RECORDS SOUTH AUSTRALIAN MUSEUM VOLUME 35 FART il MUAY! 2002 NEW SPECIES AND NEW RECORDS OF CLOACINA VON LISTOW, 1898 (NEMATODA: STRONGYLOIDEA) PARASITIC INMACROPODID MARSUPIALS FROM PAPUA NEW GUINEA I. BEVERIDGE Summary New species described from macropodid marsupials in Papua New Guinea are : Cloacina cretheis sp. nov. from tree kangaroos, Dendrolagus inustus (type host), D. dorianus, D. goodfellowi, D. matchiei and D. scottae; C. cunctabunda sp. nov. from D. mbaiso; C. eurynome sp. nov. from D. dorianus and D. scottae; C. hecale sp. nov. from D. dorianus; C. theope sp. nov. from D. matschiei and D. dorianus; C. erigone sp. nov., C. hyperaea sp. nov., C. nephele sp. nov., C. polymena sp. nov., C. praxithea sp. nov. and C. procris sp. nov. from the scrub wallaby, Macropus agilis. New host records are : C. caballeroi Mawson, 1977, C. sterope Beveridge & Speare, 1999 and C. syphax Beveridge & Speare, 1999 in Do. Hageni. Additional geographical records are given for C. cornuta in M. agilis. A key to the known species of Cloacina in Papua New Guinea is provided. NEW SPECIES AND NEW RECORDS OF CLOACINA VON LINSTOW, 1898 (NEMATODA: STRONGYLOIDEA) PARASITIC INMACROPODID MARSUPIALS FROM PAPUA NEW GUINEA I. BEVERIDGE BEVERIDGE, I. 2002. New species and new records of Cloacina von Linstow, 1898 (Nematoda: Strongyloidea) parasitic in macropodid marsupials from Papua New Guinea. Records of the South Australian Museum 35(1): 1-32. New species described from macropodid marsupials in Papua New Guinea are : Cloacina cretheis sp. nov. from tree kangaroos, Dendrolagus inustus (type host), D. dorianus, D. goodfellowi, D. matschiei and D. scottae; C. cunctabunda sp. nov. from D. mbaiso; C. eurynome sp. nov. from D. dorianus and D. scottae; C. hecale sp. nov. from D. dorianus; C. theope sp. nov. from D. matschiei and D. dorianus; C. erigone sp. nov., C. hyperaea sp. nov., C. nephele sp. nov., C. polymena sp. nov., C. praxithea sp. nov. and C. procris sp. nov. from the scrub wallaby, Dorcopsis hageni, and C. oweni n. sp. and C. papuensis n. sp. from the agile wallaby, Macropus agilis. New host records are: C. caballeroi Mawson, 1977, C. sterope Beveridge & Speare, 1999 and C. syphax Beveridge & Speare, 1999 in Do. hageni. Additional geographical records are given for C. cornuta in M. agilis. A key to the known species of Cloacina in Papua New Guinea is provided. I. Beveridge, Department of Veterinary Science, University of Melbourne, Parkville, Victoria 3052, Australia. Manuscript received 7 May 2001. Species of the nematode genus Cloacina von Linstow, 1898 occur exclusively in the stomachs of macropodid marsupials. Currently, 103 species are recognised as valid (Beveridge 1998, 1999; Beveridge and Speare 1999) although substantial numbers of species remain to be described (Beveridge 1998). Most of the species described to date are from Australian kangaroos and wallabies, with relatively few records from Papua New Guinea. This situation reflects the paucity of knowledge of the parasite fauna of macropodids from Papua New Guinea, which is based currently on a small number of incidental collections. Nevertheless, material currently available from various kangaroos and wallabies consists of a number of novel species of Cloacina. Thirteen new species are described in the current paper as well as new host and distribution records. The opportunity is taken to provide a summary of species of Cloacina known from Papua New Guinea together with a key to facilitate their identification. Finally, a preliminary comparison is made between the Australian and Papua New Guinean species and their host distributions. Beveridge (1998) noted the occurrence of a number of species of Cloacina in Papua New Guinea, specifically C. australis (Yorke & Maplestone, 1926) in the agile wallaby, Macropus agilis (Gould, 1842); C. caballeroi Mawson, 1977 in the grey scrub wallaby, Dorcopsis luctuosa (D’ Albertis, 1874) and the brown scrub wallaby, D. muelleri (Schlegel, 1866); C cloelia Beveridge, 1998 in the pademelons Thylogale calabyi Flannery, 1992 and T. stigmatica (Gould, 1860); C. cornuta (Davey & Wood, 1938) in M. agilis; C. cybele Beveridge, 1998 in T. stigmatica; and C. dahli von Linstow, 1898 in Thylogale browni Ramsay, 1877, T. calabyi and T. stigmatica. Subsequently, based on an examination of the parasites of four small scrub wallabies, Dorcopsulus vanheurni (Thomas, 1922), Beveridge & Speare (1999) described seven new species, C. sancus, C. sciron, C. sappho, C. solon, C. solymus, C. sterope and C. syphax. Beveridge (1998) noted the presence of undescribed species from the white-striped scrub wallaby, Dorcopsis hageni Heller, 1897 in the collections of the South Australian Museum, Adelaide, and from Do. luctuosa in the collections of The Natural History Museum, London. Flannery et al. (1996) also reported species of the genus Cloacina in a variety of tree kangaroos: Dendrolagus inustus Mueller, 1840; D. goodfellowi Thomas, 1908; D. dorianus Ramsay, 1883; D. scottae Flannery & Seri, 1990; and D. mbaiso Flannery, Boeadi & Szalay, 1995, a group of kangaroos not previously known to harbour Cloacina. These various undescribed collections form the basis of the current report. 2 I. BEVERIDGE MATERIALS AND METHODS Nematodes were examined from collections held in the Australian Helminthological Collection (AHC), South Australian Museum (SAM), Adelaide, the Natural History Museum (BMNH), London and the United States National Parasite Collection (USNPC), Beltsville, Maryland. Nematodes were washed in water and cleared in lactophenol. Drawings were made using a drawing tube attached to an Olympus BH2 microscope. Drawings of apical views of the heads of nematodes are oriented with the dorsal aspect uppermost; drawings of the bursa have the ventral surface uppermost. All drawings are of paratype specimens. Measurements were made with an ocular micrometer. All measurements are in millimetres and are presented as the range followed by the mean in parentheses. Morphological terminology follows Beveridge (1998). Because of the relative uniformity of the genus Cloacina, the descriptions presented are abbreviated and follow the format used by Beveridge (1998), in which a full description of the genus was followed by individual species descriptions concentrating on features of diagnostic significance. A detailed description of the genus is available in Beveridge (1998). Features which are relatively invariable, such as the disposition of ventral and lateral bursal rays, are not included in descriptions but are, nonetheless, illustrated. Types of the new species have been deposited in SAM, BMNH and USNPC. Host nomenclature follows Flannery (1995) and Flannery et al. (1995). Following previous practice (Beveridge 1998), the names of new species are based on those of classical deities since Cloacina was the Roman goddess of the toilet. An invocational quatrain on this subject, usually attributed to Byron (see Lewin 1999, p. 31), occurs in a number of variations, but the rendering by Lewin (1999) is presented here: O Cloacina, goddess of this place, Look on thy suppliants with smiling face, Soft yet cohesive let their offerings flow, Not rashly swift nor insolently slow. SYSTEMATICS Cloacina caballeroi Mawson, 1977 Material examined: 73, 132, from stomach of Dorcopsis hageni, Usino, Madang, Papua New Guinea, coll. T. Reardon, May 1987, SAM AHC 31425. Remarks Cloacina caballeroi was described by Mawson (1977) and Beveridge (1998) from Dorcopsis muelleri and Do. luctuosa from Irian Jaya and Papua New Guinea. This represents the first record from Do. hageni. Cloacina cornuta (Davey & Wood, 1938) Material examined: 23, 12, from stomach of Macropus agilis, Bula Plain, Bensbach, Papua New Guinea, coll. I. Owen, May, 1998, BMNH 1998.9.28.24-26. Remarks Cloacina cornuta was reported from Macropus agilis from Port Moresby and Cape Rodney by Beveridge (1998). The present record adds an additional locality. Data provided by Dr I. Owen indicate that in four wallabies examined, numbers of C. cornuta ranged from 1650-9600 (mean 4700). Although based on a very small number of wallabies, these data contrast strikingly with those of Speare et al. (1983) for M. agilis from northern Australia in which C. cornuta was found in only 41% of hosts examined, and with Beveridge et al. (1998) who found the nematode in 35% of wallabies in Queensland. In Australia, C. cornuta invariably occurs in much lower numbers than the synhospitalic species C. australis (unpublished observations). Cloacina cretheis sp. nov. (Figs 1-13) Types: From stomach of Dendrolagus inustus, Mt Somoro, Sandaun Province, Papua New Guinea, coll. T. Flannery, 10.iii.1990. Holotype 3, SAM AHC 31426; allotype 2, SAM AHC 31427. Paratypes: 12d, 122, SAM AHC 31428; 1d, 12, BMNH 2001.4.10.1-2; 1¢, 12, USNPC 91133. Slide preparations of male, apical views of mouth and bursa, SAM AHC 28378. Material examined: From Dendrolagus inustus: types. From Dendrolagus dorianus: 13, 49, Gunung Ki, Tembagapura, Irian Jaya, coll. T. Flannery, 19.v.1994, SAM AHC 31430; 2d, 10, Lake Trist, Papua New Guinea, coll. I. Redmond, BMNH, 1979, 4486-4505. From Dendrolagus CLOACINA FROM PAPUA NEW GUINEA 3 FIGURES 1-13. Cloacina cretheis sp. nov., specimens from Dendrolagus inustus. 1. Anterior end, lateral view. 2. Cephalic extremity, lateral view, dorsal aspect on right-hand side. 3. Cephalic extremity, dorsal view. 4. Submedian cephalic papilla. 5. Cephalic extremity, apical view. 6. Cephalic extremity, transverse optical section through buccal capsule. 7. Genital cone, ventral view. 8. Genital cone, dorsal view. 9. Gubernaculum, ventral view. 10. Bursa, apical view. 11. Distal tip of spicule, lateral view. 12. Female tail, lateral view. 13. Female genital system, lateral view. Scale bars: 1, 10, 12, 13, 0.1 mm; 2-9, 11, 0.01 mm. 4 I. BEVERIDGE goodfellowi: 5d, 58, Mt Machold, Sandaun Province, Papua New Guinea, coll. T. Flannery and K. Vula, 13.xii.1990, SAM AHC 31431. From Dendrolagus matschiei Forster & Rothschild, 1907: 26¢, 282, Huon Peninsula, Papua New Guinea, coll. J. Mayer, July 1999, USNPC 91146. From Dendrolagus scottae: 93, 52, Sweipini, Sandaun Province, Papua New Guinea, coll. T. Flannery, 15.vi.1991, SAM AHC 31429. Description Small nematodes; cervical cuticle very slightly inflated to beyond level of excretory pore; transverse cuticular annulations widely spaced, 0.030 apart. Submedian cephalic papillae prominent, 0.015 long, projecting anteriorly from peri-oral cuticle; distal segment sub-spherical, 0.005 long, shorter than cylindrical proximal segment, 0.010 long. Buccal capsule shallow, symmetrical in dorsal and lateral views, circular in transverse section, wider than deep, wall without striations. Leaf crown elements 8 in number, slightly recurved at tips; peri-oral cuticle not inflated into lobes attached to each element. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, not projecting into lumen of buccal capsule. Oesophagus slender, claviform, slightly wider at posterior end; lining without sclerotised bosses; denticles absent. Nerve ring in mid-oesophageal region; deirids posterior to Nerve ring; excretory pore at level of oesophago-intestinal junction. Male. Measurements of 5 specimens, types. Total length 6.7—9.0 (7.8); maximum width 0.35— 0.45 (0.40); buccal capsule 0.010 (0.010) x 0.023— 0.025 (0.023); oesophagus 0.46-0.57 (0.52); nerve ring to anterior end 0.20—0.24 (0.22); excretory pore to anterior end 0.39—-0.48 (0.45); deirid to anterior end 0.28—0.36 (0.32); spicules 2.34-2.62 (2.54); gubernaculum 0.03 (0.03) long. Measurements of 5 specimens from D. goodfellowi: total length 7.9-9.3 (8.8); maximum width 0.48-0.56 (0.51); buccal capsule 0.010— 0.012 (0.011) x 0.023-0.025 (0.023); oesophagus 0.53-0.64 (0.59); nerve ring to anterior end 0.23- 0.24 (0.24); excretory pore to anterior end 0.44— 0.53 (0.47); deirid to anterior end 0.20—-0.34 (0.29); spicules 2.53-2.94 (2.74); gubernaculum 0.030—0.040 (0.034) long. Measurements of 3 specimens from D. scottae: total length 6.9-9.7 (8.4); maximum width 0.43— 0.47 (0.45); buccal capsule 0.010 (0.010) x 0.023— 0.025 (0.023); oesophagus 0.48-0.62 (0.56); nerve ring to anterior end 0.20; excretory pore to anterior end 0,40-0.46 (0.42); deirid to anterior end 0.25-0.31 (0.28); spicules 2.63—2.66 (2.65); gubernaculum 0.030—0.035 (0.032) long. Measurements of single specimens from D. dorianus: total length 9.4 ; maximum width 0.43; buccal capsule 0.010 x 0.025 ; oesophagus 0.50 ; excretory pore to anterior end 0.50; deirid to anterior end 0.26; spicules 2.97; gubernaculum 0.035 long. Dorsal ray elongate; external branchlets arise at 1/3 length, before major bifurcation; angle of bifurcation acute; external branchlets as long as internals, directed postero-laterally, not reaching margin of bursa; internal branchlets originate 2/3 along dorsal ray, directed postero- laterally, almost reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, wider than long. Spicule tip blunt; ala terminates anterior to spicule tip; anterior lip of genital cone conical; posterior lip with paired projections and cuticular inflation of internal surface of bursa on either side. Female. Measurements of 5 specimens, types. Total length 7.7—9.0 (8.2); maximum width 0.52- 0.54 (0.52); buccal capsule 0.008—0.010 (0.010) x 0.023-0.025 (0.023); oesophagus 0.48-0.55 (0.53); nerve ring to anterior end 0.20-0.30 (0.23); excretory pore to anterior end 0.39-0.50 (0.44); deirid to anterior end 0.26-0.33 (0.30); tail 0.23-0.31 (0.27); vulva to posterior end 0.35—0.40 (0.38); vagina 0.50-0.73 (0.62); egg 0.075—0.080 (0.080) x 0.040—0.050 (0.045). Measurements of 5 specimens from D. scottae: total length 8.8-10.7 (10.0); maximum width 0.49-0.64 (0.59); buccal capsule 0.008—0.010 (0.009) x 0.020—-0.025 (0.023); oesophagus 0.53— 0.58 (0.55); nerve ring to anterior end 0.20; excretory pore to anterior end 0.48-0.50 (0.49); deirid to anterior end 0.29-0,32 (0.30); tail 0.23- 0.32 (0.28); vulva to posterior end 0.38-0.40 (0.39); vagina 0.53—0.58 (0.56). Measurements of 3 specimens from D. dorianus: total length 8.8—9.2 (9.0); maximum width 0.49-0.72 (0.60); buccal capsule 0.010- 0.013 (0.012) x 0.023-0.025 (0.024); oesophagus 0.53-0.60 (0.57); nerve ring to anterior end 0.23 (0.23); excretory pore to anterior end 0.48-0.51 (0.49); deirid to anterior end 0.20—0.29 (0.24); tail 0.21-0.25 (0.23); vulva to posterior end 0.49-0.50 (0.49); vagina 0.64-0.67 (0.66); egg 0.070—0.075 (0.078) x 0.040—-0.050 (0.045). Female tail short, conical; vagina elongate, convoluted; egg ellipsoidal. CLOACINA FROM PAPUA NEW GUINEA 5 Remarks C. cretheis sp. nov. is characterised by submedian papillae with a small sub-spherical distal segment, a simple buccal capsule, eight elements to the leaf crown, a slender, unornamented oesophagus, deirids posterior to the nerve ring, a dorsal ray in which the external branchlets arise before the major bifurcation and an elongate, convoluted vagina. The shape of the dorsal ray alone separates it from all congeners except C. caballeroi, C. enyo Beveridge, 1998, C. ips Beveridge, 1998, and C. syphax. It differs from all of these except C. caballeroi in having the deirid posterior to the nerve ring, although in some highly contracted specimens of C. cretheis, in which the anterior oesophagus becomes sinuous, the deirid may appear more anteriorly on the contracted cuticle. C. cretheis is further distinguished from C. syphax which has a sinuous anterior margin to its buccal capsule and submedian cephalic papillae with enlarged distal segments. C. cretheis is distinguished from C. caballeroi by spicule lengths, which are 2.34—2.97 mm in C. cretheis compared with 1.23-1.45 mm in C. caballeroi. The spicules of C. enyo are 0.68—0.88 mm long while those of C. ips are 1.36-1.39 mm in length, providing a further means of distinguishing the latter species. C. cretheis occurs in a wide range of tree kangaroo species in Papua New Guinea and was by far the most frequently encountered species in tree kangaroos in this study. It does not occur in either Dendrolagus bennettianus DeVis, 1887 or D. lumholtzi Collett, 1884 in Australia (Spratt et al. 1991). Although incomplete, the measurements of specimens from different hosts suggest that there are no major differences induced by the host species. Some of the minor differences observed are due to the state of contraction or of preservation of specimens, and the incomplete sets of measurements reflect the poor state of preservation of many of the specimens, precluding the measurement of internal organs. Cloacina cunctabunda sp. nov. (Figs 14-23) Types: From stomach of Dendrolagus mbaiso, Camp Ridge, Tembagapura, Irian Jaya, coll. T. Flannery, 24.v.1994. Holotype ¢, SAM AHC 31432; allotype 2, SAM AHC 31433. Paratype 3, 3 slides, SAM AHC 28379. Description Small nematodes; cervical cuticle slightly inflated to beyond level of excretory pore; transverse cuticular annulations widely spaced, 0.015 apart. Submedian cephalic papillae prominent, 0.010 long, projecting anteriorly from peri-oral cuticle; distal segment ovoid, 0.006 long, longer than cylindrical proximal segment, 0.004 long. Buccal capsule shallow, symmetrical in dorsal and lateral views, oval in transverse section, slightly deeper dorso-ventrally, wider than deep, wall without striations. Leaf crown elements 6 in number, recurved at tips; peri-oral cuticle, inflated; inflations not attached to each element of leaf crown. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, not projecting into lumen of buccal capsule. Oesophagus slender, claviform, slightly wider at posterior end; lining without sclerotised bosses; denticles absent. Nerve ring in mid-oesophageal region; deirids anterior to nerve ring, mid-way between nerve ring and buccal capsule; excretory pore anterior to level of oesophago-intestinal junction. Male. Measurements of holotype and paratype. Total length 3.30, 3.32; maximum width 0.16, 0.17; buccal capsule 0.008, 0.010 x 0.025, 0.025; oesophagus 0.34, 0.35; nerve ring to anterior end 0.16, 0.18; excretory pore to anterior end 0.26, 0.30; deirid to anterior end 0.11, 0.14; spicules 1.39, 1.57; gubernaculum 0.015, 0.015 long. Dorsal lobe of bursa and dorsal ray elongate, ray broad at origin; external branchlets arise at 1/2 length, immediately before major bifurcation; angle of bifurcation acute; external branchlets shorter than internals, directed postero-laterally, not reaching margin of bursa; internal branchlets originate soon after externals arise from dorsal ray, directed postero-laterally, almost reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, wider than long. Spicule tip not everted. Anterior lip of genital cone conical; posterior lip with paired projections. Female. Measurements of allotype. Total length 4.25; maximum width 0.21; buccal capsule 0.010 x 0.025; oesophagus 0.40; nerve ring to anterior end 0.18; excretory pore to anterior end 0.29; deirid to anterior end 0.11; tail 0.35; vulva to posterior end 0.55; vagina 0.35. Female tail elongate, conical; vulva immediately anterior to anus; vagina short with single convolution; egg not seen. Remarks Although described from only three specimens, 6 I. BEVERIDGE FIGURES 14-23. Cloacina cunctabunda sp. nov., specimens from Dendrolagus mbaiso. 14. Anterior end, lateral view. 15. Cephalic extremity, lateral view, dorsal aspect on left-hand side. 16. Cephalic extremity, ventral view. 17. Submedian cephalic papilla, lateral view. 18. Cephalic extremity, apical view. 19. Cephalic extremity, transverse optical section through buccal capsule. 