush-pea Pultenaea benthamii 3, 11 Bush-pea Pultenaea costata 5 Bush-pea Pultenaea d'altonii 3 Bush-pea Pultenaea hibbertioides 17, 18,26 Bush-pea Pultenaea humilis 16 Bush-pea Pultenaea laxiflora 16,43 Bush-pea Pultenaea prolifera 16, 27 Bush-pea Pultenaea prostrata Button-grass Gymnoschoenus sphaeroce- phalus Candlebark Eucalyptus rubida 39 Cassinia Cassinia sp. 4 Cherry Ballart Exocarpos cupressiformis 12, 14, 16,21,22,26,33,44 Chocolate Lily Dichopogon strictus Cleland's Beard-heath Leucopogon clelandii Clustered Everlasting Helichrysum semipapposum Coarse Dodder-laurel Cassytha melantha Coarse Twine-rush Leptocarpus brownii Coast Banksia Banksia integrifolia 1,2,6 Coast Saw-sedge Gahnia trifida • Coast Swainson-pea Swainsona lessertiifolia Common Beard-heath Leucopogon virgatus 15-20,25,27,29 Common Billy-buttons Craspedia glauca Common Correa Correa reflexa 1-3, 6-8, 10, 13-16, 18, 19, 20, 26, 27, 29 GRAMPIAN S-EDENHOPE VERTEBRATE FAUNA SURVEY 315 Common name Common Heath Common Hovea Common Rapier-sedge Cootamundra Wattle Cranberry Heath Cross-leaf Honey-myrtle Dagger Hakea Daisy-bush Daphne Heath Desert Banksia Downy Dodder-laurel Desert Hakea Drooping She-oak Dumosa Mallee Dwarf Geebung Dwarf Hakea Dwarf She-oak Early Black Wattle Erect Guinea-flower Eutaxia Fairy Wax-flower Flame Grevillea Flame Heath Flat Cord-rush Forest Wire Grass Fringed Everlasting Fringe-myrtle Furze Hakea Gold-dust Wattle Golden Heath Golden Pennants Golden Spray Golden-tip Golden Wattle Gorse Bitter-pea Grampians Fringe-myrtle Grampians Grevillea Grampians Guinea-flower Grampians Gum Grampians Parrot-pea Grampians Thryptomene Grey Box Grey Everlasting Grey Mulga Guinea-flower Guinea-flower Guinea-flower Guinea-flower Hairy Boronia Hairy Correa Hairy Geebung Scientific name Plant community numbers (see Appendix 1 ) Epacris impressa Hovea heterophylla Lepidosperma filiforme Acacia bailey ana Astroloma humifusum Melaleuca decussata Hakea teretijolia Olearia erubescens Brachyloma daphnoides Banksia ornata Cassytha pubescens Hakea muellerana Casuarina stricta Eucalyptus dumosa Persoonia chamaepeuce Hakea rugosa Casuarina pusilla Acacia decurrens Hibbertia stricta Eutaxia microphylla Eriostemon verrucosus Grevillea dimorpha Astroloma conostephioides Restio complanatus Tetrarrhena jtincea Helichrysum baxteri Calytrix tetragona Hakea ulicina Acacia acinacca Styphelia adscendens Loudonia behrii Viminaria juncea Goodia lotijolia Acacia pycnantha Daviesia ulici folia Calytrix sullivanii Grevillea confertifolia Hibbertia humifusa Eucalyptus alpina Diliwynia oreodoxa Thryptomene calycina Eucalyptus microcarpa Helichrysum obcordatum Acacia brachybotrya Hibbertia cistiflora Hibbertia obtusifolia Hibbertia procumbens Hibbertia virgata Boronia pilosa Correa aemula Persoonia rigida 1-3, 5, 6, 8, 10, 13-16, 18-20, 26, 27, 29, 39-41 8, 11, 12, 14-16,27 20 16,20-21,25,26, 33,39-41,43,44,48 4, 15,33-35,48 14, 16- -21, 26 16- -20, 27 44 21, 39- -41 48 3, 6, 8, 10, 15,20, 35,42 1-3, 10, 14-20, 26-27, 29, 31, 32 44 3 6, 14-21,25-27,31,39-41,43,44,48 9 4, 14, 16-21,44,48 6, 8, 14, 18, 20, 27 42, 43, 46, 48 6,8, 17, 18,25,26,27,31 16,21,33,39-41,44,48 13, 14, 16, 17,20 1,4,6 2-4 6, 15, 16,21,26, 39-41 46 44,48 17 48 16, 18-20,25-27,44 4,9-12, 14 14, 15, 17-20 316 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS Common name Scientific name Plant community numbers (see Appendix 1) Hairy Mint-bush Prostanthera hirtula Hare's-tail Lagurus ovatus Harlequin-flower Sparaxis grandiflora Hazel Pomaderris Pomaderris aspera 11, 12 Heath-myrtle Micromyrtus ciliata 16,26,43,48 Heath Spyridium Spyridium eriocephalum 11 Heath Tea-tree Leptospermum myrsinoides 1,2,6, 14-20,26,29, 31, 32 Hedge Wattle Acacia armata 9, 10,21,26,33,39-41,44 Holly Grevillea Greviliea ilicifolia Hooked Needlewood Hakea vittata Hop Bitter-pea Daviesia latijolia Hop Wattle A cacia stricta Horny Cone-bush Isopogon ceratophyllus 3, 6, 8, 10, 14-20, 26, 27, 29, 31 Ivy Flat-pea Platylobium triangulare 3 Juniper Wattle Acacia ulicifolia 15, 16,21,43,48 Kamarooka Mallee Eucalyptus froggattii 48 Lance Beard-heath Leucopogon lanceolatus Large-leaf Ray-flower Anthocercis frondosa Lavender Grevillea Grevillea lavandulacea 15 Leafless Bitter-pea Daviesia brevijolia 8, 14-18,26,48 Leafless Currant-bush Leptomeria aphylla 3, 8, 14, 15, 17, 18 Lightwood Acacia implexa Long-leaf Box Eucalyptus goniocalyx 15,21 Love Creeper Comesperma volubile 8, 10, 13, 16 Mallee Honey-myrtle Melaleuca neglecta Manna Gum Eucalyptus viminalis 22-26 Manuka Leptospermum scoparium 35 Matted Bush-pea Pultenaea pedunculata 15, 16 Messmate Stringybark Eucalyptus obliqua 5,7, 10,11, 14,28 Mistletoes Amyema sp. Mitchell's Wattle Acacia mitchellii 6, 14, 17, 18 Moonah Melaleuca lanceolata 4,44 Mountain Correa Correa lawrenciana Mountain Grevillea Grevillea alpina Mountain Grey Gum Eucalyptus cypellocarpa 8-12 Mountain Hickory Wattle Acacia obliquinervia 4,5 Mountain Pepper Drimys lanceolata 5,6,8, 10, 13-16, 18-20 Mountain Tea-tree Leptospermum grandifolium Mt. Byron Bush-pea Pultenaea patellifolia 15, 16 Muntries Kunzea pomifera 48 Myrtle Wattle Acacia myrtifolia 6,8, 10, 13-16, 19,28 Narrow-leaf Bitter-pea Daviesia virgata 15,20 Narrow-leaf Trymalium Trymalium d'altonii Narrow-leaf Wattle Acacia mucronata 7, 10,11,39-41 Notched Phebalium Phebalium bilobum 1,4,5 Onion-grass Romulea rosea Orange Bell-climber Marianthus bignoniaceus 5, 9, 10 Oyster Bay Pine CalUtris rhomboidea 2, 4, 6, 8, 14-20, 29, 39-41, 48 Pale-fruit Ballart Exocarpos strictits 44 Pale Turpentine Bush Beyeria leschenaultii 16,21 Pale Rush Juncus pallidus Peach Heath Lissanthe strigosa GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 317 Common name Scientific name Plant community numbers (see Appendix 1 ) Peppermint Box Pine Heath Pink Beard-heath Pink-bells Pink Swamp-heath Ploughshare Wattle Prickly Bush-pea Prickly Cryptandra Prickly Currant-bush Prickly Geebung Prickly Moses Prickly Starwort Prickly Tea-tree Purple Coral-pea Purple Eyebright Red-fruit Saw-sedge Red Parrot-pea Red Stringybark River Bottlebrush River Red Gum River Tea-tree Rock Correa Rosemary Grevillea Rosy Baeckea Rough Bush-pea Rough Coprosma Rough Mint-bush Rough Star-hair Rough Wattle Round-leaf Mint-bush Round Templetonia Ruddy Beard-heath Rust Bush-pea Rusty-pods Sallow Wattle Salt Paper-bark Saw-sedge Scarlet Bottlebrush Scarlet Coral-pea Scent-bark Scented Paper-bark Scrambling Lily Scrubby Velvet-bush Shrub Violet Shining Peppermint Shiny Tea-tree Short Purple-flag Showy Bauera Showy Parrot-pea Silky Guinea-flower Eucalyptus odorata Astroloma pinijolium Leucopogon ericoides Tetratheca ciliata Sprengelia incarnata Acacia gunnii Pultenaea juniperina Cryptandra tomentosa Coprosma quadrifida Persoonia juniperina Acacia verticillata Stellaria pungens Leptospermum juniperinum Hardenbergia violacea Euphrasia collina Gahnia sieberiana Dillwynia hispida Eucalyptus macrorhyncha Callistemon paludosus Eucalyptus camaldulensis Leptospermum obovatum Correa glabra Grevillea rosmarini folia Baeckea ramosissima Pultenaea scabra Coprosma hirtella Prostanthera denticulata Astrotricha asperifolia Acacia aspera Prostanthera rotundifolia Templetonia egena Leucopogon rufus Pultenaea hispidula Hovea longifolia Acacia longifolia Melaleuca halmaturorum Gahnia sp. Callistemon macropunctatus Kennedia pro strata Eucalyptus aromaphloia Melaleuca squarrosa Geitonoplesium cymosum Lasiopetalum dasyphyllum Hybanthus floribundus Eucalyptus nitida Leptospermum nitidum Pater sonia fragilis Bauera sessiliflora Dillwynia sericea Hibbertia sericea 48 1,4,5,7,8, 15, 16, 19,21,26 10, 16 5,8, 10, 13-15, 18-20,26 1 27 11 8 7, 9, 10, 14, 26, 28, 29, 31, 32, 33, 39-41 1-8, 10, 14, 16, 20, 23, 25-26, 28-35, 44 40 33-38 30 1,2 17, 18,48 8,9, 10, 13, 14 4, 7, 9-12 10, 21, 26 27,32 1, 6, 17, 20 14, 16,21,24,26,49-52 50 34,44 15, 16, 18-20 24-26, 29 31, 32 6,7 3, 6, 14, 29 1-4 11, 31 14 15-21,25,48 318 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS Common name Scientific name Silver Banksia Silver Daisy Stiver Wattle Slender Bog-rush Slender Cypress Pine Slender Dodder-laurel Slender Honey-myrtle Slender Hop-bush Slender Mint-bush Slender Rice-flower Slender Saw-sedge Slender Smoke-bush Slender Twine-rush Slender Velvet-bush Small-flower Grevillea Small-fruit Hakea Small Grass-tree Small-leaved Clematis Smooth Tea-tree Snow Myrtle Spike Wattle Spiny Mint-bush Spiny Wattle Spreading Brachyloma Spreading Wattle Sugar Gum Swamp Club-moss Swamp Gum Swamp Honey-myrtle Sweet Apple-berry Sweet Bursaria Sweet Quandong Tall Rice-flower Tall Sword-sedge Tasman Flax-lily Tassel Cord-rush Tassel Rope-bush Thatch Saw-sedge Thorny Bitter-pea Thyme Beard-heath Trailing Ground-berry Tree Everlasting Tree Lucerne Variable Daisy Variable Prickly Grevillea Variable Sword-sedge Varnish Wattle Veined Beard-heath Victorian Christmas-bush Victorian Flat-pea Victorian Smoke-bush Plant community numbers (see Appendix 1) Banksia marginata Celmisia asteliifolia Acacia dealbata Schoenus tenuissimus Callitris preissii Cassytha glabella Melaleuca gibbosa Dodonaea angustissima Prostanthera saxicola Pimelea I in i folia Gahnia microstachya Conospermum patens Leptocarpus tenax Lasiopetalum baueri Grevillea parviflora Hakea rnicrocarpa Xanthorrhoea minor Clematis microphylla Leptospermum glabrescens Calytrix alpestris Acacia oxycedrus Prostanthera spinosa Acacia spinescens Brachyloma depressum Acacia diffusa Eucalyptus cladocalyx Selageinella uliginosa Eucalyptus ovata Melaleuca squamea Billardierai cymosa Bursaria spinosa Santalum acuminatum Pimelea ligustrina Lepidosperma elatius Dianella tasmanica Restio tetraphyllus Hypolaena fastigiata Gahnia radula Daviesia pectinata Leucopogon thymifolius Acrotiche prostrata Helichrysum dendroideum Chamaecytisus proliferus Brachycome ciliaris Grevillea aquifolium Lepidosperma laterale Acacia verniciflua Leucopogon neurophyllus Prostanthera lasianthos Platylobium alternifolium Conospermum mitchellii 1, 4, 6, 8, 10, 14-16, 18-21, 24-27, 29-33, 35, 39-41,44 1 49 4, 6, 13-20,26 21 14, 16-19, 26 6, 10, 13,44,48 20,25 16, 19,22 1,2,4,6,8, 13, 14, 19,26,31 14, 48 14^21, 33 15 37 30, 31 44 11, 14, 15,22,28,48 48 2,7,9,14,16 22,25 1-3,6, 8, 14-16,21,26,29 15, 16,21,26,33 1,2,45 11, 31 1,2, 8, 16,27,29 GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 319 Common name Scientific name Plant community numbers (see Appendix 1) Violet Honey-myrtle Violet Kunzea Wallowa Wedge-leaf Hop-bush Western Golden-tip White Sallee Winged Spyridium Wirilda Wiry Bauer a Wombat Berry Woolly-style Heath Woolly Tea-tree Yellow Box Yellow Gum Yellow Hakea Yellow Mallee Yellow Rice-flower Zieria Melaleuca wilsonii Kunzea parvifolia Acacia calamifolia Dodonaea cuneata Goodia medicaginea Eucalyptus pauciflora Spyridium vexilliferum Acacia retinodes Bauera rubioides Eustrephus latifolius Epacris lanuginosa Leptospermum lanigerum Eucalyptus melliodora Eucalyptus leucoxylon Hakea nodosa Eucalyptus incrassata Pimelea flava Zieria 44,48 44 16, 21, 30, 43, 44 4, 16 1,2 14, 15 14, 16, 26, 31, 33 1, 6, 30 14,31,33, 39- -41 41,42 42-45 10, 35 48 44 320 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS Appendix 4. Relationships between the species of Eucalyptus occurring in the Grampians- Edenhope Area Hamilton Region Hamilton Region Ararat Region Grampians Region Edenhope Region Edenhope Region GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 321 Appendix 5. Annotated list of birds from the Grampians-Edenhope Area of southwestern Victoria DROMAIIDAE (EMU) Dromaius novaehollandiae. Emu. Abundance and distribution. Common and re- stricted in ER, GR and AR; not seen in HR. Flocks containing 50 birds were seen occasionally; However, most sightings were of family groups! Habitat. Mainly pasture lands bordering wood- land and open forest, particularly in GR. Occa- sional flocks were also in some Crown Land areas covered by woodland, shrubland, open forest and heath. It appears that the mosaic pattern of wood- land or open forest surrounded by pasture land which occurs in the N portion of the survey area is conducive to the continued success of this spe- cies in the area. Breeding. Between October and December 1974 the total number of young counted in six family groups ranged from 3 to 8 and averaged 4-5 young. On 28 July 1957 a nest with eight eggs was found near Mt. Zero, GR, in an area of Brown Stringybark with heath under- growth (Binns 1957). PODICIPEDIDAE (GREBES) Podiceps cristatus. Great Crested Grebe. Abundance and distribution. Rare. Recorded in ER near the Mt. Arapiles Block, on Jaka Lake and on Lake Wallace; in GR near Dunkeld; and in AR at Lake Fyans. Habitat. Permanent fresh- water lakes and reservoirs with areas of open deep water. Breeding. Eggs were found at Lake Wallace, ER, in late December 1972 (Mitchell 1973;McCulloch 1973). Podiceps poliocephalus. Hoary-headed Grebe. Abundance and distribution. Uncommon and widespread; usually alone or in small flocks. Habitat. Temporary and permanent freshwater lakes, swamps and flooded pastures; open water areas are favoured. Breeding. A nest containing eggs was in a flooded pasture in ER near the Mt. Arapiles Block in October 1974. Tachybaptus novaehollandiae. Little Grebe. Abundance and distribution. Common and wide- spread in ER and AR; uncommon in GR and HR. Habitat. Temporary and permanent fresh- water swamps and lakes. Breeding. A floating nest containing seven eggs was found (3 October 1974) on a pond 7 x 15 m and located between Kaliungar and Tooan Blocks in ER. Sedges and River Red Gums were on its margin, but the sur- rounding area was mainly pasture. PELECANIDAE (PELICANS) Pelecanus conspicillatus. Australian Pelican. Abundance and distribution. Uncommon. Re- corded in ER at Toolondo Reservoir; in GR at Rocklands Reservoir, Marney Swamp and Moora Moora Reservoir; and in AR at Lake Lonsdale. Habitat. Large bodies of deep standing fresh water. ANHINGIDAE (DARTERS) Anhinga melanogaster. Darter. Abundance and distribution. Rare. Recorded in GR on Rocklands Reservoir and at Cherrypool. Habitat. Large freshwater lakes and reservoirs. Breeding. A single nest on a horizontal tree branch was found near Rocklands Reservoir in December 1974 (Morris pers. comm.). PHALACROCORACTIDAE (CORMORANTS) Phalacrocorax varius. Pied Cormorant. Abundance and distribution. Rare. Recorded in AR at Lake Lonsdale. Habitat. Large permanent freshwater lakes. Breeding. One pair was nesting at Lake Lonsdale in December 1964 (Wheeler 1965). Phalacrocorax melanoleucos. Little Pied Cormorant. Abundance and distribution. Common and wide- spread. Habitat. It was present on large per- manent bodies of standing fresh water (e.g. Too- londo and Rocklands Reservoirs, and Lakes Fy- ans, Lonsdale and Charlegrark), and farm dams in all regions, on temporary freshwater swamps, on flooded pastures and woodlands particularly in ER, on streams and rivers (e.g. Boggy Creek in GR and the Glenelg River) and on estuaries (Lake Yambuk at the mouth of the Eumeralla River). Breeding. On 7 December 1974 a nesting colony consisting of at least eight nests was lo- cated in the middle of a flooded woodland on the E margin of the Tallageira Block, ER. The nests were on the branches of River Red Gums and contained well-developed young. Thirty pairs nested in AR on Lake Lonsdale between Decem- ber 1964 and February 1965 (Wheeler 1965). Phalacrocorax carbo. Black Cormorant. Abundance and distribution. Uncommon. Re- 322 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS corded in GR at Bryan Swamp, Rocklands Reser- voir and near Balmoral; in AR at Lake Lonsdale and near Mokepilly; and in HR at Cavendish. Habitat. Large deep bodies of permanent fresh water; occasionally rivers, temporary swamps and other shallow bodies of water. Breeding. Twenty pairs nested in AR at Lake Lonsdale between December 1964 and February 1965 (Wheeler 1965). Phalacrocorax sulcirostris. Little Black Cormorant. Abundance and distribution. Uncommon. Re- corded in ER at Lake Connangorach, over flooded woodland near Apsley and at Lake Wal- lace; in GR at Rocklands Reservoir and near Bal- moral; and in AR at Lake Lonsdale. Habitat. Permanent bodies of flowing and standing fresh water. Breeding. Sixty pairs nested in AR at Lake Lonsdale between December 1964 and February 1965 (Wheeler J965). ARDEIDAE (HERONS AND BITTERNS) Ardea pacifica. White-necked Heron. Abundance and distribution. Uncommon and widespread with local concentrations, particu- larly in ER and GR. Habitat. Pastures (either flooded or with new plant growth) or shallow margins of swamps and lakes. Breeding. Several nests were in River Red Gums around Barton Swamp, ER, in October 1974. Two nests, each containing at least two well-developed young, were in the Tallageira Block, ER, on 7 December 1 974. Both were in the lower branches of River Red Gums standing in the middle of flooded woodlands. Ardea novaehollandiae. White-faced Heron. Abundance and distribution. Common and wide- spread. Habitat. Margins of temporary and per- manent freshwater swamps and lakes; pastures (both flooded and those with new plant growth) are favoured foraging areas. Breeding. Nesting recorded in GR (McGarvie pers. comm.). Ardeola ibis. Cattle Egret. Abundance and distribution. Rare; recently estab- lished in Australia and recorded in GR near Dunkeld in 1962 (Austin 1962a). Recorded in HR near Nareen in 1972 and 1973 (Austin pers. comm.). Habitat. Swamp margins and pastures. Egretta alba. Large Egret. Abundance and distribution. Uncommon and widespread with local concentrations during FWD survey in ER at Lake Charlegrark and S of Mt. Arapilcs Block; and in GR at Bryan Swamp and in the Wannon River floodplain at SE end of Serra Range. Habitat. Shallow margins of fresh- water lakes and swamps and pastures (either flooded or with new plant growth). Nycticorax caledonicus. Nankeen Night-heron. Abundance and distribution. Uncommon; re- corded in ER at Lake Charlegrark and St. Marys Lake; in GR at Bryan Swamp, Cherrypool and near Balmoral; and in HR near Cavendish. Habitat. Usually in trees on margins of freshwater streams, lakes and swamps. Botaurus poiciloptilus. Brown Bittern. Abundance and distribution. Rare; not seen dur- ing FWD survey. Recorded N of Mt. Dundas Block in GR in 1956-57 (McGarvie pers. comm.); and in HR at Mt. Eccles in September 1951 (Learmonth 1951). Habitat. Confined to thick reed-beds in swamps, lakes and streams (Wheeler 1967a). PLATALEIDAE (IBISES AND SPOONBILLS) Plegadis falcinellus. Glossy Ibis. Abundance and distribution. Rare; not seen dur- ing FWD survey. Recorded in late December 1972, at Lake Natimuk (just N of the ER bound- ary) and in GR at Bryan Swamp and near Bal- moral. Habitat. Margins of rivers and swamps. Breeding. No nesting recorded in the survey area. However, it breeds at Naracoorte, South Australia (Condon 1969), which is about 20 km W of the western boundary of ER. Movements. Condon (1969) indicates that its occurrence depends on rainfall. It is interesting to note that 1972 (when flocks were in the survey area) was an unusually dry year (McCulloch 1973). Threskiornis molucca. White Ibis. Abundance and distribution. Uncommon and widespread. Habitat. Margins of freshwater swamps, lakes and rivers (e.g. Bryan Swamp in GR, Lake Lonsdale in AR, the Wannon River in GR, etc.). It forages in pastures. Breeding. Nest- ing occurred in GR at Bryan Swamp in 1960 and along the Wannon River near Bryan Swamp in 1968; and in HR at Buckley Swamp in 1968 (Cowling unpub. MS). Conservation aspects. The conservation of this species should be encouraged because it feeds upon invertebrate pest species in agricultural areas. GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 323 Threskiornis spinicollis. Straw-necked Ibis. Abundance and distribution. Common and wide- spread. Flocks in excess of 100 birds were noted in ER between the Konnepra and Jilpanger Blocks and SW of the Toolondo Block; and^in AR near Mt. Dryden. Habitat. Margins of fresh- water