20. Bursa, apical view. 21. Gubernaculum, ventral view. 22. Female tail, lateral view. 23. Female genital system, lateral view. Scale bars: 22, 23, 0.1 mm; 14-21 0.01 mm. CLOACINA FROM PAPUA NEW GUINEA 7 this species is sufficiently distinctive to allow ready recognition, and comes from a host restricted to isolated localities in Irian Jaya which is unlikely to be sampled extensively in the future (Flannery et al. 1995). Furthermore, this species of Cloacina was not found in any other species of tree kangaroo. C. cunctabunda is distinguished from all congeners except C. cretheis, C. caballeroi, C. enyo, C. ips and C. syphax in possessing a simple buccal capsule, an elongate, unornamented oesophagus and the external branchlets of the dorsal ray arising before the major bifurcation. The position of the deirid anterior to the nerve ring distinguishes the species from C. cretheis and C. caballeroi, while the shape of the buccal capsule and the submedian cephalic papillae distinguish it from C. syphax. C. cunctabunda is distinguished from C. enyo on the basis of spicule lengths, which are 1.39-1.57 mm long in C. cunctabunda compared with 0.68-0.88 mm in C. enyo. C. cunctabunda therefore most closely resembles C. ips, from which it is distinguished by the number of leaf crown elements (six in C. cunctabunda, eight in C. ips), in the absence of a dorsal oesophageal tooth which projects prominently into the buccal capsule of C. ips, in the shape of the female tail which is slender and elongate in C. cunctabunda but short and conical in C. ips, and in the shape of the vagina which is Straight in C. ips but convoluted in C. cunctabunda. Cloacina erigone sp. nov. (Figs 24-36) Types: From stomach of Dorcopsis hageni, Usino, Madang, Papua New Guinea, coll. T. Reardon, May 1987. Holotype 6, SAM AHC 31434; allotype 2, SAM AHC 31435. Paratypes: 43, 52, SAM AHC 31436, 16, 12, BMNH 2001.4.10.3-4, 1d, 12, USNPC 91134. Slide preparations of apical views of mouth and bursa, SAM AHC 28380. Material examined: types. Description Small nematodes; cervical cuticle slightly inflated to beyond level of excretory pore; transverse cuticular annulations widely spaced, 0.015 apart. Submedian cephalic papillae prominent, 0.013 long, projecting anteriorly from peri-oral cuticle; distal segment ovoid, 0.005 long, shorter than cylindrical proximal segment, 0.008 long. Buccal capsule shallow, symmetrical in dorsal and lateral views, circular in transverse section, wider than deep, wall without striations. Leaf crown elements 8 in number, slightly recurved at tips, margins prominently thickened; peri-oral cuticle not inflated into lobes attached to each element. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, projecting prominently into lumen of buccal capsule. Oesophagus claviform, robust, slightly wider at posterior end; lining with sclerotised bosses extending from anterior end, halfway to nerve ring; single dorsal denticle present in prominent preneural swelling of oesophagus. Nerve ring in mid-oesophageal region; deirids posterior to nerve ring, almost at level of excretory pore; excretory pore at level of oesophago-intestinal junction. Male. Measurements of 8 specimens, types. Total length 2.88-5.11 (4.02); maximum width 0.18—0.31 (0.26); buccal capsule 0.007—0.010 (0.008) x 0.023—0.035 (0.032); oesophagus 0.26— 0.39 (0.34); nerve ring to anterior end 0.14—0.22 (0.18); excretory pore to anterior end 0.25—0.40 (0.33); deirid to anterior end 0.23-0.37 (0.32); spicules 1.35-1.79 (1.59); gubernaculum 0.020— 0.030 (0.024) long. Dorsal ray elongate, broad at origin; external branchlets arise at 1/2 length, immediately before major bifurcation; angle of bifurcation acute; external branchlets shorter than internals, directed laterally, not reaching margin of bursa; internal branchlets originate immediately after externals, directed postero-laterally, almost reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, slightly wider than long. Spicule tip blunt; ala terminates abruptly anterior to spicule tip; anterior lip of genital cone conical; posterior lip with paired projections and cuticular inflation of internal surface of bursa on either side. Female. Measurements of 8 specimens, types. Total length 3.56-4.81 (4.39); maximum width 0.31-0.48 (0.41); buccal capsule 0.008-0.010 (0.009) x 0.033—-0.045 (0.039); oesophagus 0.35— 0.44 (0.39); nerve ring to anterior end 0.16—0.23 (0.20); excretory pore to anterior end 0.21-0.43 (0.34); deirid to anterior end 0.26—-0.37 (0.32); tail 0.16-0.19 (0.17); vulva to posterior end 0.24—0.28 (0.26); vagina 0.53-0.99 (0.75); egg 0.055—0.070 (0.062) x 0.030—0.035 (0.031). Female tail short, conical; vulva immediately anterior to anus; vagina elongate, straight; egg ellipsoidal. 8 I. BEVERIDGE FIGURES 24-36. Cloacina erigone sp. nov., types. 24. Anterior end, lateral view. 25. Cephalic extremity, lateral view, dorsal aspect on left-hand side. 26. Cephalic extremity, dorsal view. 27. Cephalic extremity, apical view. 28. Cephalic extremity, transverse optical section through buccal capsule. 29. Submedian cephalic papilla, lateral view. 30. Preneural oesophageal swelling showing dorsal denticle, dorsal view. 31. Bursa, apical view. 32. Genital cone, dorsal view. 33. Spicule tip, lateral view. 34. Gubernaculum, ventral view. 35. Female tail, lateral view. 36. Female genital system, lateral view. Scale bars: 24, 31, 35, 36, 0.1 mm; 25-30, 32-34, 0.01 mm. CLOACINA FROM PAPUA NEW GUINEA 9 Remarks Cloacina erigone sp. nov. is characterised by having submedian cephalic papillae with an elongate proximal segment, a_ shallow, unornamented buccal capsule, eight leaf crown elements, deirid posterior to the nerve ring, a dorsal ray with the external branchlets arising before the main bifurcation and a straight vagina. In these characters, C. erigone resembles C. caballeroi, C. cretheis, C. enyo, C. ips and C. syphax. It differs from all of these species, however, in having bosses lining the anterior part of the vagina and a single dorsal denticle. Species with a symmetrical buccal capsule, oesophageal bosses and a single dorsal denticle are C. australis, C. dis Beveridge, 1998, C. hecuba Beveridge, 1998, C. io Beveridge, 1998, C. laius, C. leto Beveridge, 1998, C. minor (Davey & Wood, 1938) and C. tyro Beveridge, 1998. However, in none of these species is the deirid posterior to the nerve ring, and in none do the external branchlets of the dorsal ray arise before the principal bifurcation. This species has, thus far, been found only in Do. hageni. Cloacina eurynome sp. nov. (Figs 37-49) Types: From stomach of Dendrolagus dorianus, Tembagapura, Irian Jaya, coll. T. Flannery, 19.v.1994, 23.v.1994. Holotype 6, SAM AHC 31437; allotype 2, SAM AHC 31438. Paratypes: 43,92, SAM AHC 31439, 1d, 22, SAM AHC 31440, 1d, 12, BMNH 2001.4.10.5-6, 1d, 12, USNPC 91135. Slide preparations of spicules, apical views of mouth and bursa, SAM AHC 28381. Material examined: From Dendrolagus dorianus: types. From Dendrolagus scottae: 4°, Sweipini, Sandaun Province, Papua New Guinea, coll. T. Flannery, 15.vi.1991, SAM AHC 31441. Description Small nematodes; cervical cuticle slightly inflated to beyond level of excretory pore; transverse cuticular annulations widely spaced, 0.030 apart. Submedian cephalic papillae prominent, 0.018 long, projecting anteriorly from peri-oral cuticle; distal segment ovoid, pointed apically, 0.008 long, shorter than cylindrical proximal segment, 0.010 long. Buccal capsule shallow, symmetrical in dorsal and lateral views, approximately octagonal in transverse section, wider than deep, wall without striations. Leaf crown elements 8 in number, slightly recurved at tips; peri-oral cuticle not inflated into lobes attached to each element. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, not projecting into lumen of buccal capsule. Oesophagus claviform, slender, only slightly wider at posterior end; lining without sclerotised bosses; three poorly developed denticles, one dorsal and two subventral, present in lumen of oesophagus immediately anterior to nerve ring; preneural swelling of oesophagus small. Nerve ring in mid- oesophageal region; deirids at level of nerve ring; excretory pore at level of oesophago- intestinal junction. Male. Measurements of 7 specimens, types. Total length 8.75—-10.67 (9.53); maximum width 0.39-0.48 (0.45); buccal capsule 0.007—0.010 (0.008) x 0.027—-0.030 (0.029); oesophagus 0.57— 0.64 (0.59); nerve ring to anterior end 0.26—0.28 (0.27); excretory pore to anterior end 0.54—0.62 (0.58); deirid to anterior end 0.24—0.32 (0.28); spicules 3.57—3.98 (3.77); gubernaculum 0.015— 0.020 (0.019) long. Dorsal ray elongate, broad at origin; major bifurcation occurs at 1/3 length; angle of bifurcation acute; external branchlets arise mid- way between major bifurcation and tip, much shorter than internals, directed laterally or postero- laterally, not reaching margin of bursa; internal branchlets directed postero-laterally, almost reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, wider than long. Spicule tip blunt, gently curved; ala terminates abruptly anterior to spicule tip; anterior lip of genital cone conical; posterior lip with paired projections and cuticular inflation of internal surface of bursa on either side. Female. Measurements of 10 specimens, types. Total length 10.19-13.28 (11.50); maximum width 0.52—0.75 (0.60); buccal capsule 0.005— 0.013 (0.008) x 0.030—0.035 (0.031); oesophagus 0.61-0.73 (0.65); nerve ring to anterior end 0.25— 0.28 (0.27); excretory pore to anterior end 0.52— 0.71 (0.60); deirid to anterior end 0.18—0.34 (0.27); tail 0.22-0.29 (0.25); vulva to posterior end 0.31-0.46 (0.39); vagina 1.57-1.85 (1.75); egg 0.080—0.095 (0.086) x 0.040—0.050 (0.046). Female tail slender, conical; vulva immediately anterior to anus; vagina elongate, straight distally, recurrent anterior to vestibule, recurrent section twisted in characteristic figure of eight formation; egg ellipsoidal. 10 I. BEVERIDGE FIGURES 37-49. Cloacina eurynome sp. nov., types. 37. Anterior end, lateral view. 38. Cephalic extremity, lateral view, dorsal aspect on right-hand side. 39. Cephalic extremity, ventral view. 40. Submedian cephalic papilla, lateral view. 41. Cephalic extremity, apical view. 42. Cephalic extremity, transverse optical section through buccal capsule. 43. Preneural oesophageal swelling showing denticles, lateral view, dorsal aspect on right-hand side. 44. Preneural oesophageal swelling showing denticles, ventral view. 45. Bursa, apical view. 46. Spicule tip, lateral view. 47. Gubernaculum, genital cone and thickenings of spicule sheaths, dorsal view. 48. Female tail, lateral view. 49. Female genital system, lateral view. Scale bars: 37, 45, 48, 49, 0.1 mm; 38-44, 46, 47, 0.01 mm. CLOACINA FROM PAPUA NEW GUINEA 11 Remarks The presence of three preneural denticles at the same level in the oesophagus distinguishes C. eurynome from all congeners except C. hera Beveridge, 1998, C. hermes Beveridge, 1998 and C. hestia Beveridge, 1998, all of which are parasites of grey kangaroos, Macropus fuliginosus (Desmarest, 1817) and M. giganteus Shaw, 1790 in eastern and southern Australia (Beveridge 1998) and C. daveyi Mawson, 1977 in the wallaroo, M. robustus Gould, 1841. C. eurynome is distinguished from the species in grey kangaroos in having the deirid at the level of the nerve ring rather than well anterior to it and in having eight rather than six leaf crown elements. It is distinguished from C. daveyi which has cephalic papillae with large, globose, medially directed distal segments. The denticles in C. eurynome are vestigial, a feature found only in C. hestia. C. eurynome is distinguished from all congeners other than C. syphax, from Dorcopsulus vanheurni, also from Papua New Guinea, in the morphology of the vagina, which exhibits a highly characteristic recurrent loop, anterior to the vestibule, twisted in a figure of eight formation. C. eurynome differs from C. Syphax in possessing denticles, in having a straight rather than a sinuous anterior margin to the buccal capsule, and in having the external branchlets of the dorsal ray arise after the major bifurcation Cloacina hecale sp. nov. (Figs 50-61) Types: From stomach of Dendrolagus dorianus, Lake Trist, Papua New Guinea, coll. I. Redmond, 1979. Holotype 36, BMNH 1981.4506; allotype 2, BMNH 1981.4507. Paratypes: 49d, 882, BMNH 1981.4508-4535. Slide preparations of apical views of bursa and mouth, SAM AHC 28383. Material examined: From Dendrolagus dorianus: types. Description Robust nematodes; cervical cuticle slightly inflated to beyond level of excretory pore; transverse cuticular annulations widely spaced, 0.023 apart. Submedian cephalic papillae prominent, 0.017 long, projecting anteriorly from peri-oral cuticle; distal segment conical, 0.007 long, only slightly shorter than cylindrical proximal segment, 0.010 long. Buccal capsule shallow, symmetrical in dorsal and lateral views, circular in transverse section, wider than deep, wall without striations; anterior margin slightly undulant. Leaf crown elements 8 in number, slightly recurved at tips; peri-oral cuticle not inflated into lobes attached to each element. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, not projecting into lumen of buccal capsule. Oesophagus claviform, slender, slightly wider at posterior end; lining without sclerotised bosses; denticles absent. Nerve ring in anterior oesophageal region; deirids anterior to nerve ring; excretory pore at level of oesophago-intestinal junction. Male. Measurements of 10 specimens, types. Total length 9.44-13.86 (11.82); maximum width 0.50-0.67 (0.60); buccal capsule 0.010-0.013 (0.011) x 0.035—0.040 (0.039); oesophagus 0.76— 0.95 (0.87); nerve ring to anterior end 0.30—-0.32 (0.31); excretory pore to anterior end 0.52-0.98 (0.80); deirid to anterior end 0.17-0.26 (0.23); spicules 3.90-5.05 (4.54); gubernaculum 0.040 long. Dorsal ray broad at origin; major bifurcation occurs at 1/2 length; external branchlets arise immediately after major bifurcation, as long as internals, though more robust, directed postero- laterally, not reaching margin of bursa; internal branchlets directed posteriorly, almost reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, wider than long. Spicule minutely bifid at tip; ala terminates abruptly anterior to spicule tip; anterior lip of genital cone conical; posterior lip with paired projections and cuticular inflation of internal surface of bursa on either side. Female. Measurements of 5 specimens, types. Total length 19.6-22.9 (21.3); maximum width 0.64-0.91 (0.81); buccal capsule 0.010 (0.010) x 0.040 (0.040); oesophagus 0.95-1.02 (0.97); nerve ring to anterior end 0.25-0.29 (0.27); excretory pore to anterior end 0.63—0.86 (0.70); deirid to anterior end 0.19-0.29 (0.22); tail 0.13- 0.30 (0.21); vulva to posterior end 0.18-0.43 (0.28); vagina 2.22—2.57 (2.40); vestibule 0.27; sphincter 0.22; infundibulum 0.17; egg 0.090— 0.110 (0.096) x 0.045-0.050 (0.048). Female tail short, conical; vulva immediately anterior to anus; vagina elongate, straight, extends slightly anterior to vestibule with short recurrent section; egg ellipsoidal. Remarks Cloacina hecale sp. nov. is a robust species characterised by long spicules and an elongate 12 I. BEVERIDGE 56 FIGURES 50-61. Cloacina hecale sp. nov., types. 50. Anterior end, lateral view. 51. Cephalic extremity, lateral view, dorsal aspect on right-hand side. 52. Cephalic extremity, dorsal view. 53. Submedian cephalic papilla, lateral view. 54. Cephalic extremity, apical view. 55. Cephalic extremity, transverse optical section through buccal capsule. 56. Bursa, apical view. 57. Spicule tip, lateral view. 58. Gubernaculum, ventral view. 59. Genital cone, dorsal view. 60. Female tail, lateral view. 61. Female genital system, lateral view. Scale bars: 50, 56, 60, 61, 0.1 mm; 51-55, 57-59, 0.01 mm. CLOACINA FROM PAPUA NEW GUINEA 13 vagina, slightly recurrent at its anterior extremity. The length of the vagina relative to the sizes of vestibule, sphincter and infundibulum are given in the description in this case to emphasise the extreme length of the vagina. The other features of the species are unremarkable, with cephalic papillae bearing a distal segment with an acute tip, almost equal in length to the proximal segment, eight leaf crown elements, a symmetrical buccal capsule, a_ slender, unornamented oesophagus, the deirid anterior to the nerve ring and the external branchlets of the dorsal ray arising after the principal bifurcation. These features together with the length of the spicules (> 3.0 mm) differentiate C. hecale from congeners except C. clymene Beveridge, 1998, C. curta Johnston & Mawson, 1938, C. liebigi Johnston & Mawson, 1938, C. longispiculata Johnston & Mawson, 1939, C. nike Beveridge, 1998, C. robertsi Johnston & Mawson, 1939, C. smalesae Mawson, 1975 and C. solymus. C. hecale is differentiated from C. clymene and C. robertsi which have six elements to the leaf crown and lip-like inflations of the peri-oral cuticle attached to each element of the leaf crown. C. curta, C. liebigi, C. longispiculata and C. smalesae all have six leaf crown elements rather than eight and the vagina is prominently recurrent in these species rather than having a short anterior recurrent section. In addition, C. liebigi, C. longispiculata and C. smalesae have the cervical cuticle inflated so as to form ‘shoulders’ in the oesophageal region. C. nike has submedian cephalic papillae in which the distal segment is much shorter than the proximal segment and has a sinuous vagina and a longer, slender tail in the female. C. solymus has a robust subcylindrical oesophagus but the submedian cephalic papillae are tiny and the anterior margin of the buccal capsule is undulate. Therefore, C. hecale is readily distinguishable from all known congeners. Cloacina hyperea sp. nov. (Figs 62-74) Types: From stomach of Dorcopsis hageni, Usino, Madang, Papua New Guinea, coll. T. Reardon, May, 1987. Holotype 6, SAM AHC 31442; allotype 2, SAM AHC 31443. Paratypes: 12d, 82, SAM AHC 31444, 1¢, 12, BMNH 2001.4.10.7-8, 1d, 12, USNPC 91136. Slide preparations of apical views of mouth and bursa, SAM AHC 28382. Material examined: From Dorcopsis hageni: types. Description Robust neniatodes; cervical cuticle slightly inflated to beyond level of excretory pore; transverse cuticular annulations widely spaced, 0.034 apart. Submedian cephalic papillae prominent, 0.015 long, projecting anteriorly from peri-oral cuticle; distal segment conical, 0.007 long, only slightly shorter than cylindrical proximal segment, 0.008 long. Buccal capsule shallow, symmetrically arched anteriorly in lateral views; in dorsal views, buccal capsule arches anteriorly over dorsal oesophageal gland; in ventral views, curves posteriorly; oval and dorso- ventrally elongate in transverse section, wider than deep, wall with prominent striations; anterior margin smooth, except on dorsal aspect. Leaf crown elements 6 in number, slightly recurved at tips; peri-oral cuticle slightly inflated into lobes attached to each element. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, projecting prominently into lumen of buccal capsule. Oesophagus claviform, robust; lining with sclerotised bosses extending to level of nerve ring; single dorsal oesophageal denticle present in prominent preneural swelling. Nerve ring in mid-oesophageal region; deirids at level of nerve ring; excretory pore at level of, or slightly anterior to, oesophago-intestinal junction. Male. Measurements of 10 specimens, types. Total length 4.20—-5.21 (4.56); maximum width 0.26-0.40 (0.34); buccal capsule 0.013-0.020 (0.016) x 0.055—0.060 (0.056); oesophagus 0.42— 0.50 (0.47); nerve ring to anterior end 0.20-0.24 (0.22); excretory pore to anterior end 0.35-0.51 (0.43); deirid to anterior end 0.25—0.34 (0.31); spicules 2.25-2.49 (2.37); gubernaculum 0.020— 0.030 (0.026) long. Dorsal ray broad at origin; major bifurcation occurs at 1/2 length; angle of bifurcation acute; external branchlets arise after major bifurcation, as long as internals, directed postero-laterally, not reaching margin of bursa; internal branchlets directed posteriorly, not reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, slightly wider than long. Spicule tip blunt; ala diminishes gradually in width towards spicule tip; anterior lip of genital cone conical; posterior lip with paired projections and cuticular inflation of internal surface of bursa on either side. Female. Measurements of 6 specimens, types. Total length 5.25-6.14 (5.67); maximum width 14 I. BEVERIDGE SosO O08 oscars 2000090000 . “aN | | FIGURES 62-74. Cloacina hyperea sp. nov., types. 62. Anterior end, lateral view. 63. Cephalic extremity, lateral view, dorsal aspect on right-hand side. 64. Cephalic extremity, dorsal view. 65. Cephalic extremity, ventral view. 66. Submedian cephalic papilla, lateral view. 67. Cephalic extremity, apical view. 68. Cephalic extremity, transverse optical section through buccal capsule. 69. Preneural swelling of oesophagus, dorsal view, showing denticle. 70. Bursa, apical view. 71. Spicule tip, lateral view. 72. Gubernaculum and genital cone, dorsal view. 73. Female tail, lateral view. 74. Female genital system, lateral view. Scale bars: 62, 70, 73, 74, 0.1 mm; 63-69, 71,72, 0.01 mm. CLOACINA FROM PAPUA NEW GUINEA 15 0.32-0.45 (0.39); buccal capsule 0.013-0.018 (0.016) x 0.058—-0.060 (0.059); oesophagus 0.46— 0.52 (0.49); nerve ring to anterior end 0.20-0.23 (0.22); excretory pore to anterior end 0.35—0.50 (0.45); deirid to anterior end 0.24—0.33 (0.29); tail 0.19—-0.26 (0.23); vulva to posterior end 0.36—0.43 (0.39); vagina 0.96—-1.24 (1.07). Female tail slender, conical; vulva immediately anterior to anus; vagina elongate, sinuous, extends slightly anterior to vestibule with short recurrent section; egg not seen. Remarks The anteriorly arched buccal capsule wall distinguishes C. hyperea sp. nov. from all congeners except C. circe Beveridge, 1999 and C. laius Beveridge, 1999, both of which occur in the quokka, Setonix brachyurus (Quoy & Gaimard, 1830) in the south-west of Western Australia, and from C. nephele sp. nov., described below. C. hyperea is distinguished from C. circe in possessing sclerotised bosses and a dorsal denticle in the oesophagus. It is differentiated from C. laius in having the buccal capsule less prominently arched, in having submedian cephalic papillae of a different shape, with both segments of approximately equal length, whereas in C. laius the proximal segment is almost twice as long as the distal segment. In addition, the spicules are 2.25—2.49 mm long in C. hyperea compared with 1.50-1.97 mm in C. laius and the vagina is commensurately longer, being 0.96-1.24 mm in C. hyperea compared with 0.71-0.92 mm in C. laius. In C. hyperea the internal and external branchlets of the dorsal ray are of approximately equal length whereas in C. laius the external branchlets are much shorter than the internal branchlets. C. nephele sp. nov. has lateral lips, which are absent in C. hyperaea, and a bulbous proximal segment to the cephalic papilla. Cloacina nephele sp. nov. (Figs 75-89) Types: From stomach of Dorcopsis hageni, Usino, Madang, Papua New Guinea, coll. T. Reardon, May, 1987. Holotype d6, SAM AHC 31445; allotype 2, SAM AHC 31446. Paratypes: 3d, 22, SAM AHC 31447, 16, BMNH 2001.4.10.9, 16, USNPC 91137. Slide preparations of apical views of mouth and bursa, SAM AHC 28384. Material examined: From Dorcopsis hageni: types. Description Robust nematodes; cervical cuticle prominently inflated to beyond level of excretory pore; transverse cuticular annulations widely spaced, 0.040 apart. Submedian cephalic papillae prominent, 0.020 long, projecting anteriorly from inflated peri-oral cuticle; distal segment slender, conical, 0.010 long, as long as robust, asymmetrical proximal segment, 0.010 long. Buccal capsule shallow, symmetrically arched anteriorly in lateral views; dorsally, arches anteriorly over dorsal oesophageal gland; ventrally, curves posteriorly; oval and dorso- ventrally elongate in transverse section, wider than deep, wall with prominent striations; anterior margin smooth. Leaf crown elements 6 in number, recurved at tips; peri-oral cuticle inflated into lip- like lobes attached to each element; extra dorsal and ventral projections of peri-oral cuticle present, separate from lateral arcades of amphids and submedian papillae, giving the appearance of lips. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, not projecting into lumen of buccal capsule. Oesophagus claviform, robust; lining without sclerotised bosses; denticles absent in preneural region. Nerve ring in mid- oesophageal region; deirids at level of nerve ring; excretory pore between nerve ring and oesophago- intestinal junction. Male. Measurements of 5 specimens, types. Total length 6.08—7.40 (6.66); maximum width 0.33-0.37 (0.35); buccal capsule 0.023-0.030 (0.026) x 0.090 (0.090); oesophagus 0.69-0.73 (0.71); nerve ring to anterior end 0.28—0.30 (0.29); excretory pore to anterior end 0.48-0.53 (0.51); deirid to anterior end 0.31-0.36 (0.34); spicules 1.96-2.10 (2.03); gubernaculum 0.035— 0.040 (0.039) long. Dorsal ray broad at origin, narrowing posteriorly; major bifurcation occurs at 1/2 length; angle of bifurcation acute; external branchlets arise after major bifurcation, shorter than internals, directed postero-laterally, not reaching margin of bursa; internal branchlets directed posteriorly, almost reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, slightly wider than long. Spicule tip blunt; ala diminishes gradually in width towards spicule tip; anterior lip of genital cone conical; posterior lip with paired projections and cuticular inflation of internal surface of bursa on either side. Female. Measurements of 3 specimens, types. Total length 5.94-7.87 (7.02); maximum width 0.34—0.63 (0.50); buccal capsule 0.025—0.030 16 I. BEVERIDGE FIGURES 75-89. Cloacina nephele sp. nov., types. 75. Anterior end, lateral view. 76. Cephalic extremity, lateral view, dorsal aspect on right-hand side. 77. Cephalic extremity, lateral view, dorsal aspect on right-hand side, median optical section showing leaf crown elements and inflation of cephalic collar. 78. Cephalic extremity, dorsal view. 79. Cephalic extremity, ventral view. 80. Cephalic extremity, ventral view, median optical section showing leaf crown elements. 81. Submedian cephalic papilla, lateral view. 82. Cephalic extremity, apical view. 83. Cephalic extremity, transverse optical section through buccal capsule. 84. Bursa, apical view. 85. Genital cone, dorsal view. 86. Gubernaculum, ventral view. 87. Spicule tip, lateral view. 88. Female tail, lateral view. 89. Female genital system, lateral view. Scale bars: 75, 84, 88, 89, 0.1 mm; 76-83, 85-87, 0.01 mm. CLOACINA FROM PAPUA NEW GUINEA 17 (0.027) x 0.100 (0.100); oesophagus 0.72-0.79 (0.76); nerve ring to anterior end 0,29-0.32 (0.30); excretory pore to anterior end 0.46-0.58 (0.52); deirid to anterior end 0.30-0.31 (0.31); tail 0.31-0.39 (0.35); vulva to posterior end 0.43-0.55 (0.48); vagina 0.51-0.66 (0.59); egg 0.080—0.085 (0.083) x 0.040-0.045 (0.043). Female tail slender, conical; vulva immediately anterior to anus; vagina elongate, sinuous, extends slightly anterior to vestibule with short recurrent section; egg ellipsoidal. Remarks C. nephele sp. nov. closely resembles C. circe, C. laius and C. hyperea in having an anteriorly arched buccal capsule. It differs from C. laius and C. hyperea in lacking oesophageal bosses and denticles, and differs from C. circe in having a markedly inflated cervical cuticle, cephalic papillae in which the distal segment is not oriented medially, and a recurrent vagina. It differs from all of these species in having a swollen cephalic collar and lip-like inflations of the peri-oral cuticle attached to each leaf crown element. The dorsal and ventral ‘lips’ are unique within the genus. Cloacina oweni sp. nov. (Figs 90-101) Synonyms: Cloacina sp. nov. of Beveridge, 1998, p. 506 (Macropus agilis). Types: From stomach of Macropus agilis, Bula Plain, Bensbach, Papua New Guinea, coll. I. Owen, May, 1998. Holotype ¢, SAM AHC 31448; allotype 9, SAM AHC 31449. Paratypes: 51d, 552, SAM AHC 31450, 1d, 12, BMNH 1998.9.28.11-12. Material examined: From Macropus agilis: types; 1d, 19, Dari, Bensbach, Papua New Guinea, SAM AHC 11719. Description Robust nematodes; cervical cuticle slightly inflated to level of excretory pore; transverse cuticular annulations widely spaced, 0.030 apart. Submedian cephalic papillae prominent, 0.013 long, projecting anteriorly from inflated peri-oral cuticle; distal segment ovoid, 0.004 long, shorter than cylindrical proximal segment, 0.008 long. Buccal capsule shallow, symmetrical in lateral and dorso-ventral views; octagonal in transverse section, wider than deep, wall without prominent Striations; anterior margin smooth. Leaf crown elements 8 in number, recurved at tips; peri-oral cuticle inflated into lip-like lobes attached to each element. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, not projecting into lumen of buccal capsule. Oesophagus claviform, robust; lining without sclerotised bosses; single dorsal denticle present at level of nerve ring; preneural swelling absent. Nerve ring in mid-oesophageal region; deirids in anterior oesophageal region; excretory pore between nerve ring and oesophago-intestinal junction. Male. Measurements of 10 specimens, types. Total length 5.48-7.50 (6.73); maximum width 0.26-0.43 (0.34); buccal capsule 0.015-0.025 (0.019) x 0.045-0.070 (0.056); oesophagus 0.53- 0.65 (0.60); nerve ring to anterior end 0.25—0.32 (0.27); excretory pore to anterior end 0.26-0.53 (0.44); deirid to anterior end 0.10—0.16 (0.12); spicules 2.19-2.67 (2.46); gubernaculum 0.025— 0.040 (0.031) long. Dorsal ray broad at origin; major bifurcation occurs at 1/3 length; angle of bifurcation obtuse; external branchlets arise after major bifurcation, near extremity of ray, slightly longer and more robust than internals, directed postero-laterally, not reaching margin of bursa; internal branchlets directed postero-laterally, not reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, slightly wider than long. Spicule tip blunt, slightly recurved; ala diminishes gradually in width towards spicule tip; anterior lip of genital cone conical; posterior lip with paired projections and cuticular inflation of internal surface of bursa on either side. Female. Measurements of 10 specimens, types. Total length 6.70—9.36 (8.06); maximum width 0.30-0.61 (0.46); buccal capsule 0.015—0.025 (0.020) x 0.045-0.075 (0.057); oesophagus 0.55— 0.76 (0.64); nerve ring to anterior end 0,240.33 (0.28); excretory pore to anterior end 0.32-0.61 (0.48); deirid to anterior end 0.09-0.16 (0.11); tail 0.14—0.21 (0.17); vulva to posterior end 0.27—0.34 (0.31); vagina 0.45—0.70 (0.58); egg 0.090-0.100 (0.093) x 0.040—0.050 (0.048). Female tail short, conical; vulva immediately anterior to anus; vagina elongate, convoluted, not recurrent; egg ellipsoidal. Remarks The prominent lips and symmetrical buccal capsule distinguish C. oweni sp. nov. from all congeners except C. artemis Beveridge, 1998, C. caenis Beveridge, 1998, C. clymene, C. dindymene Beveridge, 1998, C. hypsipyle, C. 18 I. BEVERIDGE FIGURES 90-101. Cloacina oweni sp. nov., types. 90. Anterior end, lateral view. 91. Cephalic extremity, lateral view, dorsal aspect on left-hand side. 92. Cephalic extremity, dorsal view. 93. Cephalic extremity, apical view. 94. Cephalic extremity, transverse optical section through buccal capsule. 95. Submedian cephalic papilla, lateral view. 96. Preneural region of oesophagus, showing denticle, dorsal view. 97. Bursa, apical view. 98. Genital cone, dorsal view and gubernaculum. 99. Spicule tip, lateral view. 100. Female tail, lateral view. 101. Female genital system, lateral view. Scale bars: 90, 97, 100, 101, 0.1 mm; 91—96, 98, 99, 0.01 mm. CLOACINA FROM PAPUA NEW GUINEA 19 linstowi Johnston & Mawson, 1940, C. maia Beveridge, 1998, C. parva, C. robertsi, C. smalesae, C. thetidis Johnston & Mawson, 1939 and C. wallabiae Johnston & Mawson, 1939. The presence of a single dorsal denticle in the oesophagus and the lack of oesophageal bosses distinguishes it from all of these species except C. dindymene. It differs from C. dindymene in the shape of the submedian cephalic papillae which have elongated distal segments in C. dindymene, and in the shape of the dorsal ray, the bifurcations of which form an acute angle in C. dindymene rather than the obtuse angle seen in C. oweni. In features of the head, C. oweni is most likely to be confused with C. robertsi, found in rock wallabies of the genus Petrogale Gray, 1837 in eastern Australia. C. oweni is readily differentiated, however, by the oesophageal denticle and the lack of a prominently recurrent vagina as exhibited by C. robertsi. C. oweni is abundant in the stomachs of agile wallabies in the Bula Plain region of Papua New Guinea, but has not been found in the same host species in northern Australia in spite of relatively intensive examination of this host in all of the northern states (Speare et al. 1983; Beveridge et al. 1998). More surprising is the close resemblance of this species to congeners occurring in rock wallabies in Australia, a relationship for which no simple explanation currently exists. Cloacina papuensis sp. nov. (Figs 102-115) Types: From stomach of Macropus agilis, Bula Plain, Bensbach, Papua New Guinea, coll. I. Owen, May, 1998. Holotype 6, SAM AHC 31451; allotype 2, SAM AHC 31452. Paratypes: 53, 62, SAM AHC 31453, 1¢, 12, BMNH 1998.9.28.13-14. Slide preparations of apical view of mouth and bursa, SAM AHC 28386. Material examined: From Macropus agilis: types. Description Small nematodes; cervical cuticle not inflated; transverse cuticular annulations widely spaced, 0.010 apart. Submedian cephalic papillae small, conical, 0.006 long, projecting anteriorly from peri-oral cuticle; distal segment ovoid, 0.002 long, shorter than cylindrical proximal segment, 0.004 long. Buccal capsule shallow, symmetrical in lateral and dorso-ventral views; circular in transverse section, wider than deep, wall without prominent striations; anterior margin smooth. Leaf crown elements 8 in number, recurved at tips; peri-oral cuticle not inflated into lip-like lobes attached to each element. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland in prominent, bilobed, tooth- like structure projecting into lumen of buccal capsule; each ventral sector with 2 triangular sclerotised projections. Oesophagus claviform, slender; lining without sclerotised bosses; single dorsal denticle present immediately anterior to nerve ring; preneural swelling present. Nerve ring in mid-oesophageal region; deirids in anterior oesophageal region; excretory pore between nerve ring and oesophago-intestinal junction. Male. Measurements of 5 specimens, types. Total length 3.18-5.09 (4.57); maximum width 0.18-0.32 (0.23); buccal capsule 0.007-—0.010 (0.008) x 0.020—0.026 (0.021); oesophagus 0.31- 0.41 (0.36); nerve ring to anterior end 0.16-0.17 (0.17); excretory pore to anterior end 0.20—0.34 (0.28); deirid to anterior end 0.08-0.11 (0.10); spicules 1.84—2.26 (2.05); gubernaculum 0.025— 0.030 (0.029) long. Dorsal ray broad at origin; major bifurcation occurs at 1/2 length; angle of bifurcation acute; external branchlets arise after major bifurcation, shorter and more robust than internals, directed postero-laterally, not reaching margin of bursa; internal branchlets directed posteriorly, not reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, slightly wider than long. Spicule tip bifid; ala diminishes gradually in width towards spicule tip; anterior lip of genital cone conical; posterior lip with paired projections and cuticular inflation of internal surface of bursa on either side. Female. Measurements of 5 specimens, types. Total length 5.14-6.05 (5.57); maximum width 0.25-0.36 (0.32); buccal capsule 0.005—0.008 (0.007) x 0.020—0.025 (0.023); oesophagus 0.35— 0.39 (0.38); nerve ring to anterior end 0.17-0.18 (0.17); excretory pore to anterior end 0.23-0.38 (0.33); deirid to anterior end 0.07-0.011 (0.09); tail 0.14—0.22 (0.19); vulva to posterior end 0.30— 0.44 (0.39); vagina 0.47—-0.63 (0.53); egg 0.070— 0.085 (0.080) x 0.040—0.050 (0.046). Female tail short, conical; vulva immediately anterior to anus; vagina elongate, convoluted, not recurrent; egg ellipsoidal. Remarks The simple, symmetrical buccal capsule, lack of 20 I. BEVERIDGE FIGURES 102-115. Cloacina papuensis sp. nov., types. 102. Anterior end, lateral view. 103. Cephalic extremity, lateral view, dorsal aspect on left-hand side. 104. Cephalic extremity, dorsal view. 105. Submedian cephalic papilla, lateral view. 106. Cephalic extremity, apical view. 107. Cephalic extremity, transverse optical section through buccal capsule. 108. Preneural region of oesophagus, showing denticle, lateral view, dorsal aspect on left-hand side. 109. Preneural region of oesophagus, showing denticle, dorsal view. 110. Spicule tips, ventral view. 111. Bursa, ventral view. 112. Bursa, lateral view. 113. Female tail, lateral view. 114. Female genital system, lateral view. 115. Genital cone, dorsal view. Scale bars: 113, 114, 0.1 mm; 102-104, 106-112, 115 0.01 mm; 105, 0.003 mm. CLOACINA FROM PAPUA NEW GUINEA 21 prominent lips and unornamented oesophagus with a single dorsal denticle differentiate C. papuensis sp. nov. from all congeners except C. cornuta, C. dirce, C. longispiculata and C. sciron. C. papuensis is differentiated from C. sciron primarily in having the deirid anterior to, rather than at the level of, the nerve ring. In C. sciron, the anterior margin of the buccal capsule arches anteriorly and is sinuous. C. papuensis differs from C. longispiculata in lacking the prominent shoulder-like inflations of the cervical cuticle, in having the excretory pore at the level of the oesophago-intestinal junction rather than well posterior to it as in C. longispiculata, and in having eight leaf crown elements rather than the six present in C. longispiculata. It differs from C. dirce in lacking the prominent cervical inflation of the cuticle, in the shape of the submedian papillae which have an extended distal segment in C. dirce, in spicule lengths (1.84—2.26 (2.05) mm long in C. papuensis, 3.48-3.95 (3.70) mm long in C. dirce), and in the shape of the vagina which is longer and more convoluted in C. dirce. Therefore, C. papuensis most closely resembles C. cornuta, also a parasite of Macropus agilis, in having a prominent dorsal oesophageal tooth. It differs from C. cornuta in lacking a cervical cuticular inflation, in the shape of the submedian papillae which have elongate distal segments in C. cornuta, in the shape of the dorsal ray which in C. cornuta terminates in very short subequal internal and external branchlets, in spicule length (1.84-2.26 (2.05) in C. papuensis, 1.38-1.62 (1.51) mm in C. cornuta), and in the length and shape of the female tail which is very short and prominently swollen in C. cornuta. Cloacina polymela sp. nov. (Figs 116-128) Types: From stomach of Dorcopsis hageni, Usino, Madang, Papua New Guinea, coll. T. Reardon, May, 1987. Holotype 6, SAM AHC 31454; allotype 2, SAM AHC 31455. Paratypes: 43, 82, SAM AHC 31456, 1d, 12, BMNH 2001.4,10.10-11, 1d, 12, USNPC 91138. Slide preparations of apical views of mouth and bursa, SAM AHC 28387. Material examined: From Dorcopsis hageni: types. Description Robust nematodes; anterior extremity deviated dorsally; cervical cuticle inflated to level of excretory pore; transverse cuticular annulations widely spaced, 0.030—0.062 apart. Submedian cephalic papillae prominent, 0.016 long, projecting anteriorly from inflated peri-oral cuticle; distal segment acute, conical, 0.010 long, longer than cylindrical proximal segment, 0.006 long. Cephalic collar prominently inflated on dorsal and ventral aspects, anterior margin of collar rugose; amphids conical, projecting above level of cephalic collar. Buccal capsule very shallow, symmetrical, anterior margin highly sinuous; due to dorsal deviation of anterior extremity, buccal capsule appears to arch dorsally in ventral views; oval and dorso-ventrally elongate in transverse section, wider than deep, wall finely striated. Leaf crown elements 8 in number, recurved at tips; peri-oral cuticle inflated into lip- like lobes attached to each element. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, not projecting into lumen of buccal capsule. Oesophagus claviform, robust, anterior section of greater diameter than region posterior to nerve ring; lining without sclerotised bosses; denticles absent in preneural region. Nerve ring posterior to mid point of oesophagus; deirids slightly anterior to level of nerve ring; excretory pore between nerve ring and oesophago-intestinal junction. Male. Measurements of 9 specimens, types. Total length 5.86-6.03 (5.93); maximum width 0.29-0.46 (0.39); buccal capsule 0.020-—0.030 (0.025) x 0.115—0.150 (0.130); oesophagus 0.66— 0.79 (0.72); nerve ring to anterior end 0.44-0.53 (0.48); excretory pore to anterior end 0.55—0.70 (0.65); deirid to anterior end 0.36-0.45 (0.39); spicules 2.61—2.71 (2.65); gubernaculum 0.035— 0.050 (0.045) long. Dorsal ray broad at origin; major bifurcation occurs at 1/2 length; angle of bifurcation acute; external branchlets arise after major bifurcation, much shorter than internals, directed postero- laterally, not reaching margin of bursa; internal branchlets elongate, directed posteriorly, almost reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, subcordate. Spicule tip blunt, slightly recurved; ala diminishes gradually in width towards spicule tip; anterior lip of genital cone conical; posterior lip with paired projections and cuticular inflation of internal surface of bursa on either side. Female. Measurements of 10 specimens, types. Total length 5.45-8.22 (7.14); maximum width 0.33-0.59 (0.46); buccal capsule 0.015—0.030 (0.025) x 0.120—0.160 (0.150); oesophagus 0.70— 22 I. BEVERIDGE 124 FIGURES 116-128. Cloacina polymela sp. nov., types. 