swamps, lakes, rivers and farm ponds. It commonly forages in pastures. Breeding. Nesting occurred in GR on Bryan Swamp in 1960 and along the Wannon River near Bryan Swamp in 1968 (Cowling, unpub. MS). A rookery is also reported in GR on Shannon Swamp near Glenisla (Middleton pers. comm.). Conservation aspects As for White Ibis (above). Platalea regia. Royal Spoonbill. Abundance and distribution. Rare. Recorded in ER near Edenhope; in GR at Bryan Swamp and near Balmoral; and in AR at Lake Lonsdale. Habitat. Margins of swamps, rivers and lakes. Platalea flavipes. Yellow-billed Spoonbill. Abundance and distribution. Common and wide- spread. Habitat. Shallows of freshwater lakes, rivers, swamps and flooded pastures. Breeding! Young were present in GR during late December 1972 (McCulloch 1973). Between 10 and 20 nests were in ER at Boikerbert Swamp (Apsley) in 1972 (Thomas pers. comm.). ANATIDAE (GEESE, SWANS, DUCKS) Dendrocygna eytonL Plumed Tree-duck. Abundance and distribution. Rare; not observed in FWD survey. Since 1970 we have reports of it in ER at Clear Lake (Middleton pers. comm.) and on Lake Natimuk along the N boundary of the survey area (McQueen pers. comm.). Habitat. Freshwater lakes and swamps. Breeding. Local residents of ER state that it occasionally breeds in the region. However, we were unable to con- firm this. Cygnus atratus. Black Swan. Abundance and distribution. Common and wide- spread. Habitat. A wide range of aquatic habitats including saline lakes (e.g. White Lake, ER), freshwater lakes (e.g. Lake Lonsdale, AR), estu- aries (e.g. Lake Nambuk, HR), flooded moun- tain streams in GR, large swamps with dense emergent vegetation (e.g. NW margin of Morea Block in ER) and flooded pastures and wood- lands throughout the survey area. Breeding. Adults with broods were first recorded on 13 August and some cygnets were still considerably smaller than the adults in December. Between 13 August and 4 October the young counted in 20 broods averaged 4-9 cygnets per brood (range, 2-8). During late December of the relatively dry year of 1972 young swans were dying of starva- tion on some of the lakes which were drvina up (Mitchell 1973). Stictonetta naevosa. Freckled Duck. Abundance and distribution. Rare; not seen in FWD survey. A flock of about 30 was recorded in March-April of 1953 near the E boundary of HR (Shanks 1953). There are few other pub- lished records of this species in the vicinity of the survey area. Habitat. The flock was on a small saline lake of about 16 ha. Cereopsis novaehollandiae. Cape Barren Goose. Abundance and distribution. Rare; not seen in FWD survey. General sightings have been re- corded from ER and AR subsequent to 1970 (Middleton and Thomas pers. comm.). Specific sightings were recorded (Morris pers. comm.) from Lake Kennedy (E boundary of HR) on 11 February 1967 (a flock of 14) and on 3 August 1967 (a flock of 3). A single bird was seen in HR at Nareen in 1970 (Austin pers. comm.) and another was at Branxholme (near W boundary of HR) in July 1975 (Middleton pers. comm.). Habitat. During February of 1967 Morris (pers. comm.) provided the following comments on Lake Kennedy: open water which is surrounded by pasture; the geese can be seen regularly in the NE corner feeding in the vegetation growing over salty ground; the lake will probably dry up later in the year. Tadorna tadornoides. Mountain Duck. Abundance and distribution. Common in ER, AR and HR and uncommon in GR; widespread throughout. Large mobile flocks are formed after fledging of young. Habitat. Both open water and margins of fresh to saline lakes and swamps. Pas- tures with new plant growth or those which are flooded are extensively utilized. Breeding. Broods were seen September through November. Of 18 broods counted in ER, the average number of young in each was 8 9. Anas superciliosa. Black Duck. Abundance and distribution. Common and wide- spread. Habitat. Bodies of standing and flowing fresh water, particularly temporary sedge swamps and flooded or unflooded pastures with new plant growth. Breeding. A nest with 8 eggs was found 324 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS on 6 September 1974 in AR. It was on the ground in dense Kangaroo Grass (25 cm tall) in a road- side reserve of Yellow Gum woodland. Adults with ducklings were also noted in ER in early October 1951 (Stow et al 1952). Anas gibberifrons. Grey Teal. Abundance and distribution. The most common duck in the survey area and widespread through- out. Groups in excess of 500 birds have been re- cently recorded in ER near Douglas and on Lake Carchap; and in GR on Rocklands Reservoir. Habitat. Bodies of water ranging from flooded woodlands through seasonal swamps to large lakes and reservoirs. Also flowing waters but usually the slower moving, vegetated portions of streams or rivers were occupied. Breeding. Many broods were seen in ER between early October and early December. Of 11 broods counted, the average number of young in each was 7-2 (range, 6-8). Anas castanea. Chestnut Teal. Abundance and distribution. Rare; seen singly or in pairs in ER during FWD survey. Local con- centrations are occasionally reported, e.g. Karnak Swamp during early 1974 (Thomas pers. comm.). Habitat. Usually on large bodies of standing water. Anas rhynchotis. Shoveller. Abundance and distribution. Uncommon and widespread with local concentrations (e.g. near Goroke, ER, early in 1974) (Thomas pers. comm.) . Recorded in ER on Connangorach Swamp, near Goroke, S of Barton Swamp, near the Tooan Block and on Lake Mullancoree; in GR near Balmoral; in AR along the Western Highway (N of Mt. Zero); and on Lake Linlith- gow (near E boundary of HR). Habitat. Usually on temporary swamps and flooded pastures in spring and congregating on permanent lakes and swamps in autumn. Breeding. A brood of 14 young was seen in AR on 1 October. Malacorhynchus membranaceus. Pink-eared Duck. Abundance and distribution. Uncommon in ER with local concentrations, such as occurred on Karnak Swamp early in 1974 (Thomas pers. comm.). Mitchell (1973) also recorded 50 on Lake Mullancoree during late December 1972. Populations in Victoria fluctuate and largest num- bers occur apparently after wet years in inland Australia (Wright pers. comm.). This appeared to be the case in 1965 when large flocks were re- ported on Lake Linlithgow (near E boundary of HR) during late February-early March (Mor- ris pers. comm.) and in ER on Lake Carchap in late May (Thomas pers. comm.). Habitat. Large bodies of standing water. Breeding. A pair of adults with 2 young were seen on a temporary swamp along the Wimmera Highway W of the Kallungur Block, ER, on 4 October 1974. Aythya australis. White-eyed Duck. Abundance and distribution. Uncommon in ER with local concentrations near Goroke and Eden- hope in 1974 and near Douglas early in 1975 (Thomas pers. comm.). More than 500 were on Lake Mullancoree in December 1972 (Mitchell 1 973 ) . Habitat. Temporary and permanent swamps and lakes. Chenonetta jubata. Wood Duck. Abundance and distribution. Common and wide- spread. Pairs and small flocks, usually of less than 10 birds, were present August through December. After December flocks of more than 100 birds were on bodies of water such as Clear Lake and Toolondo Reservoir in ER; Bryan Swamp and Rocklands Reservoir in GR; Lake Lonsdale, Pine Lake and Lake Taylor in AR; and the Wannon River in HR. Habitat. Flooded pastures and woodlands, seasonal swamps and on farm ponds as well as foraging in unflooded pastures. Large bodies of flowing and standing freshwater were occupied subsequent to nesting. Breeding. Four broods, averaging 7-5 young in each, were seen in ER and AR between October and early December. Oxyura australis. Blue-billed Duck. Abundance and distribution. Uncommon in ER; not observed elsewhere. Recorded on Jaka Lake and Lake Wallace. Habitat. Both lakes have large areas of open deep water. Biziura lobata. Musk Duck. Abundance and distribution. Uncommon and widespread with local concentrations noted in ER on St. Marys Lake, White Lake and Lake Charle- grark; and in GR on Bryan Swamp and Moora Moora Reservoir. Habitat. Large, deep bodies of standing fresh water; occasionally on rivers and seasonal sedge swamps. ACCIPITRJDAE (KITES, HAWKS, EAGLES) Elanus notatus. Black-shouldered Kite. Abundance and distribution. Uncommon and widespread over the plains of the survey area. In GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 325 1952, population numbers were larger than nor- mal in the Coleraine area, HR, and Austin (1953a) indicated that a wide range of habitat was being utilized. Breeding. Nests in trees in pas- tures of HR (Austin pers. comm.) . Milvus migrans. Black Kite. Abundance and distribution. Rare; not seen in FWD survey. In 1952 a spectacular increase in numbers occurred in Victoria (Wheeler 1967a). Single birds began arriving in HR around Hamil- ton-Coleraine in February 1952 and by June had joined into flocks of up to 30 birds (Austin 1953a). Two such flocks occurred between Hamil- ton and Coleraine where they occupied territory and food with Whistling Kites. Austin (1953a) speculated that an outbreak of myxomatosis, and the consequent large numbers of rabbits killed on the roads, may have been responsible for the in- creased population of this species. Habitat. Open forest and plains country particularly near habi- tation (Wheeler 1967a). Feeding. The birds in HR during 1952 were feeding mainly on rabbits killed on roads (Austin 1953a). Haliastur sphenurus. Whistling Kite. Abundance and distribution. Common and wide- spread; usually as solitary or paired birds soaring overhead. Habitat. Semi-cleared pastures. Feed- ing. Rabbits are reportedly taken in, or near, the survey area (Hill 1907; Austin 1953a). Breeding. A common nesting bird in the survey area; some breeding details are given by Hill (1907a) for the Ararat District (near AR), by D'Ombrain (1905a) for the Casterton District (near the W boundary of HR), and by