116. Anterior end, lateral view, showing dorsal deviation of head. 117. Cephalic extremity, lateral view, dorsal aspect on left-hand side. 118. Cephalic extremity, ventral view. 119. Cephalic extremity, dorsal view. 120. Cephalic extremity, apical view. 121. Cephalic extremity, transverse optical section through buccal capsule. 122. Submedian cephalic papilla, lateral view. 123. Amphid, lateral view. 124. Bursa, ventral view. 125. Spicule tip, ventral view. 126. Gubernaculum and genital cone, dorsal view. 127. Female tail, lateral view. 128. Female genital system, lateral view. Scale bars: 116, 124, 127, 128, 0.1 mm; 117- 123, 125, 126, 0.01 mm. CLOACINA FROM PAPUA NEW GUINEA 23 0.94 (0.82); nerve ring to anterior end 0.45-0.61 (0.54); excretory pore to anterior end 0.55—0.85 (0.75); deirid to anterior end 0.31-0.50 (0.42); tail 0.21—0.30 (0.24); vulva to posterior end 0.33-0.50 (0.38); vagina 0.49-0.68 (0.60); egg 0.080-0.085 (0.082) x 0.040—-0.045 (0.042). Female tail slender, conical; vulva immediately anterior to anus; vagina elongate, sinuous, extends slightly anterior to vestibule with short recurrent section; egg ellipsoidal. Remarks Cloacina polymela sp. nov. is a highly distinctive species which differs from all congeners in having the anterior extremity deviated dorsally. This feature appears in every specimen and therefore is not likely to be a fixation artefact. In addition, it differs from congeners except C. dryope Beveridge, 1998 and C. sappho in having an oesophagus in which the preneural region is of greater diameter than the posterior part. C. polymela is distinguished from C. dryope in having eight rather than six leaf crown elements, in having a more posterior deirid, in having the excretory pore anterior to the oesophago-intestinal junction rather than posterior to it, and in the shape of the buccal capsule which is extremely shallow in C. dryope and in which the anterior margin has only slight saliences rather than prominent undulations. C. polymela and C. sappho are the only two members of the genus in which the amphids form acutely pointed conical projections above the cephalic collar. In all other species, the amphids are dome-shaped and do not project obviously beyond the collar. C. polymela also resembles C. sappho in the shape of the dorsal ray of the bursa and in possessing eight leaf crown elements. However, it differs in having prominent inflations of the cephalic collar on the dorsal and ventral surface, a feature which resembles the lip-like projections present in C. nephele. C. polymela also differs from C. sappho in spicule length (2.61—2.71 (2.65) mm in C. polymela; 1.30—1.50 (1.38) mm in C. sappho) and in the shape of the vagina which is recurrent in C. polymela but short and straight in C. sappho. Cloacina praxithea sp. nov. (Figs 129-138) Types: From stomach of Dorcopsis hageni, Usino, Madang, Papua New Guinea, coll. T. Reardon, May, 1987. Holotype 6, SAM AHC 31457. Paratypes: 23, SAM AHC 31458, 16, BMNH 2001.4.10.12; 1d, USNPC 91139. Slide preparations of apical views of mouth and bursa, SAM AHC 28388. Material examined: From Dorcopsis hageni: types. Description Robust nematodes; cervical cuticle slightly inflated to beyond level of excretory pore; transverse cuticular annulations widely spaced, 0.015 apart. Submedian cephalic papillae prominent, 0.019 long, projecting anteriorly from peri-oral cuticle; distal segment ovoid, 0.008 long, deviated medially, slightly shorter than asymmetrical, cylindrical proximal segment, 0.011 long; submedian papillae situated anteriorly on cephalic collar; amphids situated posteriorly, below anterior margin of buccal capsule in lateral views. Buccal capsule shallow, symmetrical in lateral and dorso-ventral views, wall with faint striations; anterior margin undulate, with anterior projections associated with each leaf crown element. Leaf crown elements 8 in number, not recurved at tips; peri-oral cuticle inflated into lobes attached to each element. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, not projecting into lumen of buccal capsule. Oesophagus subcylindrical, almost claviform at base, robust; lining without sclerotised bosses or denticles; preneural swelling of oesophagus absent. Nerve ring in mid- oesophageal region; deirids anterior to nerve ring; excretory pore at level of, or slightly anterior to, oesophago-intestinal junction. Male. Measurements of 6 specimens, types. Total length 3.71-5.34 (4.86); maximum width 0.32-0.38 (0.36); buccal capsule 0.015—0.018 (0.016) x 0.050—0.055 (0.054); oesophagus 0.66— 0.78 (0.71); nerve ring to anterior end 0.34-0.38 (0.36); excretory pore to anterior end 0.43-0.49 (0.46); deirid to anterior end 0.27-0.32 (0.30); spicules 1.30-1.58 (1.43); gubernaculum 0.030— 0.040 (0.033) long. Dorsal ray broad at origin; major bifurcation occurs at 1/2 length; angle of bifurcation acute; external branchlets arise immediately after major bifurcation, much shorter than internals, directed postero-laterally, not reaching margin of bursa; internal branchlets directed posteriorly, not reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, subcordate, slightly wider than long. Spicule tip blunt; ala diminishes gradually in width then terminates abruptly anterior to spicule tip; anterior lip of genital cone conical; posterior 24 I. BEVERIDGE 136 FIGURES 129-138. Cloacina praxithea sp. nov., types. 129. Anterior end, lateral view. 130. Cephalic extremity, lateral view, dorsal aspect on right-hand side. 131. Cephalic extremity, dorsal view; arrows indicate amphids. 132. Cephalic extremity, apical view; arrows indicate amphids. 133. Cephalic extremity, transverse optical section through buccal capsule. 134. Submedian cephalic papilla, lateral view. 135. Gubernaculum, ventral view. 136. Bursa, apical view. 137. Genital cone, dorsal view. 138. Spicule tip, lateral view. Scale bars: 129, 136, 0.1 mm; 130-135, 137, 138, 0.01 mm. CLOACINA FROM PAPUA NEW GUINEA 25 lip with paired projections and cuticular inflation of internal surface of bursa on either side. Female. Not seen Remarks Cloacina praxithea sp. noy. is distinguished from all congeners by the position of the amphids on the cephalic collar. In other species the amphids are at the same level as the submedian papillae, anterior to the level of the buccal capsule, while in C. praxithea the amphids, in lateral view, are below the level of the anterior margin of the buccal capsule (Fig. 130). C. parxithea is characterised by a symmetrical, lobed buccal capsule without bosses or denticles in the oesophagus. As such, it has similarities with the Australian species C. artemis, C. dryope, C. hebe, C. hypsipyle, C. linstowi, C. maia, C. thetidis and C. wallabiae, all of which differ from it in having six rather than eight leaf crown elements. Among the species known from New Guinea, the features of the buccal capsule, as well as the presence of eight leaf crown elements, indicate affinities with C. syphax, C. solon, C. sappho and C. solymus. C. solymus has tiny cephalic papillae, in striking contrast to the prominent papillae of C. praxithea, while C. sappho and C. polymela have the amphids on or forming projections beyond the level of the cephalic collar; C. solon has a greatly inflated cervical cuticle and its spicules are 2.60— 2.95 (2.81) mm in length compared with 1.93-— 2.05 (1.99) mm in C. praxithea; C. syphax differs in having the lateral branchlets of the dorsal ray arise anterior to the principal bifurcation. Therefore, C. praxithea is readily distinguishable from all congeners even in the absence of females, which are currently unknown. Cloacina procris sp. nov. (Figs 139-153) Types: From stomach of Dorcopsis hageni, Usino, Madang, Papua New Guinea, coll. T. Reardon, May, 1987. Holotype ¢6, SAM AHC 31459; allotype 2, SAM AHC 31460. Paratypes: 16,22, SAM AHC 31461. Slide preparations of apical views of mouth and bursa, SAM AHC 28389. Material examined: From Dorcopsis hageni: types. Description Robust nematodes; cervical cuticle slightly inflated to level of excretory pore; transverse cuticular annulations widely spaced, 0.020 apart. Submedian cephalic papillae small, 0.0035 long, projecting anteriorly from peri-oral cuticle; distal segment ovoid, 0.0005 long, much shorter than cylindrical proximal segment, 0.003 long. Buccal capsule shallow, asymmetrical in lateral views, deeper on ventral side, slight anterior arching of capsule; dorsally, capsule arches anteriorly over dorsal oesophageal tooth; ventrally, curves posteriorly; oval and dorso-ventrally elongate in transverse section, wider than deep, wall with faint striations; anterior margin smooth. Leaf crown elements 8 in number, not recurved at tips; peri-oral cuticle very slightly inflated into lobes attached to each element. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, projecting prominently into lumen of buccal capsule; each sector of oesophagus bears single triangular tooth-like structure projecting into lumen. Oesophagus claviform, robust; lining with sclerotised bosses extending to level of nerve ring; single dorsal oesophageal denticle present in preneural region; preneural swelling of oesophagus absent. Nerve ring in mid-oesophageal region; deirids posterior to nerve ring; excretory pore at level of, or slightly anterior to, oesophago-intestinal junction. Male. Measurements of 3 specimens, types. Total length 3.25-4.52 (3.89); maximum width 0.17-0.32 (0.26); buccal capsule 0.015-0.018 (0.017) x 0.050—0.060 (0.054); oesophagus 0.40— 0.41 (0.40); nerve ring to anterior end 0.20-0.21 (0.21); excretory pore to anterior end 0.35—0.40 (0.38); deirid to anterior end 0.27-0.32 (0.30); spicules 1.30-1.58 (1.43); gubernaculum 0.030 (0.030) long. Dorsal lobe of bursa elongate; dorsal ray broad at origin; major bifurcation occurs at 1/2 length; angle of bifurcation acute; external branchlets arise at level of major bifurcation, much shorter than internals, directed almost laterally, not reaching margin of bursa; internal branchlets directed posteriorly, reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, slightly wider than long. Spicule tip blunt; ala diminishes gradually in width then terminates abruptly anterior to spicule tip; anterior lip of genital cone conical; posterior lip with paired projections and cuticular inflations of internal surface of bursa on either side. Female. Measurements of 3 specimens, types. Total length 4.26-5.15 (4.64); maximum width 0.37-0.47 (0.42); buccal capsule 0.018-0.020 (0.019) x 0.060-0.065 (0.062); oesophagus 26 I. BEVERIDGE FIGURES 139-153. Cloacina procris sp. nov., types. 139. Anterior end, lateral view. 140. Cephalic extremity, lateral view, dorsal aspect on left-hand side. 141. Submedian cephalic papilla, lateral view. 142. Cephalic extremity, dorsal view. 143. Cephalic extremity, ventral view. 144. Cephalic extremity, apical view. 145. Cephalic extremity, transverse optical section through buccal capsule. 146. Transverse optical section through anterior extremity of oesophagus showing bosses and three tooth-like projections in each of the oesophageal sectors. 147. Preneural region of oesophagus, dorsal view, showing denticle. 148. Gubernaculum, ventral view. 149. Genital cone, dorsal view. 150. Bursa, apical view. 151. Spicule tip, lateral view. 152. Female tail, lateral view. 153. Female genital system, lateral view. Scale bars: 139, 152, 153, 0.1 mm; 140, 142-151, 0.01 mm; 141, 2.5 um. CLOACINA FROM PAPUA NEW GUINEA 27 0.48-0.50 (0.49); nerve ring to anterior end 0.21-0.22 (0.22); excretory pore to anterior end 0.36—-0.40 (0.38); deirid to anterior end 0.29- 0.33 (0.31); tail 0.19-0.27 (0.23); vulva to posterior end 0.35—0.47 (0.40); vagina 0.52—0.60 (0.53); egg 0.050—0.060 (0.055) x 0.030—0.035 (0.032). Female tail slender, conical; vulva immediately anterior to anus; vagina elongate, straight, turns abruptly at entry to vestibule; egg ellipsoidal. Remarks Although described from only a small series of specimens, C. procris sp. nov. is quite distinctive morphologically. It is characterised by very small submedian cephalic papillae, a slightly asymmetrical, arched buccal capsule, eight leaf crown elements, bosses lining the anterior region of the oesophagus, a single, small oesophageal denticle, deirid posterior to the nerve ring, an elongate dorsal ray with the external branchlets arising immediately after the principal bifurcation, and a straight vagina. These features distinguish the species from all congeners. The asymmetrical buccal capsule, oesophageal bosses and dorsal denticle with the posterior position of the deirid distinguish the species from all others except C. eos Beveridge, 1998, C. papillata Beveridge, 1979 and C. sterope. In C. eos and C. papillata there are only six rather than eight leaf crown elements and the vagina is prominently recurrent. In C. sterope, the buccal capsule exhibits a much greater degree of asymmetry, the submedian cephalic papillae are larger and more prominent and the spicules are longer (1.67—2.07 (1.96) mm in C. sterope, 1.30- 1.58 (1.43) mm in C. procris). Cloacina sterope Beveridge & Speare, 1999 Material examined: 33, 4°, from stomach of Dorcopsis hageni, Usino, Madang, Papua New Guinea, coll. T. Reardon, May, 1987, SAM AHC 31462. Remarks Cloacina sterope was described by Beveridge and Speare (1999) from Dorcopsulus vanhearni from the Chimbu Province of New Guinea. Morphological features as well as measurements of the new specimens from D. hageni agree well with the original description. D. hageni represents a new host record. Cloacina syphax Beveridge & Speare, 1999 Material examined: 176, 92, from stomach of Dorcopsis hageni, Usino, Madang, Papua New Guinea, coll. T. Reardon, May, 1987, SAM AHC 31469; BMNH 2001.4.10.13. Remarks Cloacina syphax was described from Dorcopsulus vanheurni by Beveridge and Speare (1999). The current collection represents a new host record. Cloacina theope sp. nov. (Figs 154-166) Types: From stomach of Dendrolagus matschiei Forster & Rothschild, 1907, Huon Peninsula, Morobe Province, Papua New Guinea, coll. J. Mayer, July 1999. Holotype ¢, USNPC 91140; allotype 2, USNPC 91141. Paratypes: 2d, 49, USNPC 91142; 1d, on slides, SAM AHC 28390. Material examined: From Dendrolagus matschiei: types. From Dendrolagus dorianus: 103, 5¢, Lake Trist, Papua New Guinea, coll. I. Redmond, 1979, BMNH 1981. 4486-4505. Description Robust nematodes; cervical cuticle slightly inflated; transverse cuticular annulations widely spaced, 0.021—0.044 apart. Submedian cephalic papillae prominent, 0.017 long, projecting anteriorly from inflated peri-oral cuticle; distal segment ovoid, 0.008 long, directed slightly medially, slightly shorter than robust, asymmetrical proximal segment, 0.009 long. Buccal capsule shallow, symmetrical in lateral and dorso-ventral views; circular in transverse section, wall without prominent striations; anterior margin smooth. Leaf crown elements 8 in number, recurved at tips; peri-oral cuticle not inflated into lip-like lobes attached to each element. Dorsal sector of oesophagus bearing opening of dorsal oesophageal gland, not projecting into lumen of buccal capsule. Oesophagus claviform, slender; lining without sclerotised bosses; denticles present in preneural region; dorsal denticle prominent; two subventral thickenings of oesophageal lining giving appearance of denticles, at same level as dorsal denticle. Nerve ring in mid-oesophageal region; deirids at level of nerve ring; excretory pore at level of oesophago-intestinal junction. Male. Measurements of 4 specimens, types. Total length 4.68-5.04 (4.88); maximum width 28 I. BEVERIDGE FIGURES 154-166. Cloacina theope sp. nov., types. 154. Anterior end, lateral view. 155. Cephalic extremity, lateral view, dorsal aspect on left-hand side. 156. Cephalic extremity, dorsal view. 157. Submedian cephalic papilla, lateral view. 158. Cephalic extremity, apical view. 159. Cephalic extremity, transverse optical section through buccal capsule. 160. Preneural region of oesophagus, lateral view, dorsal aspect on left-hand side, showing denticles. 161. Preneural region of oesophagus, dorsal view, showing denticles. 162. Bursa, apical view. 163. Gubernaculum, genital cone and spicule sheaths, dorsal view. 164. Spicule tip, lateral view. 165. Female tail, lateral view. 166. Female genital system, lateral view. Scale bars: 154, 162, 165, 166, 0.1 mm; 155-161, 163-164, 0.01 mm. CLOACINA FROM PAPUA NEW GUINEA 29 0.30-0.32 (0.31); buccal capsule 0.010 (0.010) x 0.030-0.033 (0.031); oesophagus 0.51-0.60 (0.55); nerve ring to anterior end 0.18-0.20 (0.19); excretory pore to anterior end 0.31-0.41 (0.35); deirid to anterior end 0.13-0.23 (0.18); spicules 2.29-2.63 (2.52); gubernaculum 0.015— 0.020 (0.018) long. Dorsal ray broad at origin; major bifurcation occurs at 1/2 length; angle of bifurcation obtuse; external branchlets arise immediately after major bifurcation, shorter than internals, directed laterally, not reaching margin of bursa; internal branchlets directed postero-laterally, almost reaching margin of bursa. Externo-dorsal ray not reaching margin of bursa. Gubernaculum prominent, slightly wider than long. Spicule tip minutely bifid; ala diminishes gradually in width towards spicule tip; anterior lip of genital cone conical; posterior lip with paired projections and cuticular inflations of internal surface of bursa on either side. Female. Measurements of 5 specimens, types. Total length 6.09-6.71 (6.29); maximum width 0.39-0.43 (0.41); buccal capsule 0.008—0.010 (0.009) x 0.030-0.035 (0.032); oesophagus 0.52— 0.64 (0.59); nerve ring to anterior end 0.25 (0.25); excretory pore to anterior end 0.30—0.51 (0.43); deirid to anterior end 0.14—0.23 (0.20); tail 0.28— 0.36 (0.33); vulva to posterior end 0.49-0.68 (0.59); vagina 1.03-1.33 (1.19); egg 0.085-0.095 (0.089) x 0.045—0.050 (0.048). Female tail slender, conical; vulva immediately anterior to anus; vagina elongate, sinuous, extends slightly anterior to vestibule with short recurrent section; egg ellipsoidal. Remarks Cloacina theope sp. nov. differs from congeners in having three oesophageal denticles in the preneural region of the oesophagus which are at the same level, but are of dissimilar sizes. The dorsal denticle is well developed but the subventral denticles are mere thickenings of the oesophageal lining and may not warrant the designation as denticles. In all other species with three denticles at the same level in the oesophagus, C. daveyi, C. eurynome, C. hera, C. hermes, C. hestia, the denticles are equally developed, although in C. hestia and C. eurynome all three denticles may be weakly sclerotised. C. theope differs from C. daveyi, which has submedian cephalic papillae in which the distal segment is large and directed medially, and from all the remaining species, which have the deirid in the anterior oesophageal position and six rather than eight leaf crown elements. It is most similar to C. eurynome from which it differs principally in having shorter spicules (3.57-3.98 (3.77) mm in C. eurynome, 2.29-2.63 (2.52) mm in C. theope). If the subventral denticles are excluded from consideration, then the unornamented anterior oesophagus, a symmetrical buccal capsule with a smooth anterior margin, and the presence of a dorsal denticle indicates similarities with C. cornuta, C. dindymene, C. dirce, C. ernabella, C. longispiculata and C. sciron. C. ernabella differs in the shape of the cephalic papillae (globose, medially directed distal segment in C. ernabella), while all species except C. sciron differ in having the deirid in the anterior oesophageal region rather than at the level of the nerve ring. C. theope differs from C. sciron, which has an anteriorly arched buccal capsule, longer spicules (2.83-3.30 (3.00) mm in C. sciron, 2.29-2.63 (2.52) mm in C. theope), a longer dorsal ray and a vagina which is not recurrent. In C. theope the recurrent section of the vagina is twisted around the ascending arm in a manner similar to that found in C. syphax and C. eurynome. C. theope differs from C. syphax in the shape of the buccal capsule, which has an undulating anterior margin in C. syphax but is straight in C. theope. Therefore, C. theope