McCulloch (1973) for GR. Accipiter fasciatus. Brown Goshawk. Abundance and distribution. Common in GR, un- common elsewhere; widespread throughout. Habi- tat. Open forests; occasionally woodlands, shrub- lands and roadside trees in pastures. Feeding. Usually this goshawk catches and kills its own food, which consists mainly of vertebrates. How- ever, during an outbreak of myxomatosis in 1952, the goshawk was seen feeding on rabbit carcasses in HR near Coleraine (Austin 1953a). Breeding. Adults with young out of the nest were in GR near Balmoral during late December 1972 (Mc- Culloch 1973). Accipiter cirrhocephalus. Collared Sparrowhawk. Abundance and distribution. Rare; recorded in GR along the Glenelg River at Balmoral and S of Rocklands Reservoir. Habitat. The bird near Rocklands Reservoir was in a Yellow Gum/Yel- low Box woodland. Feeding. The individual in the woodland unsuccessfully chased a New Hol- land Honeyeater through the tree canopies. Haliaeetus leucogaster. White-breasted Sea-eagle. Abundance and distribution. Rare; not seen in FWD survey. Recorded in ER at Lake Wallace; in GR at Rocklands and Moora Moora Reser- voirs; and in AR at Lakes Lonsdale and Fyans. Habitat. Large freshwater lakes. Aquila audax. Wedge-tailed Eagle. Abundance and distribution. Common and wide- spread. The literature indicates that the popula- tion of this species in SW Victoria has remained relatively constant during the past 80 years. How- ever, precise information is lacking; this is un- fortunate because it would be relatively simple to locate ail the nests of this species within a de- fined area and then to follow the breeding popu- lation fluctuations over a long period of time. Habitat. It soars over all habitats, but most com- monly over pastures, woodlands and open rock formations. Feeding. Reported feeding on dead rabbits on roads in GR (Mitchell 1973). Breed- ing. About the turn of this century 10 or 12 pairs were nesting in the Ararat District (just E of AR) (Hill 1907a). Hill indicated that they sometimes nested on dead trees in open, level country, but more often in growing timber on hillsides. Hieraaetus morphnoides. Little Eagle. Abundance and distribution. Uncommon and widespread; most sightings during FWD survey were in ER. Habitat. Our sightings were of birds over pastures. However, Austin (1953a) saw five over timbered country. Breeding. Young were in nests in GR near Balmoral in late December 1972 (McCulloch 1973). Circus assimilis. Spotted Harrier. Abundance and distribution. Rare and wide- spread; not seen in FWD survey. Habitat. Grass- lands and croplands (Wheeler 1967a). Circus aeruginosus. Swamp Harrier. Abundance and distribution. Uncommon and widespread. Habitat. Swamps, lake and river mar- gins, pastures, heathy areas and croplands. Breed- ing. D'Ombrain (1905a) noted that this species often nested in the middle of a crop in the Cas- terton District (W boundary of HR), and that 326 W. B. EMISON f J. W. PORTER, K. C. NORRIS and G. J. APPS breeding commenced in September. Clutches generally contained three and occasionally five eggs. FALCONIDAE (FALCONS) Falco subniger. Black Falcon. Abundance and distribution. Rare; not seen in FWD survey but population increases occasionally occur. Between April 1951 and June 1952, at least 30 sightings of this species were made by Austin (1953a) in SW Victoria. He usually saw it singly, but on one occasion he did record a pair. Occasionally seen in GR near S end of Victoria Range in 1952-53 (McGarvie pers. comm.). Habitat. Austin (1953a) noted that all birds were in areas with little or no timber, and the only trees used as perches were isolated and dead. The observations by McGarvie were also in cleared areas. Feeding. An Australian Magpie Lark and a Common Skylark were taken (Austin 1953a). He also describes this falcon as hunting above and behind a Swamp Harrier as it flew over long grass. Falco per egr huts. Peregrine Falcon. Abundance and distribution. Common in GR and uncommon but with local concentrations else- where; widespread throughout. Habitat. Cliff faces which provide protection for nest sites and perching areas. Most cliffs overlooked pastures although a few overlooked open forests and woodlands. Feeding. Skeletal material found in eyries indicates that the introduced Domestic Pigeon is a major food source, but a wide range of other avian species are also taken (e.g. Gang- gang and Sulphur-crested Cockatoos, Crimson Rosellas, Common Starlings, White-faced Herons, cormorants, magpies and Masked Plovers). The location of eyries may influence the types of prey taken. For example, eyries above bodies of water contained remains of more aquatic species than did the eyries overlooking either pastures or open forests. Breeding. Nesting was docu- mented in ER and GR. Copulation was re- corded between 3 September and 4 October 1974. We followed the progress of two active eyries through the 1974 nesting season: one eyrie, lo- cated in a small recess close to the top of a cliff face, was found on 3 October; it contained three eggs which were being incubated. The nesting platform was level, dirt-covered and about 0-7 x 0-7 m in size. The cliff was near a major road in an area subjected to fairly intensive recreational use; the eyrie was easily accessible by a short climb up the back of the cliff. On 5 October three eggs and an adult were still present, but by 9 October only two eggs were in the eyrie and on 20 October the nest was empty. The second eyrie, located under an overhanging ledge at the base of a sheer cliff, was found on 12 September; it con- tained two eggs and one adult was present. To reach the eyrie a large, steep, rounded rock had to be climbed; the top of this rock was relatively level and extended to the base of the cliff where the eyrie was situated. The location of the eyrie was near a minor road in an area which, po- tentially, could be important for recreation. For- tunately, it has only light usage at the present time. On 30 September both eggs were present and two adults were flying over the eyrie. On 10 October both eggs were fractured and the young birds inside were vocalising; two adults were flying overhead. On 29 October, 2 November and 7 November two downy young and one adult were present; both chicks were banded on the last-mentioned visit. Our final inspection of the eyrie was made on 11 December when we ob- served one adult and two juveniles flying over- head. Conservation aspects. The abundance of this species in the survey area is of particular importance in view of the nearly world-wide population decline of this species (Hickey 1968). Studies initiated now, while this species is still in substantial numbers in Victoria, could assist in preventing similar, local population declines. Obviously, the data presented on the breeding of this species in the survey area suggest that access- ible eyries are vulnerable to human interference. Because of the tendency of this falcon to have tra- ditional nesting sites (White 1968) it is essen- tial that management authorities in areas which are, or will be, predominantly recreation-oriented be informed as to eyrie locations. Such informa- tion (which should not be made public) would not only allow a discreet surveillance of these sites (to discourage human interference) but, more importantly, would prevent the management authority from constructing roads, car parks, camp sites, picnic areas, nature walks, etc., near documented eyrie sites. Falco longipennis. Little Falcon. Abundance and distribution. Rare and wide- spread. Recorded in ER near North Lake; in GR near Balmoral; and in HR near Coleraine and near Wannon. Habitat. The bird near North Lake was over a pasture adjacent to a large block of Brown Stringybark shrubland. Feeding. We ob- served it pursuing starlings. GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 327 Falco berigora. Brown Falcon. Abundance and distribution. One of the most abundant raptors throughout the survey area. Usually seen singly or in pairs and only occa- sionally in threes or fours (family groups?). Habi- tat. Semi-cleared pasture and crop lands and oc- casionally woodlands, heathy areas, over swamps and on margins of open forests. Breeding. D'Ombrain (1905a) gave the following informa- tion on nests in the Casterton District (W bound- ary of HR) : five nests were found during Sep- tember 1903, all in River Red Gums. The clutch size was three in every case. Falco cenchroides. Nankeen Kestrel. Abundance and distribution. Uncommon and widespread. Usually seen singly, as pairs or, rarely, in groups of up to six birds. Habitat. Semi-cleared pasture and crop lands, woodlands, open forests, heathy areas, rock cliffs and rocky offshore pinnacles. Breeding. D'Ombrain (1905a) noted that this species nested in the Cas- terton District (W boundary of HR) between October and November. He stated that the clutch is generally four, but frequently five, and gave specific information on a nest found in a hole in a River Red Gum on 28 October 1903. PHASIANIDAE (QUAILS) Coturnix pectoralis. Stubble Quail. Abundance and distribution. Common and wide- spread except in timbered and mountainous areas. During the past 15 years large numbers have sometimes been recorded near Lake Linlithgow (E boundary of HR), in the S portion of AR and in the Natimuk area of ER by Morris and Thom- as (pers. comra.), Habitat. Pastures and crop lands, particularly those containing wheat, lin- seed, hog weed or new growths of grass. Breed- ing. Reported from various places in the survey area (Morris, Thomas, Austin and McGarvie pers. comra.). Coturnix australis. Brown Quail. Abundance and distribution. Not seen during FWD survey. Recorded in GR at Marney and Bryan Swamps and at Rocklands Reservoir dur- ing late December 1972 (Mitchell 1973). Habit- at. Margins of freshwater swamps and reser- voirs. Breeding. McCulloch (1973) noted young in GR during late December 1972. x TURN1CIDAE (BUTTON-QUAILS) Turnix