is clearly distinguishable from all congeners. DIscussION The new records and new species reported in this paper suggest that a diverse array of species of Cloacina exists in macropodid marsupials in Papua New Guinea. Conclusions need to be guarded as the current collections have been obtained from a small number of individuals of each host species, in some instances from a single animal. As yet, there have been no comprehensive surveys of New Guinean macropodids for parasites, and several species of Thylogale, Dorcopsis, Dorcopsulus and Dendrolagus have apparently yet to be examined for helminths (Spratt et al. 1991). In addition, Beveridge (1998) and Beveridge and Speare (1999) have listed museum records of additional undescribed species from some of these hosts which cannot currently be named due to lack of adequate material. In spite of the relatively rudimentary knowledge of the New Guinean parasite fauna, some preliminary comparisons can be made with species present in Australia. In terms of morphological characters, the species of Cloacina present in New 30 I. BEVERIDGE Guinea are virtually as diverse as those present in Australia. Of the various morphological features of the genus, only the spiral twisting of the proximal, recurrent region of the vagina (C. syphax, C. eurynome, C. theope) and the presence of amphids on conical projections (C. praxithea, C. sappho) appear to be restricted to species from New Guinea. Two other apparently autapomorphic characters, using Arundelia dissimilis (Johnston & Mawson, 1939) as an outgroup, are the posterior position of the amphids in C. praxithea and the dorsal deviation of the head in C. polymela. A. dissimilis is the sole member of a closely related genus within the tribe Cloacininea (see Beveridge 1987) and is therefore considered to be an appropriate outgroup for polarising morphological characters. Characters such as the origin of the external branchlets of the dorsal ray anterior to the major bifurcation are more common in species from New Guinea, but are also found in species from northern Australia. No obvious patterns are detectable in the distribution of other characters. While only tentative conclusions can be drawn in the absence of a formal phylogenetic analysis, there do not appear to be any major distinctions which can be drawn between species occurring in New Guinea compared with those present in Australia. In several instances (C. australis, C. cloelia, C. cornuta, C. cybele, C. dahli), the same species of Cloacina are found in Australia and in New Guinea, a phenomenon explicable in terms of host distribution. C. australis and C. cornuta are both parasites of M. agilis, a wallaby which is common in northern Australia (Strahan 1995) and which invaded southern New Guinea relatively recently from Australia (Flannery 1995). Similarly, C. cloelia, C. cybele and C. dahli occur in pademelons of the genus Thylogale, and at least one species, T. stigmatica, is thought to be a recent arrival in New Guinea (Winter 1997). However, the occurrence of these same nematodes in species of Thylogale restricted to New Guinea suggests that they have been present for a longer period of time than the recent invasion of T. stigmatica. The remaining species of Cloacina currently described from New Guinea occur in hosts which are endemic. Species of scrub wallabies Dorcopsis and Dorcopsulus do not occur in Australia, while the tree kangaroos, Dendrolagus spp., have apparently radiated in New Guinea and invaded north-eastern Australia secondarily (Flannery et al. 1996). The recent arrival of the genus Dendrolagus in Australia may have contributed to the absence of species of Cloacina in D. lumholtzi and D. bennettianus. While the records of species of Cloacina from New Guinea are still fragmentary, it appears that a substantial New Guinean fauna exists, and its relationships with its hosts and the comparisons that can be made with Australian representatives may provide insights into the evolution and hence the diversity of this nematode genus. KEY TO SPECIES OF CLOACINA IN MACROPODIDS FROM PAPUA NEW GUINEA Denticles present in preneural region of oesophageal lumen... eeeeeeeeeeeeeeee 2 — Denticles absent... eee 13 Paired ventral oesophageal denticles anterior to dorsal denticle ......... C. dahli — Single dorsal denticle, or three denticles at same level in oesophagus .............00 3 Anterior part of oesophagus lined with SClerotised BOSSES oo... eeeeeeeeeeeteeereeeeens 4 — Anterior oesophagus without sclerotised DOSSES sire ..- 02 saceste cots sutoacesntcenons dlusecrsurehs 8 Anterior oesophageal bosses much larger than other bosses; deirid anterior to nerve ring; female tail swollen ..... C. australis — Oesophageal bosses of similar size; deirid at level of nerve ring or posterior to it; female tail not swollen «0.0... eee 5) Buccal capsule symmetrical in lateral view BEST OM tos daa. Se. ALAS eee 2 ee C. erigone — Buccal capsule asymmetrical in lateral view, more shallow on dorsal aspect ... 6 Buccal capsule prominently arched anteriorly 2.8, ents ef, oe C. hyperea — Buccal capsule not prominently arched ANLETIOTLY reeiceesk filecseatebiccessteveteettaessls 7 Spicules 1.30-1.58 mm; distal segment of submedian papilla less than 1/4 length of proximal segment ................ C. procris — Spicules 1.67—2.07 mm; distal segment of submedian papilla only slightly shorter than proximal segment ......... C. sterope Dorsal denticle and 2 vestigial subventral denticles present ...... ce eeeeceeeeeteeeeeeeees 9 — Single dorsal denticle present ............ 10 Spicules 3.57—3.98 mm .....C. eurynome — Spicules 2.29-2.63 mm .......... C. theope 10. — Prominent lips present ............. C. oweni == ‘Lips absent: 4. shen ld aanieuace 11 11. — _ Deiridatlevel of nerve ring; anterior margin CLOACINA FROM PAPUA NEW GUINEA 31 of buccal capsule arched......... C. sciron — Deirid in anterior oesophageal region, anterior margin of buccal capsule NOtzZontale A. ace iba esa 12 12. — Cervical cuticle and female tail inflated, distal segment of submedian papilla almost as long as proximal segment, spicules 1.38-1.62 MM... eee C. cornuta — Cervicalcuticle and female tail not inflated, distal segment of submedian cephalic papilla much shorter than proximal segment, spicules 1.84—2.26 mm............ ee tint de Mawngrdedideigereetchs C. papuensis 13. — Anteriormargin of buccal capsule undulate ogedeusecersih Mlanadooowadesepagensedcbaregethyivpectesens 14 — Anterior margin of buccal capsule not MNGUIALC 2.0, astsies AO ME Mae eel 18 14. — Amphids on or forming conical elevations projecting beyond cephalic collar...... 15 — Amphidsembedded within cephalic collar aptacabng db guy ke EOE 16 15. — Head deviated dorsally, cephalic collar elevated between adjacent submedian papillae 2s nce a cere C. polymela — Head directed anteriorly, cephalic collar depressed between adjacent submedian PAPAS. is sivscsospeedyeetesueideseter C. sappho 16. — External branchlets of dorsal ray arise before major bifurcation ........ C. syphax — External branchlets of dorsal ray arise after major bifurcation ..........c cee 17 17. — Cervical cuticle prominently inflated, submedian papillae project prominently beyond cephalic collar .............. C. solon — Cervical cuticle not inflated, submedian papillae very small, barely project above cephalic collar... C. solymus 18. — External branchlets of dorsal ray arise before major bifurcation ...............0.. 19 — External branchlets of dorsal ray arise after major bifurcation .........0.ccee 21 REFERENCES BEVERIDGE, I. 1987. The systematic status of Australian Strongyloidea (Nematoda). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4éme série 9: 107-126. BEVERIDGE, I. 1998. Taxonomic revision of the genus Cloacina von Linstow (Nematoda: Strongyloidea) 19. 20. 21. 22. 23. 24. 25. Deirid anterior to nerve ring ..............006 pabsstbesebadedosecdeaeaenegengyyeied C. cunctabunda Deirid posterior to nerve ring ............ 20 Spicule length 2.34-2.97 mm............. Spicule length 1.23-1.45 mm............... pbabbdeecvdiaretdacsacsssouns tha tgllenay C. caballeroi Buccal capsule sinuous in apical views, appearing as extra thickenings of wall in lateral or dorso-ventral ViewS ...........006. sendepeh dnesiesnsyasoatemhadcernaced edeetiv diy seieiodarged 22 Buccal capsule prominently arched AMLETIOTL YE ceil. pereredeteabteceed C. nephele Buccal capsule not prominently arched . Pighiethdrgr brig tadessegdepateenescude ry hs itteaesedy 23 Amphids posterior to anterior margin of buccal capsule 0. C. praxithea Amphids anterior to anterior margin of buccal capsule wo... cee eseeeeteeeeeteees 24 Oesophagus with sclerotised bosses, spicule tip foot-shaped .......... C. cloelia Oesophagus lacking bosses, spicule tip SUIMPIC: ssh s.2-,escaedencsecenacuecenrssdicenedevidvass 25 Spicule length 3.90-5.05 mm, spicule tip SIMPIC ts. 50, cihersseere?Bepugenctetesteers C. hecale Spicule length 2.30-2.90 mm, spicule tip with enlarged flange, ala terminating abruptly anterior to spicule tip ............... ACKNOWLEDGMENTS Sincere thanks are due to Tim Flannery, Terry Reardon, Ian Redmond, Joerg Mayer and Ifor Owen for collecting the material, sometimes under trying circumstances, on which this paper is based and to Eileen Harris and Janet Martin for making the material available for study. Rosemary Harrigan is thanked for excellent technical assistance and Dave Spratt for comments on a draft of the manuscript. from macropodid marsupials. Invertebrate Zoology 12: 273-508. BEVERIDGE, I. 1999. New species of Cloacina von Linstow, 1898 (Nematoda: Strongyloidea) parasitic in the stomach of the quokka, Setonix brachyurus (Marsupialia: Macropodidae), from Western 32 I. BEVERIDGE Australia. Transactions of the Royal Society of South Australia 123: 17-30. BEVERIDGE, I. & SPEARE, R. 1999. New species of parasitic nematodes from Dorcopsulus vanheurni (Marsupialia: Macropodidae) from Papua New Guinea. Transactions of the Royal Society of South Australia 123: 85-100. BEVERIDGE, I., CHILTON, N. B., JOHNSON, P. M., SPEARE, R., SMALES, L. R. & SPRATT, D. M. 1998. Helminth parasite communities of kangaroos and wallabies (Macropus spp. and Wallabia bicolor) from north and central Queensland. Australian Journal of Zoology 46: 473-495. FLANNERY, T. F. 1995. ‘Mammals of New Guinea’. Reed Books: New South Wales. FLANNERY, T. F., BOEADI, & SZALAY, A. L. 1995. A new tree kangaroo (Dendrolagus: Marsupialia) from Irian Jaya, Indonesia, with notes on ethnography and the evolution of tree kangaroos. Mammalia 59: 65-84. FLANNERY, T. F., MARTIN, R. & SZALAY, A. L. 1996. ‘Tree kangaroos: a curious natural history’. Reed Books: Melbourne. LEWIN, R. A. 1999. ‘Merde. Excursions into scientific, cultural and sociohistorical coprology’. Aurum Press: London. MAWSON, P. M. 1977. Cloacina cornuta (Davey & ~ Wood) and C. caballeroi sp. nov. (Nematoda : Cloacininae) from macropods from Papua. Excerta Parasitologica en memoria del Doctor Eduardo Caballero y Caballero. Instituto de Biologia, Publicaciones Especiales 4: 455-458. SPEARE, R., BEVERIDGE, I. & JOHNSON, P. M. 1983. Parasites of the agile wallaby, Macropus agilis (Marsupialia). Australian Wildlife Research 10: 89- 96. SPRATT, D. M., BEVERIDGE, I. & WALTER, E. L. 1991. A catalogue of Australasian monotremes and marsupials and their recorded helminth parasites. Records of the South Australian Museum, Monograph Series, 1: 1-105. STRAHAN, R. 1995. ’The mammals of Australia’. Reed Books, Chatswood. WINTER, J. W. 1997. Responses of non-volant mammals to late Quaternary climatic changes in the wet tropics region of north-eastern Australia. Wildlife Research 24: 493-511. DARWIN FORMATION (EARLY CRETACEOUS, NORTHERN TERRITORY) MARINE REPTILE REMAINS IN THE SOUTH AUSTRALIAN MUSEUM BENJAMIN P. KEAR Summary Marine reptile remains from Early Cretaceous (Aptian/Albian) deposits of the Darwin Formation, Northern Territory, are described from the collections of the South Australian Museum. The material includes vertebra and limb girdle elements from ichthyosaurs and plesiosauroids, as well as the first described remains from the Northern Territory. The pliosaurid specimen (a single femur) is attributed to the small-bodied (up to 3 m in length) freshwater/near-shore marine taxon Leptocleidus, supporting interpretation of the Darwin Formation as representing a near-shore marine depositional environment. A comparison of the Darwin Formation marine reptile fauna with those known from Early Cretaceous deposits elsewhere in Australia, indicates greatest compositional similarity to the Hauterivian-Barremian Birdrong Sandstone fauna of Western Australia. A potential link between the distribution of some marine reptile taxa (eg pliosaurids) and the prevailing depositional environment is also suggested. DARWIN FORMATION (EARLY CRETACEOUS, NORTHERN TERRITORY) MARINE REPTILE REMAINS IN THE SOUTH AUSTRALIAN MUSEUM BENJAMIN P. KEAR KEAR, B. P. 2002. Darwin Formation (Early Cretaceous, Northern Territory) marine reptile remains in the South Australian Museum. Records of the South Australian Museum 35(1): 33-47. Marine reptile remains from Early Cretaceous (Aptian/Albian) deposits of the Darwin Formation, Northern Territory, are described from the collections of the South Australian Museum. The material includes vertebra and limb girdle elements from ichthyosaurs and plesiosauroids, as well as the first described pliosaurid remains from the Northern Territory. The pliosaurid specimen (a single femur) is attributed to the small-bodied (up to 3m in length) freshwater/near-shore marine taxon Leptocleidus, supporting interpretation of the Darwin Formation as representing a near-shore marine depositional environment. A comparison of the Darwin Formation marine reptile fauna with those known from Early Cretaceous deposits elsewhere in Australia, indicates greatest compositional similarity to the Hauterivian—Barremian Birdrong Sandstone fauna of Western Australia. A potential link between the distribution of some marine reptile taxa (eg pliosaurids) and the prevailing depositional environment is also suggested. Benjamin P. Kear [kear.ben@saugov.sa.gov.au], *South Australian Museum, North Terrace, Adelaide, South Australia 5000; and Vertebrate Palaeontology Laboratory, School of Biological Sciences, University of New South Wales, UNSW Sydney, New South Wales. 2052; *address for correspondence. Revised manuscript received | February 2002. Sometime in the 1960s (exact date unknown), a small collection of fossil marine reptile remains from the Early Cretaceous (Aptian/Albian) deposits of Casuarina Beach, Darwin Formation, Northern Territory (see Murray 1987, Fig. 1, p. 95 for locality map) was donated to the South Australian Museum. In 1994 several additional Darwin Formation marine reptile specimens were acquired as part of an exchange with the Museum and Art Gallery of the Northern Territory. A recent examination of these remains, which include isolated elements from both ichthyosaurs and plesiosauroids, has also revealed the presence of a pliosaurid femur. This specimen is attributable to the genus Leptocleidus Andrews, 1922 and represents the first documented pliosaur material from the Northern Territory. This paper describes Darwin Formation marine reptile remains currently housed in the South Australian Museum and provides a brief palaeoecological analysis based on faunal comparison with other known Early Cretaceous marine reptile bearing deposits. Marine reptile remains were first reported from the Darwin Formation by an anonymous author (1924) who described a fragmentary ichthyosaur skeleton from Fannie Bay near Darwin. This specimen was later attributed to Ichthyosaurus australis (Tiechert & Matheson 1944); however, Murray (1985) re-assigned the material, along with other ichthyosaur remains from Nightcliff, north of Fannie Bay, to Ichthyosauria gen. et sp. indet. A more recent examination by Wade (1990) suggested that all ichthyosaur material from the Darwin Formation might be synonymous with Platypterygius longmani from the Aptian—Albian of Queensland. Murray (1987) described isolated ichthyosaur and elasmosaurid remains from Casuarina Beach, assigning the material to Platypterygius sp. and Elasmosauridae gen. et sp. indet., respectively. The presence of elasmosaurid and pliosaurid remains in the Casuarina Beach deposits was also noted by Cruickshank et al. (1999) but no descriptions were provided. A comparison of the Darwin Formation marine reptile fauna with those known from Early Cretaceous deltaic-marine deposits elsewhere in Australia (Wallumbilla Formation [Doncaster Member], Allaru Mudstone, Toolebuc Formation and Mackunda Formation, Queensland; Bulldog Shale, South Australia; Wallumbilla Formation {Doncaster Member], New South Wales; Birdrong Sandstone and Barrow Group, Western Australia) indicates greatest compositional similarity to the Hauterivian—Barremian Birdrong Sandstone fauna, with ichthyosaurs, elasmosaurids and the small-bodied pliosaurid Leptocleidus represented. A potential link between the distribution of some 34 B. P. KEAR marine reptile taxa (eg pliosaurids) and the prevailing depositional environment is also suggested. The presence of Leptocleidus, a pliosaurid genus known otherwise from freshwater and near-shore marine deposits (Andrews 1911, 1922; Strémer 1935; Cruickshank 1997; Cruickshank & Long 1997; Cruickshank et al. 1999) supports interpretation of the Darwin Formation as representing a shallow near-shore marine depositional environment (sensu Smart & Senior 1980; Murray 1985; Henderson 1998). MATERIALS AND METHODS Institutional abbreviations used: SAM, South Australian Museum, Adelaide; NTM, Museum and Art Gallery of the Northern Territory, Darwin; AM, Australian Museum, Sydney. Lithostratigraphic nomenclature follows Mory (1988) for Early Cretaceous deposits of the Darwin area and Burton & Mason (1998) for the opal-bearing sediments of White Cliffs, New South Wales. Systematic terminology follows Montani (1999) for Ichthyopterygia and Brown (1981) for Plesiosauria except in the subdivision of Pliosauroidea, in which Polycotylidae is retained as a separate family (sensu Williston 1925: White 1940; Persson 1963; Welles 1962; Carpenter 1996, 1997). Functional sub-division of the vertebral column in ichthyopterygians follows Buchholtz (2001), with designation of structural units as neck, trunk, tail stock and fluke. Interpretation of plesiosaur pelvic and femoral musculature follows Robinson (1975) and Lingham-Soliar (2000). All measurements were taken using callipers and are in millimetres (mm), GEOLOGICAL SETTING AND PRESERVATION The marine reptile-bearing deposits of the Darwin Formation are typically characterised by glauconitic sandstone and radiolarian mudstone with basal quartzose conglomerate and localised nodular phosphorite horizons (Henderson 1998). The unit forms part of the Bathurst Island Group, which rests nonconformably on Precambrian basement within the Money Shoals Platform (see Fig. 1, p. 117 and Fig. 6, p. 124 of Henderson 1998 for locality and stratigraphic setting). At its base, the Bathurst Island Group is composed of the laterally equivalent Darwin Formation and Marligar Formation, which are overlain in turn by the Wangarlu Mudstone and Moonkinu Formation. The Darwin Formation has historically been considered Albian in age on the basis of its macroinvertebrate (Day 1969; Skwarko 1966, 1968; Henderson 1990) and palynomorph (Morgan 1980) assemblages; however, more recent analyses (Henderson 1998; D. Megirian pers. comm. 2000) suggest an upper Aptian age. The Marligar Formation has been dated as Neocomian/Aptian by palynological studies (Burger 1978) and upper Aptian because of its laterally equivalent relationship with the Darwin Formation (Henderson 1998). The overlying Wangarlu Mudstone is regarded as upper Albian— Cenomanian (possibly with a basal lower Albian component, Henderson 1998) on the basis of ammonite (Henderson 1990) and palynological (Norvick & Burger 1975; Burger 1978) data. The uppermost Moonkinu Formation is considered Cenomanian—Turonian on account of its ammonite (Wright 1963) and bivalve/gastropod/ scaphopod (Skwarko 1983) faunas. The glauconitic sandstones and predominant radiolarian mudstones of the Darwin Formation reflect a near-shore marine/paralic depositional environment (Smart & Senior 1980; Henderson 1998). The presence of a limited benthic invertebrate fauna also suggests unfavourable, possibly anoxic bottom conditions (Henderson 1998). The Darwin Formation marine reptile remains generally occur as heavily weathered isolated elements, although several fragmentary skeletons have been recovered (Murray 1985; D. Megirian pers. comm. 2000). The disarticulated nature of the remains led Murray (1985) to speculate that an inshore and/or tidal depositional environment may have facilitated stranding of decaying carcasses on mudflats prior to burial. SYSTEMATICS DIAPSIDA Osborn, 1903 ICHTHYOPTERYGIA Owen, 1860 ICHTHYOSAURIA de Blainville, 1835 OPHTHALMOSAURIA Appleby, 1956 (sensu Montani 1999) Platypterygius von Huene, 1922 cf. Platypterygius longmani DARWIN FORMATION MARINE REPTILE REMAINS w mn FIGURE 1. SAM P35039, ef. Platpterygius longmani dorsal vertebra in (A) anterior and (B) lateral views. Scale bar is 40 mm. Material trunk centra; SAM P35427 (Table 1), three SAM P35039 (Figs 1A, B; Table 1), four associated anterior tail stock centra; SAM P35429 associated mid-posterior trunk vertebral centra; (Table 1), two associated sections of eight and SAM P35426 (Table 1), two articulated anterior four articulated anterior? tail stock centra; SAM FIGURE 2, SAM P35430, cf. Platpterygius longmani caudal vertebra in (A) anterior, (B) lateral, (C) ventral and (D) dorsal views. Scale bar is 20 mm. 36 B. P. KEAR TABLE 1. Measurements (mm) of cf. Platypterygius longmani vertebral centra from the Darwin Formation, Northern Territory. Specimen Length Width Height SAM P35039 39.6 91.2 98.9 33.1 88.9 90.2 30.2 80.1 89.2 32.3 62.6 66.1 SAM P35426 40.6 85.5 88.9 37.0 86.1 89.7 SAM P35427 40.7 94.0 96.4 30.1 86.2 92.4 25.8 102.8 109.6 *SAM P35429 24.9 81.6 69.9 25.8 83.1 75.2 26.4 86.4 79.1 25.2 64.8 83.9 20.5 - 78.2 21.6 58.3 60.1 21.1 58.9 50.8 22.3 - 57.2 17.4 - 55.1 18.9 - 46.6 14.1 - 46.9 SAM P35430 20.8 68.9 59.7 20.8 61.1 62.1 18.7 55.7 56.1 14.6 53.2 50.4 15.3 47.9 48.3 15.3 42.8 42.6 15.2 43.6 45.0 12.9 41.5 41.3 * some centra represented by fragments only P35430 (Figs 2A, B, C, D; Table 1), nine associated anterior tail stock centra (juvenile). Locality Casuarina Beach, north of Darwin, Northern Territory. Darwin Formation, Aptian or Albian. Description All of the preserved centra are disc-like and anteroposteriorly compressed, with the dorsal length subequal to the ventral. Tail stock centra (Figs 2A, B, C, D) exhibit a more marked degree of anteroposterior compression relative to those from the trunk region (Figs 1A, B) and are slightly ovoid in outline, with height being approximately 1.5 times the transverse width. The articular surfaces of all centra are deeply amphicoelous, with a distinct central notochordal pit. The neural canal is generally broad and shallow, becoming narrower in centra from the tail stock region. The neural arch facets are narrow and rectangular in all centra. Where preserved, both the diapophyses and parapophyses of the trunk centra are distinctly raised and subcircular. This is unlike centra from the tail stock region, in which only the ovoid diapophysis is present and positioned low on the lateral centrum surface. The ventral surfaces of at least two of the recovered centra from the tail stock region show distinct haemal arch facets. These are slightly concave and saddle-shaped in outline with low ridge-like margins. Medially, the haemal arch facets border the very weakly concave, spool-shaped depression for the haemal canal. Remarks All of the ichthyosaur vertebral centra examined can be distinguished from those of basal ichthyopterygians by their disc-like shape and strong anteroposterior compression (the more plesiomorphic Grippidia having cylindrical centra, Montani 1999). The absence of any further diagnostic features restricts attribution of the specimens to Ichthyosauria gen. et sp. indet. However, tentative assignment of the remains to cf. Platypterygius longmani is justifiable on the basis of: 1) overall similarity to P. longmani vertebral material described from the Allaru Mudstone and Toolebuc Formation, Queensland (Wade 1990) and Darwin Formation (Murray 1985, 1987 sensu Wade 1990); and 2) current recognition of P. longmani as the only valid ichthyosaur species from the Aptian—Albian of Australia (Wade 1990; Bardet et al. 1994; Arkhangelsky 1998; Montani 1999). Buchholtz (2001) discussed differentiation of structural units within the vertebral column of Jurassic ichthyosaurs with relation to variation in swimming styles between taxa. Following on from this study, it can be suggested that differences observed in the Darwin Formation centra (relating to regionalised morphological variation along the vertebral column) can be used to predict preferred swimming style in the Cretaceous P. longmani. The marked increase in height proportionate to width, and anteroposterior compression of the tail stock relative to trunk centra, in the Darwin Formation specimens is comparable to the condition recorded in derived thunniform-model DARWIN FORMATION MARINE REPTILE REMAINS 37 ichthyosaurs such as Ophthalmosaurus icenicus and Stenopterygius quadrissicus (Buchholtz 2001). Wade (1990) and Broili (1907) noted that the anterior fluke centra in P. longmani and P. platydactylus were distinctly higher than wide, a feature also conforming to the predicted thunniform condition. Taken together therefore, these characteristics may suggest that Platypterygius spp. employed an axial oscillatory swimming mode and probably shared many other key traits (see Montani 1999; Buchholtz 2001) with derived thunniform-model ichthyosaurs. SAUROPTERYGIA Owen, 1860 PLESIOSAURIA de Blainville, 1835 PLESIOSAUROIDEA (Gray, 1825) Welles, 1943 Plesiosauroidea gen. et sp. indet. Material SAM P35431 (Figs 3A, B, C), incomplete and badly weathered element tentatively identified as the ventral plate portion of a left ischium. Orientation is based on position of the lateral margin and its attitude relative to the horizontal axis; SAM P35434 (Figs 4A, B, C), right ischium lacking part of anteromedial margin. Locality Casuarina Beach, north of Darwin, Northern Territory. Darwin Formation, Aptian or Albian. Description Dimensions of remains are: SAM P35431, total length of 96.7 mm; total width is 98.9 mm; maximum height is lateral extremity is 22.5 mm; maximum length of lateral extremity is 55.3 mm; maximum height of ventral plate is 45.9 mm. SAM P35434, total length is 74.6 mm; total width FIGURE 3. SAM P35431, Plesiosauroidea gen. et sp. indet. partial ischium in (A) dorsal, (B) medial and (C) posterior views. Scale bar is 40 mm. 38 B. P. KEAR FIGURE 4. SAM P35434, Plesiosauroidea gen. et sp. indet. ischium in (A) dorsal, (B) posterior and (C) lateral views. Scale bar is 40 mm. is 97.4mm; maximum height of the of lateral articular surface is 52.1 mm; maximum length of lateral articular surface is 46.1 mm; maximum height of ventral plate is 10.8 mm. Both SAM P35431 and SAM P35434 are hatchet-shaped in dorsal outline with an anteroposteriorly expanded ventral plate. The lateral articular extremity of SAM P35434 is massive but has been lost in SAM P35431. Medially, the ventral plate portion of both elements is dorsoventrally compressed with an inclined medial edge for contact with the opposing ischium. This surface is oriented approximately 10° ventral to a horizontal plane through the lateral articular extremity, and is deeply pitted, suggesting the presence of cartilage. Both the dorsal and ventral surfaces of the ventral plate in SAM P35434 are rugose, possibly for attachment of the m. puboischiofemoralis internus and m. p. externus, respectively. A weak ridge on the posteromedial margin of the ventral plate may also have supported slips of the m. ischiotrochantericus. The ventral plate of both specimens is not greatly anteroposteriorly expanded (with total length being just slightly less than the total transverse width of each element) and is separated from the lateral articular extremity by a constricted waist-like midsection. This is ovoid to circular in cross-section, becoming increasingly dorsoventrally thickened towards the lateral articular surface (not preserved in SAM P35431). The articular surface itself is convex, with the rectangular anterior pubis facet DARWIN FORMATION MARINE REPTILE REMAINS 39 offset approximately 195° relative to the longitudinal axis of the articular surface. The pubis facet is separated posteriorly from the rectangular median acetabular facet, and in turn from the lobate posterior-most ilial facet (offset approximately 150° relative to the longitudinal axis of the articular surface) by weak parallel ridges. The articular surface margin is produced into a crennate rim, which is continuous ventrally, with a low rugose tuberosity possibly marking part of the m. puboischiofemoralis internus attachment. Remarks Despite being heavily weathered, SAM P35431 is identified as the ventral plate portion of a plesiosaurian left ischium on the basis of its hatchet-shaped outline and general morphology. Nevertheless, the specimen is unusual in its markedly dorsoventrally deep, medial articular margin. SAM P35431 was initially registered as an ichthyosaurian humerus belonging to Platypterygius sp. However, such an identification is unlikely as the humeri of Platypterygius spp. are much more robust, with rectangular to spool-shaped dorsal outline and prominent ridge-like tuberosities on both the dorsal and ventral surfaces (eg Kiprijanoff 1881; Broili 1907; Kuhn 1946; McGowan 1972; Wade 1984; Murray 1987; Choo 1999). SAM P35431 also differs from the larger girdle elements of Platypterygius spp., in which the coracoid is subcircular in outline with dorsoventrally deep glenoid/intercoracoid facets (Broili 1907; Wade 1984), and both the scapula and ischiopubis are strap-like with predominantly narrow, ridge-like articular surfaces (Broili 1907; Nace 1939; Murray 1987). SAM P35434 probably represents a juvenile individual because of its small size and poor ossification of the articular surfaces. Both it and SAM P35431 can be assigned to Plesiosauroidea on the basis of their anteroposteriorly short ventral plates. This differs from the condition in pliosauroids in which the ventral plate of the ischium bears a marked posterior expansion (Mehl 1912; Andrews 1913; Tarlo 1959, 1960; Brown 1981). The presence of a relatively weak posterior expansion in Eurycleidus (Cruickshank 1994) and Leptocleidus (pers. obs, AM F99374 from the Aptian—Albian Bulldog Shale of Coober Pedy, South Australia), however, suggests that this character state may not be universal for the group. SAM P35434 is unusual in its proportionately massive articular head relative to the ventral plate. The phylogenetic significance of this feature, if any, is uncertain because of its immature stage of development. ELASMOSAURIDAE Cope, 1869 Elasmosauridae gen. et sp. indet. Material SAM P35432 (Figs SA, B, C, D), a single weathered posterior cervical vertebral centrum. Locality Casuarina Beach, north of Darwin, Northern Territory. Darwin Formation, Aptian or Albian. Description The surface of the centrum is badly weathered with very little periosteal bone remaining. Total length is 72.8mm; width of anterior surface is 109.5 mm; depth of anterior surface is 85.2 mm. In general proportions, the centrum is anteroposteriorly short relative to both the height and width of the anterior articular surface. Both the lateral and ventral surfaces are shallowly concave, with the ventral surface being pierced by two large foramina. The articular surfaces are poorly preserved but clearly platycoelous, with a raised convex rim. The neural arch facets are indistinct but prominent bosses high on the lateral surface of the centrum indicate positioning of the rib facets. Remarks SAM P35432 is tentatively assigned to Elasmosauridae on the basis of its platycoelous articular surfaces. Murray (1987) reported an elasmosaurid cervical with very shallowly concave articular facets from the Casuarina Beach locality. Brown (1981, 1993) and Bardet et al. (1999) regarded the presence of platycoelous articular surfaces on the cervical centra to be a potential synapomorphy for Elasmosauridae. The character state has also been recorded in the Maastrichtian cryptoclidid Morturneria (Chattergee & Small 1989), though Bardet et al. (1991) suggested that this taxon may represent a derived elasmosaurid. The proportionately short centrum length relative to height of SAM P35432 differs from the common condition in elasmosaurids (such as that described by Murray 1987), in which the cervical centra are markedly elongate (Brown 1993; Bardet et al. 1999). The presence of relatively short cervical centra is considered plesiomorphic among 40 B. P. KEAR FIGURE 5. SAM P35432, Elasmosauridae gen. et sp. indet. posterior cervical centrum in (A) dorsal, (B) ventral, (C) anterior and (D) lateral views. Scale bar is 40 mm. plesiosauroids (Brown 1993; Bardet ef al. 1999). However, the feature has been described in the potential elasmosaurid taxa (sensu Bardet et al. 1991) Aristonectes (Cabrera 1941) and Morturneria (Chattergee & Small 1989) as well as in indeterminate elasmosaurid posterior cervicals from unspecified (probably Albian) deposits near Oodnadatta (Freytag 1964) and the Neales River region (SAM P6181, Persson 1960), South Australia; Toolebuc Formation, Queensland (Kear 2001); and Molecap Greensand, Western Australia (Long & Cruickshank 1998). The presence of proportionately elongate cervicals may therefore not necessarily be universal amongst elasmosaurids, with some taxa exhibiting relatively short centra, particularly in the posterior cervical region. SAM P35432 is notable for its possession of a raised convex rim surrounding the central concavity of the articular surface. Brown et al. (1986) and Brown (1993) considered this a plesiomorphic condition commonly associated with members of the Cryptoclididae and Plesiosauridae. In contrast, elasmosaurids, including basal taxa such as Occitanosaurus (Bardet et al. 1999) and Muraenosaurus (Andrews 1910; Brown 1981, 1993), generally exhibit a more open V-shape, in which the articular surface margins form an abrupt angle with the centrum sides. A similar trait is evident in elasmosaurid material from the Toolebuc Formation of Queensland (Kear 2001), and is indicated (but not described) in the figured Darwin Formation specimen (NTM P8727-70, Fig. 1, p. 97) of Murray (1987). Carpenter (1996), however, considered articular rim morphology to be a variable character amongst plesiosaurs and of questionable diagnostic value. DARWIN FORMATION MARINE REPTILE REMAINS 41 PLIOSAUROIDEA Seeley, 1874 PLIOSAURIDAE Seeley, 1874 Leptocleidus Andrews, 1922 Leptocleidus sp. Material SAM P35053, cast of NTM P913-5 (Figs 6A, B, C) right femur. Locality Casuarina Beach, north of Darwin, Northern Territory. Darwin Formation, Aptian or Albian. Description A near complete right femur lacking part of posterodistal margin. Total length is 260.8 mm; FIGURE 6. SAM P35053 (cast of NTM P913-5), Leptocleidus sp. femur in (A) anterior, (B) dorsal and (C) proximal views. Scale bar is 40 mm. total proximal length is 47.9 mm; total proximal height (including trochanter) is 51.5 mm; width across base of dorsal trochanter is 37.2 mm; total distal length (not including missing posterior margin) is 91.6 mm; total distal height is 25.9 mm. The anterior edge of the femur is weakly sigmoidal, with a strongly convex distal extremity. The posterior edge is uniformly concave along its entire length. Both the dorsal and ventral surfaces are shallowly concave along their longitudinal axis; however, the ventral surface becomes convex and upturned towards the distal extremity. Proximally, the femur is greatly dorsoventrally expanded and separated from its distal section by a constricted neck. The dorsal trochanter is robust and anteroposteriorly constricted at its base. It is separated from the ovoid glenoid articular surface by a shallow groove. The surfaces of both the dorsal trochanter and glenoid articulation are deeply pitted, suggesting the presence of cartilage. The femoral shaft is weakly anteroposteriorly constricted and ovoid in cross-section, with its posterior margin bearing a raised rugose tuberosity, possibly for attachment of the m. caudifemoratis. A second low tuberosity is present on the median ventral surface and may represent part of the m. puboischiofemoralis insertion. The distal femoral extremity is fan-shaped in dorsoventral outline, with a weakly projecting anterodistal margin. The distal articular surface is narrow and elliptical, with a raised median ridge separating the very large radial facet from the remains of the ulnar facet. Remarks Despite the poor existing knowledge of femoral morphology in Leptocleidus spp., SAM P35053 (NTM P913-5) can be readily distinguished from the proportionately shorter and stouter propodials of elasmosaurids (Brown 1981; Murray 1987) by its anteroposteriorly broad, flat femoral shaft, greatly expanded distal extremity and sigmoidal anterior margin. These characteristics are very similar to those of the fragmentary femora of L. capensis (Andrews 1911) and well-preserved material (AM F99374, SAM P15980) from the Bulldog Shale of South Australia (Figs 7A, B). The femur of L. clemai (Cruickshank & Long 1997) differs in its possession of a concave anterior margin. This feature, along with a proportionately more slender shaft, is also present in the femora of polycotylids and most other pliosauroids (Andrews 1913; Welles 1943, 1962; Brown 1981; Storrs 1999). A sigmoidal anterior femoral and humeral margin has, however, been 42 B. P. KEAR TABLE 2. Marine reptile groups from Early Cretaceous deltaic-marine deposits of Australia. Pliosauroids are separated into small-bodied (eg Leptocleidus) and large-bodied (eg Kronosaurus) forms because of their differing ecological roles. The poorly known taxon ‘Cimoliasaurus maccoyi’ is recorded separately because of its unknown affinity. Indeterminate plesiosaur material from the marine Bungil Formation, Queensland and freshwater deposits of the Griman Creek Formation, New South Wales, and Strzeleki and Otway Groups, Victoria is excluded pending a more thorough examination of the remains. Taxa from the opal bearing deposits of White Cliffs, New South Wales are placed within the Wallumbilla Formation in accordance with the lithostratigraphic nomenclature of Burton & Mason (1998). Source texts include: Etheridge (1904)', Longman (1924), Tierchert & Matheson (1944)3, Persson (19604, 19635, 1982°), Ludbrook (1966)’, Condon (1968)’, Pledge (1980), Smart & Senior (1980)!, Gaffney (1981)"', Molnar (1982!7, 199119), Wade (1984"4, 1990!5), Murray (1985!°, 19877), Hocking et al. (1987)!8, Burger (1988)!°, Bardet (1992), Thulborn & Turner (1993)?!, Krieg & Rodgers (1995), Cruickshank & Long (1997)3, Henderson (1998)4, Long (1998), Long & Cruickshank (1998), Cruickshank et al. (1999)?’, Choo (1999)*8, Kear (2001). Darwin Formation Wallumbilla Formation (Money Shoals Platform) Aptian/Albian (Eromanga Basin) Aptian—Albian Ichthyosauria*!3!5.16.17 Ichthyosauria!3 Elasmosauridae™3!” Elasmosauridae**!2:!325 Small pliosaurid”’ Large pliosaurid !245:12.1321.25 Polycotylidae**3 *C. maccoyi’'43 Shallow near-shore Coastal / offshore shallow marine™ / paralic!? / marine!®!9 possibly tidal'® cited as a potentially diagnostic character state for the polycotylid genus Polycotylus (Storrs 1999). DIscussION The Darwin Formation sediments have been interpreted as representing a shallow near-shore marine/paralic (Smart & Senior 1980; Henderson 1998) to possibly tidal (Murray 1985) depositional environment. The presence of the small pliosaurid taxon Leptocleidus, known elsewhere from freshwater and near-shore marine deposits (Andrews 1911, 1922; Str6mer 1935; Cruickshank 1997; Cruickshank & Long 1997; Cruickshank et al. 1999), is consistent with this interpretation. A comparison (Table 2) of the Darwin Formation marine reptile fauna with those known from Early Cretaceous deltaic-marine deposits elsewhere in Australia (Wallumbilla Formation [Doncaster Member], Allaru Mudstone, Toolebuc Formation and Mackunda Formation, Queensland; Bulldog Shale, South Australia; Wallumbilla Formation [Doncaster Member], New South Wales; Birdrong Sandstone and Barrow Group, Allaru Mudstone (Eromanga Basin) Albian Toolebuc Formation (Eromanga Basin) Albian Ichthyosauria!*:!3-14.15.25 Elasmosauridae**!?:13.25 Chelonioidea’3*5 Ichthyosauria!2!314.15.25 Elasmo sauridae*>®! 2,13,21,25,29 Large pliosaurid?