varia. Painted Button-quail. Abundance and distribution. Uncommon and widespread. Habitat. Open forests, woodlands and shrublands, particularly where there is little ground cover. Turnix velox. Little Button-quail. Abundance and distribution. Not seen during FWD survey. Hill (1907b) recorded it during some summers before 1907 E of Ararat (near E boundary of AR) and Austin (1951) recorded it in 1949-50 near Casterton (just W of HR) and near Edenhope, ER. Habitat. Austin (1951) saw this quail in light bracken in open country. Breed- ing. The birds near Casterton were reported nest- ing in 1949-50. PEDIONOMIDAE (PLAINS-WANDERER) Pedionomus torquatm. Plains-wanderer (Plate 21, Fig. 6). Abundance and distribution. Rare; not seen in FWD survey. A dead bird was found N of Horsham (N boundary of survey area) in May 1973, and was sent to NMV where it is lodged as a specimen (W 9092). D'Ombrain (1926) noted it in HR near Hamilton in 1922. Habitat. Flat, open, grass-covered plains (Frith 1969). Conser- vation aspects. Early during European settlement it was common on the plains in Victoria (D'Om- brain 1926), but a dramatic population decline has occurred throughout its range and today it is seldom seen. Studies to determine its present Victorian status are required. RALLIDAE (RAILS, CRAKES, SWAMPHENS) Rallus philippensis. Land Rail. Abundance and distribution. Not seen in FWD survey; recorded recently in AR at Stawell (Mc- Cann pers. comm.) and in HR at Nareen (Aus- tin pers. comm.) but there are few other recent literature records for the survey area. Some old, general records of it occurring in or near the survey area include: Anon. (1892); Hill (1907b); and Sullivan (1929a). Habitat. Swampy con- ditions, crops and cultivation, and, at times, fores- ted areas (Wheeler 1967a). Breeding. Five eggs were found in a nest at Warrnambool (just E of HR) during late October 1928 (Sullivan 1929a). 328 W. B. KMISON, J. W. POR'IhR, K. C. NORRIS and G. J. APPS Rallus pcctoralis. Water Rail. Abundance and distribution. One bird was cap- tured alive in a mammal trap in GR on the E side of Lake Wartook in FWD survey; it was later released in the same area. Also recently re- ported in GR at Wannon Bridge (McCann pers. comm.) and in ER at Horsham (Thomas pers. comm.). Habitat. The bird trapped near Lake Wartook was in dense wet sedges under a shrub layer of Scented Paperbarks. Porzana pusilla. Marsh Crake. Abundance and distribution. Not seen in FWD survey and there are few literature references from the survey area. However, because of the dense nature of its habitat, it may be more abund- ant than is suggested by the few recorded sight- ings. Recorded in GR near Balmoral (Mitchell 1973); and in AR- at Lake Fyans (McCann pers. comm.). Habitat. Dense vegetation in swamps. Porzana jiuminca. Spotted Crake. Abundance and distribution. Not seen in FWD survey. The general remarks made for the Marsh Crake also apply here. Mitchell (1973) reported it as common in GR near Balmoral in December 1972; and McCann (pers. comm.) has recently observed it in AR at Lake Lonsdale and near Stawell. The NMV has specimens from Eden- hope and Stawell. Habitat. Dense vegetation bor- dering swamps and rivers. Porzana tabuensls. Spotless Crake. Abundance and distribution. Not seen in FWD survey. Recorded in ER along the Glenelg River near Harrow (Austin pers. comm.); in GR near Balmoral (Mitchell 1973); and in AR at Lake Lonsdale (McCann pers. comm.). Habitat. Dense vegetation in swampy areas. Gallinula ventralis. Black-tailed Native-hen. Abundance and distribution, A few solitary and paired birds were in ER (usually along the N boundary) during FWD survey. The numbers of this species had been high for two or three years before our survey (Middleton pers. comm.). Such population fluctuations are normal and Wheeler (1967a) remarks that it appears in great numbers during, and immediately after, flood periods. Habitat. Temporary or permanent freshwater swamps, lakes and flooded pastures. Breeding. A pair with a brood of chicks was in ER near the Mt. Arapiles Block in early October 1974. Gallinula tenebrosa. Dusky Moorhen. Abundance and distribution. Uncommon and widespread. Habitat. Large permanent bodies of fresh water; also flooded woodlands and other temporary bodies of water during spring. Breed- ing. Adults with chicks were seen in a flooded River Red Gum woodland in ER near the Talla- geira Block on 7 December 1974. Young moor- hens were seen also in GR near Balmoral during late December 1972 (McCulloch 1973). Porphyrio porphyrio, Swamphen. Abundance and distribution. Common and wide- spread. Habitat. Margins of seasonal swamps and flooded pastures or woodlands; shorelines of large permanent bodies of water are important during late summer and early autumn. Breeding. Young were on swamps in GR near Balmoral in late December 1972 (McCulloch 1973). A regu- lar breeder in AR at Lake Fyans (McCann pers. comm.). Fuiica atra. Coot. Abundance and distribution. Common and wide- spread. Habitat. Freshwater and saline lakes, swamps and flooded pastures; occasionally on slow-flowing rivers and streams. Seen both on open water and on vegetated margins. Breeding. McCulloch (1973) noted adults with young in GR in late December 1972. GRUIDAE (CRANES) Grus rubicitnda. Brolga. Abundance and distribution. Uncommon and widespread. Habitat. Pastures or swamps on the plains, particularly semi-cleared pastures with some low-lying, poorly drained areas. Occasion- ally in stubble fields, but usually near swamps. Breeding. Nesting is reported in all regions of the survey area. Conservation aspects. We suggest that a field study to determine the status of this species in Victoria be conducted. The results of this study might suggest areas which, if set aside as reserves, could lead to a permanent nu- cleus of successful breeding birds being estab- lished in the State. BURH1NIDAE (STONE-CURLEWS) Burhinus magnirostris. Bush Stone-curlew. Abundance and distribution. Rare; not seen in FWD survey. Hill (1907b) presented the follow- ing information on the status of this species in the Ararat District, shortly after the turn of the GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 329 century: 'fairly numerous on the timbered rises, but, according to old residents, they are less numerous than formerly, which may be accounted for by the increase in the number of foxes'. Other factors possibly also responsible for their decline include predation by feral Cats, clearing of the woodlands and agricultural activities. Between 1919 and present other sightings or calls have been reported in ER near Lake Wallace (Wheeler 1964); in GR (Audas 1920) and N of Mt. Dun- das Block (McGarvie pers. comm.); in AR at Illawarra Block (Middleton pers. comm.) and along the Wimmera River near Horsham (Thomas pers. comm.); and in HR near Caven- dish and Wannon (Wheeler 1967b; Mitchell 1973). Habitat. Noted in Yellow Gum wood- land in the Illawarra Block (Middleton pers. comm.) and in River Red Gum areas N of Mt. Dundas Block (McGarvie pers. comm.). Con- servation aspects. Studies are required to deter- mine the reasons for the population declines of this species. In particular, the effects of the loss and alteration of its woodland habitat should be examined. ROSTRATULIDAE (PAINTED SNIPE) Rostratula benghalensis. Painted Snipe. Abundance and distribution. Rare. Four in GR at McGlashan's Swamp (near Balmoral) in late December 1972 (Mitchell 1973) and two N of Mt. Dundas Block in October and November 1957 (McGarvie pers. comm.). Habitat. Swamps. HAEMATOPODIDAE (OYSTERCATCHERS) Haematopus ostralegus. Pied Oystercatcher. Abundance and distribution. Common along the S boundary of HR (Isles pers. comm.). Habitat. Sandy ocean beaches and tidal flats. CHARADRIIDAE (LAPWINGS, PLOVERS AND DOTTERELS) Vanellus miles. Masked Plover. Abundance and distribution. Common and wide- spread. Pairs or small flocks were present during spring and early summer; in mid-December a few larger flocks were evident and by March-April flocks containing 20 to 50 birds were common. Habitat. Margins of permanent and temporary lakes, swamps, reservoirs, etc., particularly where receding water has left exposed flats. Also areas of flooded vegetation, pastures with new growth and crops such as lucerne. Breeding. Adults with young were seen during September and October 1974. A nest of four eggs was found in the Vic- toria Valley, GR, on 5 September 1974. It was in a depression on a slight rise and was lined with grasses. Another nest with four eggs was on a mud bank near Hamilton, HR, on 27 August 1971 (RAOU nest card). Vanellus tricolor. Banded Plover. Abundance and distribution. Not seen in FWD survey. Flocks of up to 75 birds were in GR near S end of Victoria Range in May 1954 (Mc- Garvie pers. comm.). Two were in GR along Victoria Range Road in December 1972 (Mit- chell 1973). Reported regularly at Concongella (just E of AR) near Stawell (McCann pers. comm.). Habitat. Swamps, lagoons, and a strong preference for grassy plains often far from water (Condon and McGill 1974). The birds seen by McGarvie were in open country. Breeding. Nests regularly at Concongella (McCann pers. comm.). Nesting recorded in HR near Coleraine (RAOU nest cards). A regular nesting bird in GR near S end of Victoria Range (McGarvie pers. comm.). Erythrogonys cinctus. Red-kneed Dotterel. Abundance and distribution. Rare; only recorded during FWD survey along the N boundary of ER at Mitre Lake (a flock of 4 in October 1974). McCann (pers. comm.) has seen it in AR at Lake Lonsdale and it has been reported elsewhere in the survey area by Sullivan (1929a), Wheeler (1964) and Mitchell (1973). Habitat. Margins of standing bodies of water. Charadrius rubricollis. Hooded Dotterel. Abundance and distribution. Common along the S boundary of HR (Isles pers. comm.). Habitat. Ocean beaches, Charadrius bicinctus. Double-banded Dotterel. Abundance and distribution. Common during win- ter months along S boundary of HR (Isles pers. comm.). Rare inland; recorded in ER at Lake Wallace (Wheeler 1 964) . Habitat. Ocean beaches and, occasionally, inland swamps and lake margins. Movements. A winter migrant to Vic- toria, arriving as early as February and leaving in August (Wheeler 1967a). Breeds in New Zea- land (Frith 1969). Charadrius ruficapillus. Red-capped Dotterel. Abundance and distribution. Uncommon in ER. Most sightings were of small flocks, each with less than 10 birds, in October 1974, but a flock of about 80 birds was near Douglas on 15 April 330 W. B. EMISON, J. W. PORTER, K. C. NORRIS and O. J, APPS 1975. Habitat. Dry margins or beds of freshwater and, in particular, salt lakes. Breeding. Young were seen in ER at White Lake on 5 October 1957 (Bald 1957a). Also recorded nesting in GR (McGarvie pcrs. comm,). Charadrius mclanops. Black-fronted Dotterel. Abundance and distribution. Uncommon and widespread. Habitat. Margins, particularly of sand or mud, of permanent and temporary bodies of water. Sometimes it nests far from water. Breeding. A nest in a depression in the ground and lined with bark contained two eggs on 3 Oc- tober 1974. It was in a Yellow Box/ Yellow Gum woodland about 100 m from a large temporary lake in the Toolondo Block, ER. The undergrowth was sparse and appeared heavily grazed. Also re- corded nesting in GR (McGarvie pers. comm.); in AR near Stawell (McCann pers. comm.); and in HR near Coleraine (RAOU nest card). RECURVIROSTRIDAE (STILTS, AVOCETS) Himantopus himantopus. Pied Stilt. Abundance and distribution. Uncommon and widespread; most sightings are from ER. Re- corded in ER at Lake Wallace, lakes near Doug- las, Lake Natimuk (N boundary of ER) and 2 km SW of Mt. Arapiles Block; in AR at Lake Lonsdale; and in HR near Macarthur. Habitat. Margins of lakes and swamps. Movements. Rela- tively large numbers are seen during dry years such as 1972 (Mitchell 1973) and few arc seen during wet years such as 1974 (FWD survey). Possibly they move out of the arid interior to the few permanent lakes and swamps remaining on the periphery of their range during dry periods. Cladorhynchus leucocephahis. Banded Stilt. Abundance and distribution. Uncommon, only seen in ER. Recorded at Centre Lake (9 birds in October 1974) and White Lake (40 birds in April 1975). A flock of 800 was on White Lake and another flock of approximately the same size was on Centre Lake on 5 October (Bald 1957a). Austin (1962b) recorded 10,000 birds on the lakes at Douglas and implied that such numbers might be smaller than usual. Habitat. Shallow open margins or sandbars of salt lakes. Occasion- ally associated with Pied Stilts, Red-necked Avo- cets and Black Swans. Breeding. Of 9 birds at Centre Lake, two had full chest bands and the rest were without or had indistinct bands. About one-third of the 40 on White Lake had full chest bands. Most of the 800 birds on White Lake in 1957 had full chest bands (Bald 1957a). Con- servation aspects. The salt lakes in ER near Douglas appear to be one of the main inland areas for this species in Victoria. Recurvirostra novaehoUandiae. Red-necked Avocet. Abundance and distribution. Uncommon, only seen in ER. Reported on Centre Lake (70 birds) in October 1974 and on Mitre Lake and a lake N of Lake Natimuk (N boundary of ER). Austin (1962b) reported 10,000 on the lakes near Doug- las in 1962. They were with about 100 Pied Stilts and 10,000 Banded Stilts. Habitat. Shallow open margins of salt lakes. SCOLOPAC1DAE (SANDPIPERS, SNIPE) Tringa ncbularia. Greenshank. Abundance and distribution. Rare; not seen in FWD survey. Mitchell (1973) recorded the fol- lowing for late December 1972: Two L. Nati- muk; three McGlashan's Swamp; two L. Mulla'. All three localities are either in or near ER. Habitat. Inland swamps and shallow lakes. Move- ments. Breeds in N Europe and N Asia and mi- grates in winter through E Asia to Australia (Frith 1969). Gallinago hardwlcku. Japanese Snipe. Abundance and distribution. Uncommon and widespread. Habitat. Dense vegetation in marshes, swamps and wet heaths. Movements. Breeds in Japan and migrates S in winter through New Guinea to Australia (Frith 1969). Calldris acuminata. Sharp-tailed Sandpiper. Abundance and distribution. Uncommon; not seen in FWD survey. Flocks of up to 500 birds have been reported in or near ER at White and Centre Lakes, Lake Natimuk and Lake Mullancoree; and in GR on Bryan and Marney Swamps. Habitat. Margins of saline and freshwater lakes and swamps. Movements. Breeds in NE Siberia and migrates through E Asia to Australia (Frith 1969). Calldris ruficollis. Red-necked Stint. Abundance and distribution. Common during summer months along S boundary of HR (Isles pers. comm.). Uncommon inland; a flock of about 70 was in ER on the margin of a lake near the Mt. Arapiles Block in October 1974. Also re- corded on White and Centre Lakes in October 1957 (Bald 1957a). Habitat. Ocean and estuary GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 331 beaches, tidal flats, margins of saline and fresh- water lakes and swamps. Movements. Breeds in NE Siberia and W Alaska and migrates through Asia to Australia (Frith 1969). Calidris ferruginea. Curlew Sandpiper. Abundance and distribution. Rare; not seen in FWD survey. Recorded at Lake Natimuk (N boundary of ER) in December 1972 (Mitchell 1973). Habitat. Margins of lakes and swamps. Movements. Breeds in E Arctic Asia and mi- grates in winter through E Asia to Australia (Frith 1969). Calidris alba. Sanderling. Abundance and distribution. Rare; a flock of 80 was in HR near Lake Yambuk on 23 July 1976 (Isles pers. comm.). Habitat. Ocean beaches. Movements. Breeds N of the Arctic Circle, some migrate to Australia where they are widespread from November to May. Many birds overwinter in Australia (Condon 1969). LARIDAE (GULLS, TERNS) Larus novaehollandiae. Silver Gull. Abundance and distribution. Uncommon and widespread. Usually in flocks of 10-100 individu- als. Habitat. Saline and freshwater lakes and swamps; flocks often occur around food sources far from water. Chlidonias hybrida. Whiskered Tern. Abundance and distribution. Uncommon and widespread. Occasionally in flocks of up to 300 birds (Sullivan 1929b) . Habitat. Saline and freshwater lakes and swamps, flooded pastures and freshly ploughed croplands (where they feed on invertebrates). Breeding. Three nests were found in GR near Dunkeld on 22 December 1968 (RAOU nest cards). Gelochelidon nilotica. Gull-billed Tern. Abundance and distribution. Rare; in 38 years Austin (pers. comm.) recorded three small flocks flying over Nareen in HR. COLUMBIDAE (PIGEONS AND DOVES) Colomba livia. Domestic Pigeon. Abundance and distribution. Uncommon and widespread. Usually in small flocks, seldom of more than 20 birds. Analyses of wing bones found in Peregrine Falcon eyries in the survey area indicate that some nesting pairs rely on the pigeons as a food source when feeding young. Habitat. Areas extensively modified by human oc- cupation (e.g. towns, farm buildings, cleared pas- tures, bridges, etc.). Geopelia striata. Peaceful Dove. Abundance and distribution. Uncommon in ER, GR and AR; not seen in HR. Habitat. In ER at the Mt. Arapiles Block we observed this species in Grey Box and Yellow Gum woodlands. Breed- ing. Commonly breeds in AR near Stawell; it has colonised this area within the past 25 years (McCann pers. comm.). Geopelia cuneata. Diamond Dove. Abundance and distribution. Rare; possibly only an accidental visitor to the survey area. Seen in ER near Lake Wallace in March 1964 (Wheeler 1964). Habitat. No data available for survey area. Phaps chalcoptera. Common Bronzewing. Abundance and distribution. Common and wide- spread in ER, GR and AR; not seen in HR. Habitat. Dense shrub layers. Dominant tree spe- cies are probably less important than the density of the shrub layers. Recorded in such diverse tree associations as Yellow Gum/ Yellow Box, Brown Stringy bark/ Small Grass Tree, Messmate, and Brown Stringybark/ Manna Gum. Breeding. Nests in AR near Stawell (McCann pers. comm.) . Phaps elegans. Brush Bronzewing. Abundance and distribution. Rare; recorded in GR at Balmoral, Victoria Valley and along Gram- pians Road near Mt. William turn-off; and in HR near Hamilton. Habitat. In Victoria Valley the birds were in heaths under Scent-barks (p.c.-26). Ocyphaps lophotes. Crested Pigeon. Abundance and distribution. Uncommon and re- stricted mainly to N portions of ER and AR. Also recorded in ER S of Toolondo Block (Mitchell 1973) and near Stawell along Concongella Creek (just E of AR) (McCann pers. comm.). Habitat. Sightings along the N boundary of the survey area were usually in pasture lands. Breeding. Nests along Concongella Creek, an area it has colonised within the past 25 years (McCann pers. comm.) . CACATUIDAE (COCKATOOS) Calyptorhynchus magnificus. Red-tailed Black Cockatoo. Abundance and distribution. Rare and restricted to ER. Recorded in the Kadnook, Tallegeira and Jilpanger Blocks during FWD survey. Previously 332 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS reported in the Edetthope and Harrow districts (Austin 1951), the Edenhope region (Attiwill I960), in the Toolondo Block (Middleton pens. COmm.), on the Moree Bridge Road (Wheeler 1^64) and along the Edcnhopc Road (Mitchell 1973). The NMV has three specimens from ER. Habitat. Brown Stringybark shrubland. Feeding. Our observations of this species feeding on un- ripened Brown Slringybark seeds confirm similar observations made by Austin (1951) and Attiwill (I960). The latter author also states The birds also feed on the seeds of banksias, hakcas, small casuarinas and other native shrubs, particularly after a bushfire has passed over a patch of this type of country. Very rarely, they have been ob- served on the ground feeding on prickly burrs, am! they also bite into dead timber in search of the larva of wood-boring beetles'. Breeding. Atti- will (I960) gives six recent nesting records for ER: three during October/November and three