*5!2.3 Polycotylidae?!?527 Chelonioidea!!:!2.13.25 Shallow marine!” Shallow marine’® Western Australia) indicates greatest compositional similarity to the Hauterivian— Barremian Birdrong Sandstone fauna, with ichthyosaurs, elasmosaurids and the small-bodied pliosaurid Leptocleidus represented. Throughout the Early Cretaceous marine deposits, ichthyosaurs show a wide distribution (occurring in all currently recognised deposits except the Albian Mackunda Formation and unspecified Berriasian subsurface sediments of the Barrow Group), as do elasmosaurids, which have been recorded from all localities except those of the Albian Mackunda Formation and Berriasian Barrow Group. Cruickshank et al. (1999) indicated the possible presence of cryptoclidids in the Aptian—Albian Bulldog Shale. This is supported by more recent observations of the South Australian fauna (Kear unpubl.) but as yet the group has not been recorded from any other Australian deposit. The distribution of small- and large-bodied pliosaurid taxa in the Australian Early Cretaceous is variable, with smaller forms (represented by at least two species of Leptocleidus, Cruickshank et al. 1999) confined to the Darwin Formation, TABLE 2. (cont.) Mackunda Formation (Eromanga Basin) Albian Polycotylidae’ DARWIN FORMATION MARINE REPTILE REMAINS Bulldog Shale (Eromanga Basin) Aptian—A Ibian Birdrong Sandstone (Carnarvon Basin) Hauterivian—Barremian Ichthyosauria®'3° Ichthyosauria*!378 Barrow Group (Carnarvon Basin) Berriasian Small pliosaurid?¢ 43 Elasmosauridae®”’ Small pliosaurid®?5?7 Large pliosaurid’ Cryptoclididae?”’ Paralic’® Shallow marine? / offshore’ Bulldog Shale and Birdrong Sandstone. Larger pliosaurid remains occur only in the Aptian— Albian Wallumbilla Formation (Doncaster Member), Albian Toolebuc Formation and Bulldog Shale deposits. This division may be linked to the different habitat preferences and ecological roles of small- and large-bodied pliosaurid taxa, with offshore environments preferentially supporting larger-bodied forms. An exception is seen in the shallow offshore marine (Ludbrook 1966; Krieg & Rogers 1995) Bulldog Shale, which includes both small- and large- bodied pliosaurid remains (the latter being represented by an isolated tooth, SAM P22525, from the opal-bearing sediments of Coober Pedy, figured by Pledge 1980, p. 8). Distribution of polycotylids in the shallow marine (Smart & Senior 1980; Burger 1988) Wallumbilla Formation, Toolebuc Formation and paralic (Smart & Senior 1980) Mackunda Formation may, as with large-bodied pliosaurids, be related to a preference for predominantly offshore marine environments. However, individuals also appear to have readily Elasmosauridae”©?? Small pliosaurid?3?? Coastal near-shore Fluviatile-deltaic'® shallow marine!®/ paralic® frequented more inshore habitats, as suggested by their occurrence in the Mackunda Formation deposits. Restriction of chelonioids to the upper Albian Allaru Mudstone and Toolebuc Formation may be the product of both the group’s temporal distribution (the earliest-known chelonioid being recorded from the upper Aptian — lower Albian Santana Formation of Brazil, Hirayama 1997, 1998) as well as environmental factors such as prevailing water temperatures. Studies of sedimentary sequences (Frakes & Francis 1988; Frakes & Krassay 1992; Frakes et al. 1995; Constantine et al. 1998), climatic modelling (Barron & Washington 1982) and isotope data (Gregory et al. 1989; Pirrie et al. 1995) have suggested that strongly seasonal climates with winter freezing and at least seasonal sea ice characterised high latitudes in the Early Cretaceous of Australia. Such conditions may well have been unfavourable to chelonioids and provided an effective barrier to the group’s dispersal into the Australian region prior to the upper Albian. 44 B. P. KEAR FIGURE 7. SAM P15980 juvenile Leptocleidus sp. humerus (A) and femur (B) from Andamooka, South Australia. Scale bar is 20 mm. ACKNOWLEDGMENTS I wish to thank Dirk Megirian of the Museum and Art Gallery of the Northern Territory for assistance with locating literature on the Darwin Formation deposits, and generous provision of NTM P913-S for study. Tim Flannery, Neville Pledge, Jo Bain and Ben Head of the South Australian Museum provided support, enthusiasm, preparation facilities and materials. This manuscript benefited greatly from the comments of Dirk Megirian and Arthur Cruickshank of the Leicester City Museums Service, New Walk Museum. Origin Energy, The Advertiser, the Waterhouse Club and the Coober Pedy Tourism Association provided financial support for this research, REFERENCES ANDREWS, C. W. 1910. ‘A Descriptive Catalogue of the Marine Reptiles of the Oxford Clay’. Vol. 1. British Museum (Natural History): London. ANDREWS, C. W. 1911. 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TENTACULARIID CESTODES OF THE ORDER TRYPANORHNCHA (PLATYHELMINTHES) FROM THE AUSTRALIAN REGION H. W. PALM & 1. BEVERIDGE Summary The present study summarises information on tentaculariid trypanorhynchs from Australian waters. A total of 19 species from the genera Nybelina Poche, 1926, Heteronybelinia Palm, 1999, Mixonybelinia Palm, 1999, Kotorella Euzet & Radujkovic, 1989 and Kotorelliella gen. nov. were identified: N. aequidentata (Shipley & Hornell, 1906); Nybelina africana Dollfus, 1960; N. hemipristis sp. nov.; N. jayapaulazariahi Reimer, 1980; N. mehlhorni sp. nov.; N. schmidti Palm, 1999; N. scoliodoni (Vijayalakshmi, Vijayalakshmi & Gangadharam, 1996); N. strongyla Dollfus, 1960; N. thyrsites Korotaeva, 1971; N. victoriae sp. nov.; Heteronybelinia australis sp. nov.; H. estigmena (Dollfus, 1960); H. pseudorobusta sp. nov.; Mixonybelinia beveridgei (palm, Walter, Schwerdtfeger & Reimer, 1997); M. cribbi sp. nov.; M. edwinlintoni (Dollfus, 1960); M. southwelli Palm & Walter, 1999; Kotorella pronosoma (Stossich, 1901) and Kotorelliella jonesi gen. et sp. nov. The new genus Kotorelliella is characterised by a homeocanthous, heteromorphous metabasal armature and a basal armature with additional interpolated hooks on the external surface of the tentacle, thus appearing heterocanthous. The new species appears to be closely related to Kotorella pronosoma. The tentaculariid trypanorhynch fauna in Australian waters is pecies rich, with 22 (48%) of the total of 46 known species occurring in waters of the region. Eleven new locality, and 20 new host records are reported. TENTACULARIID CESTODES OF THE ORDER TRYPANORHYNCHA (PLATYHELMINTHES) FROM THE AUSTRALIAN REGION H. W. PALM & I. BEVERIDGE PALM, H. W. & BEVERIDGE, I. 2002. Tentaculariid cestodes of the order Trypanorhyncha (Platyhelminthes) from the Australian region. Records of the South Australian Museum 35(1): 49-78. The present study summarises information on tentaculariid trypanorhynchs from Australian waters. A total of 19 species from the genera Nybelinia Poche, 1926, Heteronybelinia Palm, 1999, Mixonybelinia Palm, 1999, Kotorella Euzet & Radujkovic, 1989 and Kotorelliella gen. nov. were identified: N. aequidentata (Shipley & Hornell, 1906); Nybelinia africana Dollfus, 1960; N. hemipristis sp. nov.; N. jayapaulazariahi Reimer, 1980; N. mehlhorni sp. nov.; N. schmidti Palm, 1999; N. scoliodoni (Vijayalakshmi, Vijayalakshmi & Gangadharam, 1996); N. strongyla Dollfus, 1960; N. thyrsites Korotaeva, 1971; N. victoriae sp. nov.; Heteronybelinia australis sp. nov.; H. estigmena (Dollfus, 1960); H. pseudorobusta sp. nov.; Mixonybelinia beveridgei (Palm, Walter, Schwerdtfeger & Reimer, 1997); M. cribbi sp. nov.; M. edwinlintoni (Dollfus, 1960); M. southwelli Palm & Walter, 1999; Kotorella pronosoma (Stossich, 1901) and Kotorelliella jonesi gen. et sp. nov. The new genus Kotorelliella is characterised by a homeoacanthous, heteromorphous metabasal armature and a basal armature with additional interpolated hooks on the external surface of the tentacle, thus appearing heteroacanthous, The new species appears to be closely related to Kotorella pronosoma. The tentaculariid trypanorhynch fauna in Australian waters is species rich, with 22 (48%) of the total of 46 known species occurring in waters of the region. Eleven new locality, and 20 new host records are reported. H, W. Palm, Institut fiir Zoomorphologie, Zellbiologie und Parasitologie, Heinrich-Heine- Universitat Diisseldorf, UniversitatsstraBe 1, D-40225 Diisseldorf, Germany, current address: Centre for Coastal and Marine Resource Studies, Faculty of Fisheries and Marine Sciences, Bogor Agricultural University, 16680 Bogor, Indonesia; email: hpalm@indo.net.id, hpalm@ gmx.net; and I. Beveridge, Department of Veterinary Science, University of Melbourne, Parkville, Victoria 3052, Australia. Manuscript received 20 November 2000. Palm (1999), Palm et al. (1997), Palm and Overstreet (2000) and Palm and Walter (1999, 2000) partially revised the trypanorhynch cestode family Tentaculariidae Poche, 1926 on the basis of material deposited in museums in London, Paris, Vienna and the United States. Following the erection of two new genera, Heteronybelinia Palm, 1999 and Mixonybelinia Palm, 1999 by Palm (1999), Palm and Walter (2000) gave a summary of the current state of knowledge within the tentaculariid trypanorhynch genera. Thus, to date, a total of 39 species of Kotorella, Nybelinia, Heteronybelinia and Mixonybelinia are considered valid (Palm & Walter 2000). There is littke known about the tentaculariid trypanorhynchs from Australian coastal waters. The first collections were made by a French scientific expedition under the command of Nicolas Baudin between 1801 and 1803 (Beveridge & Campbell 1996), but the cestodes, Tentacularia coryphaenae Bosc, 1797 and an unidentified species of Nybelinia were not described until 1942 (Dollfus 1942). Korotaeva (1971) named the Nybelinia species of Dollfus (1942) N. thyrsites and subsequently (Korotaeva 1974) reported additional unidentified Nybelinia spp. from other fishes from southern Australia. Lester et al. (1988) and Sewell and Lester (1988) reported Tentacularia sp. from orange roughy, Hoplostethus atlanticus Collett, 1889 off the coasts of South Australia and Tasmania; while Sewell and Lester (1995) found Nybelinia sp. in jewfish, Rexea solandri (Cuvier & Valenciennes, 1832) from southern Australian waters. Beveridge and Campbell (1996) recorded Tentacularia coryphaenae Bosc, 1797, Nybelinia thyrsites Korotaeva, 1971 and N. sphyrnae Yamaguti, 1952 from Australian fishes; while Jones and Beveridge (1998) added N. queenslandensis from a carcharhinid shark as a new species. Palm (1999) recorded Nybelinia lingualis (Cuvier, 1817) from southern Australian waters and Speare (1999) 50 H. W. PALM & I. BEVERIDGE reported Nybelinia sp. from sailfish, [stiophorus platypterus (Shaw & Nodder, 1792) off the coast of Queensland. Thus, a total of five tentaculariid species has been described from Australia. Last and Stevens (1994) summarised knowledge of the Australian chondrichthyan fauna, the definitive hosts of trypanorhynchan cestodes, as being extremely rich and consisting of 296 species, 54% of them endemic. In terms of host-specificity and zoogeography' of trypanorhynch cestodes, the Australian fauna might therefore harbour a high number of endemic and locally distributed tentaculariid trypanorhynchs, such as was observed within the related family Eutetrarhynchidae by Beveridge (1990). Palm (1999) and Palm and Walter (2000) suggested that many tentaculariids, for example Tentacularia coryphaenae and Nybelinia lingualis, exhibit a cosmopolitan or circumtropical distribution pattern. However, records from the south-west Pacific and south-east Indian Ocean are scarce. For example, the tentaculariids Tentacularia coryphaenae, Nybelinia africana and N. scoliodoni were only recently recorded from Indonesian coastal waters (Palm 2000). Other records which might confirm a wide distribution and possibly a low degree of host specificity for these trypanorhynchs are still lacking. The present study was carried out to examine the species of Nybelinia, Heteronybelina, Mixonybelinia and Kotorella deposited in museum collections in Australia as well as in the collection of the junior author. Where necessary, the specimens are described and illustrated as emendations to currently available descriptions. Apart from the establishment of new host and locality records, species identifications provide further insight into the zoogeographical distribution of tentaculariid cestodes. The comparison of the specimens with earlier descriptions allows comments to be made on the extent of intraspecific morphological variability within tentaculariid trypanorhynchs. Together with the study of deposited tentaculariids from other collections (Palm 1999; Palm & Overstreet 2000; Palm and Walter 1999, 2000), the present study summarises the current state of knowledge on the genera of tentaculariid cestodes. MATERIAL AND METHODS Specimens were examined from the Australian Helminthological Collection (AHC) housed in the South Australian Museum (SAM), Adelaide, as well as the Queensland Museum (QM), Brisbane. Special attention was given to unidentified specimens deposited simply as Nybelinia sp. Unmounted specimens were stained in Celestine Blue, dehydrated in ethanol, cleared in methyl salicylate and mounted in Canada balsam. Pieces of strobilae were embedded in paraffin, serial sections were cut at a thickness of 5S fim and stained with haematoxylin and eosin. Drawings were made using an Olympus BH2 microscope with an ocular micrometer and drawing tube. Terminology for morphological features peculiar to the Trypanorhyncha follows Dollfus (1942) and Campbell and Beveridge (1994). The following measurements were made: scolex length (SL), scolex width at level of pars bothridialis (SW), pars bothridialis (pbo), pars vaginalis (pv), pars bulbosa (pb), pars postbulbosa (ppb), velum (vel), appendix (app), bulb length (BL), bulb width (BW), bulb ratio (BR), scolex proportions of pbo:pv:pb (SP), tentacle width (TW) and tentacle sheath width (TSW). If possible, the tentacle length (TL) was estimated. The tentacular armature was described as follows: armature homeomorphous or heteromorphous, number of hooks per half spiral row (hsr), total hook length (L) and the total length of the base of the hooks (B). The abbreviation ‘nm’ (not measured) indicates that no measurement was taken. All measurements are given in micrometers unless otherwise indicated. Specimens belonging to the same species from different hosts or localities were measured in the same order as cited in the material examined. If more than three measurements were taken from a single host species, the mean is given with the range in parentheses. Because tentaculariid cestodes are morphologically uniform and the genera are well defined, the species descriptions presented here are restricted to key differential features. Illustrations are provided if useful for future species identification; otherwise the reader is referred to illustrations given by other authors. The classification utilised follows Palm (1995, 1997). Host identity follows Last and Stevens (1994) for the elasmobranchs and FishBase 1998 (Froese & Pauly 1998) for the teleosts. SYSTEMATICS A total of 19 species were identified, and 11 new locality and 20 new host records were established. Seven new species are described and TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 51 a new tentaculariid genus is erected: Nybelinia hemipristis sp. nov., N. mehlhorni sp. nov., N. victoriae sp. nov., Heteronybelinia australis sp. nov., H. pseudorobusta sp. nov., Mixonybelinia cribbi sp. nov. and Kotorelliella jonesi gen. et sp. nov. Detailed information on individual specimens with comments on their taxonomy and distribution are provided below. Order TRYPANORHYNCHA Diesing, 1863 Superfamily TENTACULARIOIDEA Poche, 1926 Family TENTACULARIIDAE Poche, 1926 Genus Nybelinia Poche, 1926 Nybelinia aequidentata Shipley & Hornell, 1906 (Figs 1-2) Material examined From Dendrochirus zebra (Cuvier, 1829): 1 postlarva, Noumea, New Caledonia, Nov. 1997, coll. S. Pichelin (QM G 218031). Description SL=7900 (Fig. 1); SW =1775; pbo = 2975; pv = 4950; pb = 1575; vel = 925; app = 1400; BL = 1417 (1400-1450); BW = 280 (270-290); BR =5.1:1; SP = 1.9:3.1:1. Basal tentacular swelling absent. TW basal = 100-110, TW metabasal = 110-120. Tentacle sheaths straight, TSW = 80-100, prebulbar organs and muscular rings around basal part of tentacle sheaths not visible. Retractor muscles originate in basal parts of bulbs. Tentacular armature homeoacanthous, homeomorphous (Fig. 2); distinctive basal armature absent. Hooks falciform, with slender shaft, stout base and strongly recurved tip, L = 57.5-65.0; B = 17.7—22.5. Hooks decrease in size towards apical region and towards base L = 35.0-57.5; B = 12.5-17.5. Smallest hook measured L = 17.5; hsr = 7-8. Remarks The present specimen has one of the largest scoleces among Nybelinia species. In addition, the postlarva is characterised by falciform hooks over 60 in length and a bulb ratio greater than 4. The postlarva is most similar to Nybelinia aequidentata Shipley & Hornell, 1906, described by Shipley and Hornell (1906), Pintner (1927) and Palm (1999). Shipley and Hornell (1906) recorded the type specimen as having a scolex 4-5 mm long. The hook shape of the present specimen corresponds with those of the type, though the hooks in the present specimen are larger (57-65 compared with 49 in the type according to Shipley and Hornell (1906) or 33-38 reported by Palm (1999). Pintner (1927) redescribed the type and added information on the bulb ratio (4.3:1), hook length (48) and scolex measurements (SW = 1900, pbo = 1500, pb = 1600 and vel = 600). He also remarked that the hooks were not uniform in size but increased from the base towards the metabasal region and then decreased in the apical region of the armature. Thus, although the scolex and hook sizes of the present specimen are distinctly larger than those described from the type specimen, it has been identified as N. aequidentata. The present finding represents a new host and locality record. Nybelinia syngenes Pintner, 1929 is the only other species of Nybelinia which has been described as having tentacular hooks with a length greater than 60 and with a similar hook form and armature pattern. However, the scolex of this species, described originally from Sphyrna zygaena (Linnaeus, 1758), is distinctly smaller. Whether N. aequidentata exhibits variation in scolex size, as described for N. lingualis (Cuvier, 1817) and Heteronybelinia yamaguti (Dollfus, 1960) (see Dollfus 1942; Palm et al. 1997; Palm 1999; Palm & Walter 2000) is not known. Palm (1999) has suggested that this might be the case for a postlarva from Lepturacanthus savala (Cuvier, 1829) with a scolex length of 3400, a bulb ratio of 3.3:1 and slender falciform hooks 33-38 in length. Nybelinia africana Dollfus, 1960 (Figs 3-5) Material examined From unidentified shark: 1 adult, Queensland, coll. J. C. Pearson (QM G 218169). Description SL = 1780 (Fig. 3); SW = 1134; pbo = 930; pv = 840; pb = 523; vel =417; BL = 439 (423- 459); BW=163 (157-171); BR=2.7:1; SP = 1.8:1.6:1. TW metabasal = 47-50. Basal tentacular swelling absent. Tentacle sheaths spirally coiled; TSW = 65-70. Prebulbar organs absent, muscular rings around basal part of tentacle sheaths not seen. Retractor muscles 52 ‘ H. W. PALM & I. BEVERIDGE FIGURES 1-2. Nybelinia aequidentata Shipley & Hornell, 1906 from Dendrochirus zebrae. QM G 218031. 1. Scolex. 2. Homeomorphous armature with falciform hooks. Scale bars: Figure 1, 500 pm; Figure 2, 100 pm. FIGURES 3-5. Nybelinia africana Dollfus, 1960 from an unidentified shark. QM G 218169. 3. Scolex. 4. Basal and metabasal armature. 