during February/ March. The nests were in large hollows in cuealypts. Conservation aspects. The Victorian range of this species has apparently de- creased since the early days of settlement (Wheeler 1967a) and it is now restricted to the SW part of the State with ER being one of its last strongholds. Conservation of some 6f the larger areas of Brown Stringybark in this region is es- sential to the local survival of this species. Calyptorkynchus ftmercus. Yellow-tailed Black Cockatoo. Abundance and distribution. Uncommon and widespread. Habitat. Usually seen in flight over a wide range of habitats. The few sightings of birds not in (light were made in woodlands (Yel- low Gum/ River Red Cium, Yellow Gum/banksia and River Red Gum/Silver Banksia), Brown Stringybark shrubland and a pine plantation. Feeding. Five were feeding on seeds from Desert Banksia cones in the Tallageira Block (ER) and three were carrying pine cones when they flew from a roadside stand o( introduced pines W of the Jilpanger Block (ER). Breeding. An adult was on a nest 5 m up in a burnt Messmate on 22 November 1969 (Mt. William, GR) (RAOU nest card). Callocephalon fimbriatunu Gang-gang Cockatoo. Abundance and distribution. Uncommon and widespread in GR; rare in ER. Seldom seen further W than the Victoria-South Australia bor- der (Condon 1969); Middleton (pers. eomm.) suggests that the N boundary of ER is the N limit of its range in W Victoria. Habitat. Mainly open forests, particularly of Brown Stringybark. Rare in woodlands (Yellow Gum/Yellow Box). Feeding. Observed feeding on Messmate and Scrub Cypress Pine seeds during FWD survey. Breeding. McCulloch (1973) recorded young out of the nest in GR during December 1 972. Caeatua roseicapilla, Galah. Abundance and distribution. Uncommon and widespread with local concentrations in ER near the Mt. Arapiles Block. Flocks usually contained less than 10 birds and occasionally Galahs were with flocks of either Sulphur-crested Cockatoos, Long-billed Corellas or Red-rumped Parrots. Comments on the southward range expansion of this species in the W portion of the survey area are made by Austin (1951). Habitat. Mainly pas- tures; rarely woodland remnants (Grey Box and Yellow Gum/Grey Box) on pasture land mar- gins. Caeatua tenuirostris. Long-billed Corella. Abundance and distribution. Common and wide- spread; usually in flocks, occasionally of up to several hundred birds, with Sulphur-crested Cockatoos. On 3 October 1974, we counted a total of 182 Long-billed Corellas along a N-S drive of 31 km between the Mt, Arapiles Block and the E side of the Toolondo Block (ER). These birds were in nine separate flocks which ranged in size from 1 to 60 individuals. Habitat. Pastures. Large woodland trees such as River Red Gum and Yellow Gum are selected for roost- ing and nesting. These trees are either isolated in pastures or are parts of extensive woodlands, par- ticularly near areas of open water. Feeding. Oats, wheat and Onion-grass bulbs are taken (Sullivan 1929c; Smith 1962). Breeding. Adults with fledged young were in ER near Noradjuha in No- vember 1961, and they were also nesting in trees in a swamp about 32 km SE of Noradjuha (Smith 1962). Nesting also occurs in AR in dead trees in Lake Fyans (Smart pers. comm.). Con- servation aspects. The survey area is one of the last strongholds of this species in southern Aus- tralia and a concerted effort to ensure its con- tinued survival in SW Victoria is required. Im- mediate studies should be conducted to elucidate its life history as well as its impact on cereal crops grown in the area. It is important that methods designed to alleviate its impact on cereal crops arc critically reviewed before implementa- tion to ensure that no adverse long-term effects on the population will occur. GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 333 Cacatua galerita. Sulphur-crested Cockatoo. Abundance and distribution. Common and wide- spread; usually in flocks of up to several hundred birds with Long-billed Corellas. Habitat. Pastures. Large woodland trees such as River Red Gums and Yellow Gums are used for nesting and roost- ing. Feeding. Seen feeding in GR on Onion-grass bulbs and on oats. Breeding. Nesting recorded in GR (McGarvie pers. comm.). LORIIDAE (LORIKEETS) Trichoglossus haematodus. Rainbow Lorikeet. Abundance and distribution. Rare and widespread. Recorded in ER between Benayeo and the Talla- geira Block, near Edenhope, S of the Tooan Block, W of the Toolondo Block, and Horsham; in GR at S end of Victoria Range, Victoria Val- ley, Balmoral and N of Rocklands Reservoir; and in AR at Dadswell Bridge and Stawell. Habitat. Woodlands, particularly those containing Yellow Gum, Yellow Box or River Red Gum. Conserva- tion aspects. As for Musk Lorikeet (below) . Glossopsitta concinna. Musk Lorikeet. Abundance and distribution. Common and wide- spread. Habitat. Yellow Gum woodlands, some- times in association with Yellow Box, River Red Gum or both. Feeding. Nectar/pollen from Yel- low Gum blossoms is commonly taken; also seen feeding on unopened flowers of River Red Gums. Breeding. Recorded nesting in GR at Cherrypool in 1954 (McGarvie pers. comm.). Conservation aspects. The extensive loss of woodlands in the survey area may ultimately result in population de- clines of this species. The few segments of wood- land remaining in the survey area should be care- fully managed to provide a continual nectar source for nomadic birds such as the lorikeets. Glossopsitta porphyrocephala. Purple-crowned Lorikeet. Abundance and distribution. Uncommon and widespread; locally common in ER at Mt. Ara- piles Block in April 1975. Habitat. Woodlands; our records include River Red Gum and Grey Box woodlands. Feeding. At Mt. Arapiles Grey Box blossoms were being selected. The appear- ance of lorikeets in these trees coincided with the appearance of the blossoms. Conservation aspects. As for Musk Lorikeet (above). Glossopsitta pusilla. Little Lorikeet. Abundance and distribution. Uncommon and widespread; not seen in FWD survey. Recorded in ER near Edenhope (Wheeler 1964); in GR near Balmoral and Rocklands Reservoir (Mitchell 1973); and in AR near Stawell (McCann pers. comm.). Habitat. Woodlands. Conservation as- pects. As for Musk Lorikeet (above). POLYTELITIDAE (LONG-TAILED PARROTS) Nymphicus hollandicus. Cockatiel. Abundance and distribution. Not seen in FWD survey. Reported in most suitable areas in GR around Balmoral and the Black Range in Decem- ber 1972 (Mitchell 1973). Wheeler (1967a) describes this species as one whose numbers vary considerably from year to year. PLATYCERCIDAE (BROAD-TAILED PARROTS) Melopsittacus undulatus, Budgerygah. Abundance and distribution. Rare; not seen in FWD survey. Recorded in GR near Balmoral in December 1972 (Mitchell 1973). Habitat. Mainly open forests in the northern districts (Wheeler 1967a). Movements. Predominantly an inland spe- cies which experiences occasional northern move- ments (Wheeler 1967a; Condon 1969). Lathamus discolor. Swift Parrot. Abundance and distribution. Rare and wide- spread; not seen in FWD survey. Recorded in ER near Edenhope in March (Wheeler 1964) and near Lar-arum in May (Middleton pers. comm.); in GR N of the Mt. Dundas Block in September 1956 (McGarvie pers. comm.); in AR near Sta- well in October (McCann pers. comm.); and in HR at Coleraine in March (Austin 1953b). Habi- tat. Woodlands. Feeding. A flock of 12 was feed- ing on blossoms of a Red-flowering Gum {Euca- lyptus ficifolia) at Coleraine (Austin 1953b). Movements. Breeds in Tasmania (Frith 1969; Sla- ter 1970) and generally is only present in SE Aus- tralia between March and September. Conserva- tion aspects. As for Musk Lorikeet (above). Platycercus elegans. Crimson Rosella. Abundance and distribution. Common and wide- spread; particularly abundant in GR. Habitat. Open forests, woodlands, shrublands; occasional in heaths and in trees remaining in pastures. Brown Stringybark associations were particularly well populated, but also seen in plant associa- tions composed of one or more of the following trees: River Red Gum, Scent-bark, Manna Gum, 334 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS Yellow Gum, Yellow Box, Mountain Grey Gum, Messmate, Grey Box, Shining Peppermint and Swamp Gum. Breeding. Nesting recorded in GR (McGarvie pers. comm.). Platycercus elegans. Crimson Rosella. Abundance and distribution. Common and wide- spread, although few birds occur in the mountain ranges of GR. Habitat. Woodlands and semi- cleared pastures. Most common in River Red Gum associations, but also in plant associations of Yellow Gum, Yellow Box, Grey Box or Manna Gum. Breeding. Nesting recorded in GR (Mc- Garvie pers. comm.). Psephotus haematonotus. Red-rumped Parrot. Abundance and distribution. Common and wide- spread, although few birds occur in the mountain ranges of GR. Particularly abundant along N and W boundaries of ER. Habitat. Semi-cleared pas- tures where the few remaining trees arc either River Red Gum, Yellow Gum or Grey Box. Un- like the Eastern Rosella, this parrot seldom occurs in woodlands but uses woodland margins that open onto pasture. Breeding. Adults were feeding fledged young in ER between Apsley and the Tallagcira Block on 7 December 1974. Adults also noted as feeding young out of the nest in GR in late December 1972 (McCulIoch 1973). Neophema chrysostoma. Blue-winged Parrot. Abundance and distribution. Uncommon. One was in ER in the Kadnook Block in December 1974 (FWD survey) and Wheeler (1964) saw one near Wrattonbully, South Australia (about 10 km SW of Lankoop in ER) in March 1964. Storr et al. (1952) recorded it in HR near Hay- wood (about 15 km S of the Stones Block) in October 1951 and we saw a flock of about 20 im- mature Neophema parrots, which could have been either Blue-winged or Elegant Parrots, in the Stones Block in February 1975. Five were near Lake Yambuk (HR) on 27 July 1975 (Isles pers. comm.). Habitat. The bird in the Kadnook Block was in Brown Stringybark shrubland, the imma- ture parrots were in Manna Gum open forest and the birds near Lake Yambuk were on a primary dune. Feeding. The individual in the Kadnook Block was feeding on Leucopogon sp. fruits and the five near Lake Yambuk were eating seeds of Cakile sp. Neophema elegans. Elegant Parrot. Abundance and distribution. Rare. Sullivan (1929a) recorded two flocks near Lake Condah (HR) in July 1928 and we saw a flock of imma- ture Neophema parrots, which could have either been Elegant or Blue-winged Parrots, in the Stones Block in February 1975. Habitat. The birds near Lake Condah were feeding among heaps of scattered straw and the flock of imma- tures was in a Manna Gum open forest. CUCULIDAE (CUCKOOS) Cuculus pallidus. Pallid Cuckoo. Abundance and distribution. Uncommon and widespread. This migratory species has been re- corded in the survey area as early in the spring as 9 August (Sullivan 1929a). Interestingly, it was present in substantial numbers in GR on 18-19 October 1972 (Wheeler and Pescott 1973), but only two juveniles were recorded there dur- ing late December of the same year (Mitchell 1973). Thus, it may follow a similar pattern to that in the Canberra area where, after breeding, the adults move away in the early weeks of the New Year but juveniles remain until the end of April (Frith 1969). Habitat. Brown Stringybark shrublands, heaths, open forests and trees in pas- tures. Breeding. Juveniles have been seen being fed by White-naped, New Holland and Fuscous Honeyeaters. Movements. Not well known and apparently complex, but it is absent from SE Australia during winter. Cuculus pyrrhophanus. Fan-tailed Cuckoo. Abundance and distribution. Uncommon and widespread; usually seen from September to March (inclusive), but one was in HR near Lake Yambuk on 27 July 1976 (Isles pers. comm.). Habitat. Brown Stringybark and Manna Gum open forests, Yellow Gum/Yellow Box woodland and River Red Gum in pastures. Breeding. Nest- ing recorded in GR (McGarvie pers. comm.). Movements. Not well known, apparently there is some migration away from SE Australia during winter, but it may not involve the entire popula- tion (Slater 1970). Wheeler (1967a) indicates that many winter in Victoria. Chrysococcyx osculans. Black-eared Cuckoo. Abundance and distribution. Rare; not seen in FWD survey. Recorded in GR near Balmoral in December 1972 (Mitchell 1973). Also recorded as rare in the Ararat District (E of AR) by Hill ( 1 907b) . Habitat. Open forests and plains (Wheeler 1967a). Breeding. Hill (1907b) re- corded eggs in nests of Speckled Warblers and Superb Blue Wrens. Movements. Not well under- GRAMPIANS-EDENHOPE VERTEBRATE FAUNA SURVEY 335 stood, but most S populations move N during winter (Macdonald 1973). Chrysococcyx basalts. Rufous-tailed Bronze- cuckoo. Abundance and distribution. Uncommon and widespread. More plentiful than the Shining Bronze-cuckoo in the survey area (Middleton pers. comm.). Habitat. Open forests (particularly of Manna Gum) and woodlands. Breeding. Foster parents in the Ararat District (E of AR) include Scarlet Robins, BufT-rumped Thornbills and Su- perb Blue Wrens (Hill 1907b). Movements. Largely migratory, especially S breeding popula- tions, most of which leave for the North in March and return S in September (Macdonald 1973). Many winter in Victoria (Wheeler 1967a). Chrysococcyx lucidus, Shining Bronze-cuckoo. Abundance and distribution. Uncommon and widespread; less plentiful than the Rufous-tailed Bronze-cuckoo in the survey area ( Middleton pers. comm.). Habitat. Open forests and wood- lands. Breeding. Fledglings have been seen in ER in the Mt. Arapiles Block (Middleton pers. comm.). Movements. A summer migrant to Vic- toria and it winters within the State only on rare occasions (Wheeler 1967a). STRIGIDAE (OWLS) Ninox strenua. Powerful Owl. Abundance and distribution. Uncommon and re- stricted to GR; seldom recorded far from the mountain ranges in the survey area. Recorded from Victoria Valley (FWD survey), Pomonal and Roses Gap (McCann pers. comm.), N of Yarram Park (Grey pers. comm.) and Billywing (Austin 1962c; Hall and McKean 1962). Habi- tat. Open forest. The bird in Victoria Valley was in an area of Swamp Gum with dense sedge un- dergrowth. Ninox novaeseelandiae. Spotted Owl. Abundance and distribution. Common and wide- spread. Habitat. Most areas which support tree growth. Breeding. D'Ombrain (1905a) mentions five nests, two of which had clutches of three, in the Casterton District (near the W boundary of HR) during October and November. Three young birds out of the nest were in GR near Balmoral in late December 1972 (Mitchell 1973; McCulloch 1973). Ninox connivens. Barking Owl. Abundance and distribution. Uncommon; not seen in FWD survey. Recorded in GR at Victoria Val- ley (Braithwaite pers. comm.); in AR near Sta- well (McCann pers. comm.); and in or near HR by D'Ombrain (1905a), Wheeler (1967b) and Middleton (pers. comm.). Habitat. Woodlands and open forests. Breeding. A pair has nested for a number of years in the NW portion of HR (Austin pers. comm.). TYTONIDAE (BARN OWLS) Tyto alba. Barn Owl. Abundance and distribution. Common and wide- spread except in the mountain ranges of GR. Habitat. Woodlands, semi-cleared pastures and settled areas and it roosts in tree hollows, old buildings, caves, thick foliage or down wells (Frith 1969). Breeding. D'Ombrain (1905b) gives details on nesting of this species in the Cas- terton District (near W boundary of HR) around the turn of this century. Tyto novaehollandiae. Masked Owl. Abundance and distribution. Rare; not seen in FWD survey. Recorded in the Casterton District (near W boundary of HR) during the early part of this century (D'Ombrain 1903; 1905a). Wake- field (1963) suggests that an accumulation of bones found in a sandstone recess in GR was from the disgorged pellets of this species. It may be significant that the composition of the mammal remains indicated that this recess had not been used for many years. Habitat. Forested areas (Wheeler 1967a). Breeding. Details of a nest and its contents (one fledgling and a partly intact egg shell) found in the Casterton District in De- cember 1902 are described by D'Ombrain (1903). PODARGIDAE (FROGMOUTHS) Podargus strigoides. Tawny Frogmouth. Abundance and distribution. Common and wide- spread. Habitat. Most areas where large trees are present. Breeding. A common breeder in the Casterton District (near W boundary of HR) where D'Ombrain (1905b) found eight nests be- tween August and November 1903. Sullivan (1929d) found three nests in the Western District in 1928. It breeds regularly in AR near Stawell (McCann pers. comm.). 336 W. B. EMISON, J. W. PORTER, K. C. NORRIS and G. J. APPS AEGOTHELIDAE (OWLET-NIGHTJARS) Aegotheles cristatus. Owlet-nightjar. Abundance and distribution. Uncommon and widespread. Recorded in ER at Mt. Arapiles Block (observed on three occasions since 1967); in GR near Rocklands Reservoir; and in AR near Stawell. Habitat. Open forests and settled areas (Wheeler 1967a). At Mt. Arapiles sightings have been made in Brown Stringybark shrubland and in Long-leaf Box low open forest (p.c.-21). Breeding. Recorded nesting near Stawell (Mc- Cann pers. comm.). CAPRIMULGIDAE (NIGHTJARS) Caprimulgus gattatus. Spotted Nightjar. Abundance and distribution. Rare; a nightjar was seen in GR near Moora Moora Reservoir on 31 October 1974. Our identification was only positive to genus, but previously published ma- terial on the distribution of nightjars suggests that it was a Spotted Nightjar. Also recorded in GR near Billywing (Hall and McKean 1962) and Middleton (pers. comm.) reported a road-killed specimen near Wail (N of ER). Habitat. The bird near Moora Moora Reservoir was in an open forest with a dense understorey. APODIDAE (SWIFTS) Hirundapus caudacutus. Spine-tailed Swift. Abundance and distribution. Uncommon and widespread. Most sightings of this migratory spe- cies in the survey area have been made between December and March although Frith (1969) states that it occurs in Australia between early October and mid-August. Habitat. Most of its time in Australia is spent in the air and it occurs over all types of habitats. Movements. It migrates to Japan and Siberia where it breeds between May and August (Frith 1969). Apus pacificus. Fork-tailed Swift. Abundance and distribution. There are few re- cords from the survey area. Middleton (pers. comm.) records it as occasional, but not as regu- lar as the Spine-tailed Swift. Habitat. As for Spine-tailed Swift (above). Movements. It mi- grates to E Asia where it breeds between May and August. ALCEDINIDAE (KINGFISHERS) Ceyx azureus. Azure Kingfisher. Abundance and distribution. Rare. One recorded in GR at Cherrypool in March 1954 (McGarvie pers. comm.). A specimen was taken in either GR or AR in November-December 1891 (Anon. 1892). Hill (1907b) recorded this species along the Wimmera River (E of AR) and Sullivan (1929d) remarked that he had seen only one in the Western District of Victoria. Habitat. Along rivers, streams and lake margins (Wheeler 1967a). Breeding. Hill (1907b) reported it nest- ing along the Wimmera River in December. Dacelo novaeguineae. Kookaburra. Abundance and distribution. Common and wide- spread. Habitat. Most areas containing trees. Breeding. Adults were feeding young out of the nest in GR in late December 1972 (McCulloch 1973). Halcyon sancta. Sacred Kingfisher. Abundance and distribution. Uncommon with lo- cal concentrations; widespread. Recorded in ER near Apsley and along the Edenhope to Harrow road; in GR near Balmoral and Rocklands Reser- voir; in AR in the Illawarra Block; and in HR near Wannon. Habitat. Woodlands, particularly those with River Red Gums, sometimes near bodies of fresh water. Rarely in open forest. Breeding. Nests with eggs and young, and adults feeding young out of nests were in GR in late December 1972 (McCulloch 1973). Movements. A migrant to S Australia and is absent in winter (Frith 1969). MEROPIDAE (