5. Mature proglottid. Scale bars: Figure 3, 200 um; Figure 4, 20 um; Figure 5, 100 pm. TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 53 originate in basal parts of bulbs. Tentacular armature consists of homeomorphous hooks along tentacle (Fig. 4). Tentacles not completely evaginated; metabasal armature differs distinctly from basal armature. Basal 2—3 rows of rosethorn- shaped hooks with distinct anterior extension of base, L = 14-16, 16-17; metabasal hooks larger, falciform with small base, strongly recurved at tip, L = 26-27; B = 10-13. Strobila 57 mm long, with about 215 acraspedote segments which enlarge in size towards end of strobila, maximum width 1.48 mm; velum straight or very slightly scalloped; first segments 15-30 x 600-675, mature segments (Fig. 5) wider than long, 220-470 x 650-970 (n=4). Pre-gravid segments (with some eggs) 440-670 x 1150-1480 (n = 5); terminal pre-gravid segment with rounded end. In mature segments, genital pore ventro-submarginal, in anterior half or anterior third of segment; genital pore to anterior end 90-150 in mature and 180-200 in pre-gravid segments; genital pores alternate irregularly. Cirrus sac thin-walled, elongate, 370— 480 x 70-100 (n=5), length:width ratio 4.5:1 (3.7-5.0:1), distal pole directed anteromedially, nearly reaching anterior end of proglottid, cirrus unarmed; internal and external seminal vesicle absent. Vas deferens greatly coiled, extends to midline of segment, then posteriorly towards female genital complex. Testes of varying shape, 60-90 in diameter, arranged in single layer centrally and in double layer peripherally; testis number 50-71 per segment, between 5 and 8 testes anterior to cirrus (n= 5). Ovary follicular, in centre of segment, bilobed, each lobe 190-260 x 140-220, increasing in size along strobila. Vitelline follicles encircle medulla, follicles 30- 50 in diameter. Ventral and dorsal osmoregulatory canals 30 in diameter, testes extend external to ventral canal but not beyond dorsal canal. Remarks The present specimen closely resembles N. africana Dollfus, 1960 as described by Dollfus (1960) and Palm (1999) in having rosethorn-shaped basal hooks, changing to falciform metabasal hooks of similar size and shape. Additionally, the segment morphology with number and size of segments, size of cirrus sac, testes and vitelline follicles, as well as arrangement of the genital complex is similar. However, some differences were observed between the present specimen and WN. africana. The specimen from Australia differs in having 2 to 3 rows of basal hooks with a distinct anterior extension of the base (Fig. 4), which has not been reported in other specimens of N. africana. However, Dollfus (1960, figs 14, 17) illustrated comparable basal hooks, some with an anterior extension of the base. The scolex is larger in the Australian specimen (1780) compared with other material (1118-1568) (Palm et al. 1997) and the testis number is smaller. However, as most other morphological characters correspond, the specimen is identified as N. africana and the observed differences are considered to be due to intraspecific morphological variability. The present finding is a new locality record for the species. Nybelinia hemipristis sp. nov. (Figs 6-9) Types Holotype from stomach of Hemipristis elongata Klunzinger, 1871, Balgal, Queensland, coll. B. G. Robertson, 16.ix.1985 (SAM AHC 28309). Paratypes: 3 specimens, Hemipristis elongatus, Marchinbar Island, Northern Territory, coll. G. Cuthbertson, 25.v.1985 (SAM AHC 28310). Material examined Types. Description SL = 2030 (Fig. 6); SW = 1230; pbo = 1000; pv = 860; pb = 560; vel = 630; BL=553 (550- 560); BW=218 (210-220); BR = 2.5:1; SP = 1.8:1.5:1. Tentacles short, massive, decreasing in diameter towards apical region; basal tentacular swelling absent. TW basal and metabasal = 75-85, TW apical = 37.5-62.5. Tentacle sheaths straight (TSW = 50-60), prebulbar organs and muscular rings around basal part of tentacle sheaths not seen. Retractor muscles originate in basal parts of bulbs. Tentacular armature homeoacanthous, homeomorphous (Fig. 7); distinctive basal armature absent (Fig. 8). Hooks slender, rosethorn-shaped with anterior elongation of base, increasing in size from basal (L = 15.5-30.0; B = 16.3-20.0) towards metabasal regions (L = 35.0-40.0; B = 25.0-27.5), then decreasing in size towards apical (L = 27.5-32.5; B = 15.0— 20.0) region; hsr = 6. Strobila acraspedote, velum straight; 33 mm long, maximum width 1030, number of segments 300. Mature segments (Fig. 9) wider than long, 600-670 x 750-920. Genital pore submarginal, 200 from anterior end, alternating irregularly. Cirrus sac short and stout, 180-210 x 50-100 (n=5S), length:width ratio 2.9:1 (2.0-3.6:1), distal pole directed 54 H. W. PALM & I. BEVERIDGE SAN ae fe weay a Les Boy < es FIGURES 6-9. Nybelinia hemipristis sp. nov. from Hemipristis elongata. Holotype, SAM AHC 28309. 6. Scolex. 7. Metabasal armature. 8. Basal armature. 9. Mature proglottid. Note short cirrus sac, the uterine duct (ud), and the Mehlis’ gland (mg). Scale bars: Figure 6, 200 um; Figures 7-8, 40 um; Figure 9, 100 pm. FIGURES 10-11. Nybelinia jayapaulazariahi Reimer, 1980 from a ‘sole’. QM G 207318. 10. Metabasal armature. 11. Basal armature. Scale bars: Figures 10-11, 10 um. TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 55 anteromedially, not reaching anterior end of segments, cirrus unarmed; internal and external seminal vesicle absent. Vas deferens coiled, extends to midline of segment, then posteriorly towards female genital complex. Testes 84-97 in number, 55-85 in diameter centrally and 40-60 peripherally, distributed in single layer; arranged in 2 lateral groups, confluent posterior to ovary; extend between ovarian lobes, about 6—7 testes anterior to cirrus sac. Ovary in centre of segment, ovarian lobes 200-260 x 120-160. Vagina ventral to cirrus sac. Vitelline follicles encircle medulla; follicles 2540 in diameter. Remarks The present specimens, with rosethorn-shaped hooks diminishing in size towards the apical armature and a strobila consisting of acraspedote segments, resemble N. anthicosum Heinz & Dailey, 1974. However, the species differ in having different scolex forms, tentacles, tentacular armatures and sizes. N. anthicosum has an elongated prominent velum, long tentacles and tentacular hooks which are distinctly spaced (see Heinz & Dailey 1974). The largest hooks occur in the 7th to 9th rows. N. hemipristis sp. nov. has a massive scolex with a shorter velum (Fig. 6), short tentacles and tentacular hooks which are tightly spaced along the tentacle. The largest hooks are in the 10th to 14th rows. In the last few hook rows, the size and number of hooks per half spiral row decrease rapidly (Fig. 7). In N. anthicosum the hook size diminishes gradually towards the apical part of the tentacle. As with N. anthicosum, N. hemipristis is a species without a characteristic basal armature and with a smaller basal than metabasal hook size, therefore belonging to species subgroup ‘Aa’ of Palm (1999). The description of the strobilar characters of N. hemipristis demonstrates that this species has a very characteristic, short, stout cirrus sac, with a length: width ratio of 2 in some segments, a feature unusual in most species of Nybelinia. However, this character might prove to be of taxonomic significance in the future. The new species is named after elasmobranch host genus, Hemipristis. the Nybelinia jayapaulazariahi Reimer, 1980 (Figs. 10-11) Material examined From ‘sole’, either Synaptura nigra Macleay, 1880 or Aseraggodes macleayanus (Ramsay, 1881): 1 postlarva, Moreton Bay, Queensland, coll. J. C. Pearson, 1968 (QM G 207318). Description SL = 1060; SW = 700; pbo = 470; pv = 450; pb = 365; BL =323 (320-325); BW=119 (85- 135); BR =2.7:1; SP = 1.3:1.2:1. TW = 32.5-35; hsr = 6. Basal tentacular swelling absent. Retractor muscles originate in basal parts of bulbs. Tentacular armature consists of homeomorphous slender uncinate hooks, L = 15.0-16.3, B = 11.2-13.7 (Fig. 10). Size of hooks diminishes towards basal part of tentacle, L = 11.2-13.7, B = 8.7-11.2 (Fig. 11). Remarks N. jayapaulazariahi was originally described from Cynoglossus sp. from the Bay of Bengal, India by Reimer (1980). Palm (1999) redescribed the species from another host, Harpodon nehereus (Hamilton-Buchanan, 1822), also from India. The tentacular armature has characteristic, slender, regularly curved hooks which increase in size towards the metabasal part of the tentacle. The present specimen, also from a ‘sole’, has a similar hook shape (compare Fig. 4 with Fig. 5 of Palm 1999) and bulb ratio, while it differs in having a larger scolex (1060 in the present specimen compared with 530 described previously) and hook size (11.2-16.3 in the current specimen compared with 5.6—11.2 in previous descriptions). However, the present specimen was in a poor state of preservation, and the larger scolex might be related to the slightly larger hook size. A small scolex of about 0.5-1.0 mm, together with the uncinate hook form, is characteristic for N. jayapaulazariahi, and soles seem to be important intermediate hosts. The present finding represents a new locality record for the species. Nybelinia mehlhorni sp. nov. (Figs 12-14) Types Holotype from stomach of Hemigaleus microstoma Bleeker, 1852, Heron Island, Queensland, coll. P. McBoarman, 20.xii.1995 (QM G 218032); paratype, same data (QM G 218033). Material examined Types. Description SL = 560 (Fig. 12); SW = 400; pbo = 330; 56 H. W. PALM & I. BEVERIDGE FIGURES 12-14. Nybelinia mehlhorni sp. nov. from Hemigaleus microstoma. Types, QM G 218032-3. 12. Scolex. 13. Metabasal and apical armature. 14. Mature proglottid. Note the uterine duct (ud) and the seminal receptacle (sr). Scale bars: Figure 12, 50 ym; Figure 13, 10 ym; Figure 14, 50 pm. FIGURES 15-16. Nybelinia scoliodoni (Vijayalakshmi, Vijayalakshmi & Gangadharam, 1996) from Diodon hystrix. QM G 218035-7. 15. Scolex. 16. Basal and metabasal armature, external surface. Scale bars: Figure 15, 150 ym; Figure 16, 25 um. TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 57 pv = 125; pb = 250; vel = 200; BL= 178 (150- 200); BW=58 (50-70); BR=3.1:1; SP = 1.3:0.5:1. Tentacles short; basal tentacular swelling absent; TW = 22.5-27.5. Tentacle sheaths sinuous to spirally coiled (TSW = 17.5— 20.0); muscular rings around basal part of tentacle sheaths seen. Retractor muscles originate in basal parts of bulbs. Tentacular armature homeoacanthous, homeomorphous (Fig. 13); distinctive basal armature absent. Hooks falciform, with stout base, elongated handle and strongly recurved tip. Hooks increase in size from basal (L = 3.5-7.5, B = 2.5—5.0) to metabasal region (L = 15.0-17.5, B = 6.3-8.7) then decrease towards apex (L = 11.3-13.8, B = 2.5-4.3); hsr = 7. Small worms, length 25 (16) mm, maximum width 950 (700) with 135 (90) segments. Strobila acraspedote, velum straight; pre-mature (testes visible) proglottids (Fig. 14) 80-280 long x 560-790 wide; mature segments (female genitalia developed) wider than long, 380-600 x 600-900; terminal segment 830 x 800. Genital pore almost lateral, 130-200 from anterior end (190 in terminal segment); pores alternate irregularly. Cirrus sac elongate, 170-220 x 40-60 (n=5) in mature segments, length:width ratio 4.1:1 (2.8-5.5:1), distal pole directed anteromedially, not reaching anterior end of segments; cirrus unarmed, internal and external seminal vesicles absent. Vas deferens greatly coiled in midline of segments, extending posteriorly towards female genital complex. Testes 103-120 in number, 80 x 95 in size, smaller testes peripheral, 35-40 in size, distributed in single layer; testes in 2 lateral groups confluent posterior to ovary; 10-16 testes anterior to cirrus sac. Vagina tubiform, 30 in diameter, ventral to cirrus sac, passes anteromedially to midline, then posteriorly to ovary; seminal receptacle present, 60-90 x 25-50. Ovary in centre of segment, ovarian lobes 250— 300 x 160-210. Vitelline follicles encircle medulla, follicles 30-45 in diameter. Uterine pore absent. Remarks Following Palm (1999), N. mehlhorni sp. nov. belongs to Nybelinia species without a characteristic basal armature and a basal hook size smaller than the metabasal one, thus belonging to the subgroup ‘Aa’. On the basis of the hook shape, the species resembles N. aequidentata (Shipley & Hornell, 1906) and N. goreensis Dollfus, 1960. While the former is much larger than the present specimen, the latter species has different scolex proportions as well as a different hook arrangement, as illustrated recently by Palm and Walter (2000). N. mehlhorni sp. nov. also resembles N. bengalensis Reimer, 1980 in its falciform hooks and in hook size (Reimer 1980). However, N. bengalensis can be distinguished by having a distinctly different scolex form and size, a bulb ratio of about 2, long and slender tentacle sheaths and tentacles with spaced falciform hooks (Reimer 1980). By contrast, N. mehlhorni sp. nov. has short tentacles with hooks tightly spaced along the tentacle. The scolex (0.56 mm in total length) is much smaller. The new species is named in honour of Prof. Heinz Mehlhorn, Heinrich-Heine-University, Diisseldorf. Nybelinia schmidti Palm, 1999 Material examined From /surus oxyrhinchus Rafinesque, 1810: 1 adult, Bicheno, Tasmania, coll. B. G. Robertson, 24.iv.1987 (SAM AHC 28313). Description SL = 1600; SW = 850; pbo = 1040; pv = 680; pb = 340; vel = 600; BL = 298 (270-330); BW = 131 (110-150); BR = 2.3:1; SP = 3.0:2.0:1. TW basal = 45-48, TW metabasal = 30.0-32.5; hsr = 5-6. Basal tentacular swelling absent. Tentacle sheaths spirally coiled, TSW = 25—40. Prebulbar organs absent, muscular rings present around the tentacle sheaths. Retractor muscles originate in basal part of bulbs. Tentacular armature homeoacanthous, homeomorphous, consisting of massive rosethorn-shaped hooks with anterior extension of base, size in metabasal region L = 13.8-15.0, B = 11.3—12.5, decreasing towards base, L=8.8-11.3, B = 6.2-8.8. Characteristic basal armature absent. Strobila immature with few segments, acraspedote; segments wider than long (660-690 x 15-60). Remarks The present specimen closely resembles that of the type, also from Jsurus oxyrhinchus (syn. Tsurus glaucus), in having similar scolex proportions and almost identical hook sizes. Other species with a similar tentacular armature are Nybelinia strongyla Dollfus, 1960 and Heteronybelinia australis sp. nov. (vide infra). While the latter species differs in having homeomorphous hooks of different sizes on the 58 H. W. PALM & I. BEVERIDGE internal and external tentacle surfaces as well as smaller basal hooks, N. strongyla differs in scolex proportions and larger hook sizes. The present record extends the range of distribution from the South African to the southern Australian coast. Nybelinia scoliodoni (Vijayalakshmi, Vijayalakshmi & Gangadharam, 1996) (Figs 15-16) Material examined From Diodon hystrix Linnaeus, 1758: Heron Island, Queensland, 2 postlarvae, coll. J. Sakanari, 23.viii.1986 (QM G 218034); 8 postlarvae, coll. M. K. Jones, 6.vii.1995 (QM G 218035-218037); 8 postlarvae, coll. T. H. Cribb, Jan. 1997 (QM G 218038-218041). From Makaira indica (Cuvier, 1832): 1 postlarva, Cape Bowling Green, Queensland, coll. P. Speare, 18.ix.1987 (SAM AHC 21351). From Istiophorus platypterus (Shaw & Nodder, 1792): Whitsunday Island, Queensland, coll. P. Speare, 5.xii.1988 (SAM AHC 21351). From Carcharhinus limbatus (Valenciennes, 1839): 1 adult, stomach, Bremer Island, Northern Territory, coll. J. Stevens, 29.v.1985 (SAM AHC 28314). Description SL = 1350, 1115 (Fig. 15); SW = 860, 800; pbo = 670, 520; pv = 230, 210; pb = 300, 320; vel = 330, 330; app = 310, 200; BL = 288 (280— 300), 282 (260-300); BW = 115 (110-120), 125 (120-130); BR =2.5:1, 2.3:1; SP = 2.2:0.8:1, 1.6:0.7:1. Tentacles nearly completely evaginated, TL = 390, 375; basal tentacular swelling absent. TW at basal armature 35, 30; TW at metabasal armature 50, 45; TW at apex 30, 25. Tentacle sheaths sinuous, TSW = 30, 40; prebulbar organs and muscular rings around basal part of tentacle sheaths not visible. Retractor muscles originate in basal parts of bulbs. Metabasal armature homeoacanthous, homeomorphous; distinctive basal armature present (Fig. 16). Basal armature consists of about 10 rows with compact rosethorn- shaped hooks, increasing in size from L = 4.5, B =3.9 (row 1) to L= 19.5, B=11.7 (row 10). From row 11, hook form changes to slender spiniform, L = 22.7, B = 13.0. Metabasal and apical hooks L = 29.9, with small base, B= 11. Number of hooks per half row (hsr) 6-7 in basal region, decreasing to 5 in metabasal and apical regions. Remarks Nybelinia scoliodoni is a widely distributed tentaculariid and is here recorded for the first time from Australian waters. The species is easily recognised by its rosethorn-shaped basal armature of about 11 rows which is followed by long, spiniform metabasal hooks. The present specimens correspond with the description given by Palm (1999) in having a similar armature and bulb ratio. They differ in having a larger scolex and hook size. However, they correspond closely to the original description of Nybelinia (= Tentacularia) scoliodoni of Vijayalakshmi et al. (1996). The present findings constitute three new host records. C. limbatus has been recorded previously as a definitive host for N. scoliodoni by Palm (1999). The specimens described here are the same as those reported by Speare (1999). Nybelinia strongyla Dollfus, 1960 (Figs 17-20) Material examined From Argyrosomus hololepidotus Lacépéde, 1802: 2 postlarvae from stomach, Murray Mouth, South Australia, coll. M. G. O’Callaghan, May 1992 (SAM AHC 28345). From Johnius vogleri (Bleeker, 1853): 1 postlarva from body cavity, Heron Island, Queensland, coll. J. Reddich, Jan 1997 (QM G 218109). Description Postlarvae from A. hololepidotus: SL = 1303, 1566 (Fig. 17); SW = 648, 796; pbo = 700, 781; pv = 647, 764; pb = 292, 355; vel = 332, 326; app = 374, 429; BL = 286 (244-292), 355 (334— 380); BW = 86 (82-90), 98 (92-107); BR = 3.3:1, 3.6:1; SP = 2.4:2.2:1, 2.2:2.2:1. Tentacles not completely evaginated; basal tentacle swelling absent. TW = 50-58, 43-45. Tentacle sheaths sinuous, TSW = 36-40, 36-40; prebulbar organs and muscular rings around basal part of tentacle sheaths not visible. Retractor muscles originate in basal part of bulbs. Tentacle armature homeoacanthous, homeomorphous (Fig. 18); distinctive basal armature absent. Hooks slender, rosethorn-shaped, increasing in size from basal towards metabasal part of tentacle. Metabasal hooks L = 17.5—20.0; B = 13.8-16.3; basal hooks L= 10.0-12.5; B = 10.0-12.5; hsr = 7. Postlarva from J. vogleri: SL = 1880; SW = 1660; pv = 680; pb = 440; BL = 308 (300- 325); BW = 129 (115-150); BR = 2.4:1; TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 59 FIGURES 17-18. Nybelinia strongyla Dollfus, 1960 from Argyrosomus hololepidotus. SAM AHC 28345. 17. Scolex. 18. Metabasal armature. Scale bars: Figure 17, 150 um; Figure 18: 15 um. FIGURES 19-20. Nybelinia strongyla from Johniops vogleri. QM G218109. 19. Metabasal armature. 20. Basal armature. Scale bars: Figure 19-20: 10 um. FIGURES 21-24. Nybelinia thyrsites Korotaeva, 1971 from Trachurus declivis. QM G 214194-S. 21. Scolex. 22. Basal armature. 23. Metabasal armature with falciform hooks. 24. Apical armature with rosethorn- shaped hooks. Scale bars: Figure 21, 200 um; Figures 22—24, 25 um. 60 H. W. PALM & I. BEVERIDGE SP = -:1.5:1. TW. basal = 37-40, TW metabasal = 30-33. Basal tentacular swelling absent. Tentacle sheaths straight; TSW = 20-25. Prebulbar organs absent, muscular rings around basal part of tentacle sheaths absent. Retractor muscles originate in basal part of bulbs. Armature homeoacanthous, homeomorphous (Fig. 19); characteristic basal armature absent (Fig 20). Hooks rosethorn-shaped with anterior extension of basal plate; hooks in basal part of tentacle smaller (L = 7.5-10.0; B = 7,.5—9.5) than in metabasal (L = 13.8-16.3; B = 10.0—12.5) armature. Number of hooks per half spiral (hsr) diminishes from 6—7 in metabasal region to 5-6 in apical part of tentacle. Remarks Nybelinia strongyla Dollfus, 1960 was described from a postlarva from the west coast of Africa, and is characterised by the homeoacanthous tentacular armature with slender rosethorn-shaped hooks, reaching a size of up to 16 in the metabasal armature. Palm and Walter (2000) described adult specimens of N. strongyla from Africa. The present specimens correspond closely in their tentacular armature as well as in scolex proportions to those specimens described by Dollfus (1960) and Palm and Walter (2000). Minor variations in the hook pattern are observed within N. strongyla but are difficult to define, and many scolex characters within tentaculariids appear to be variable (Palm 1999, Palm and Walter 2000). Therefore, the postlarvae described above are identified as N. strongyla. However, some doubt surrounding the identification remains. Further studies are needed to determine whether postlarvae with rosethorn-shaped hooks as described above are conspecific with the African material, or whether they belong to a new species of Nybelinia. Argyrosomus hololepidotus is a new host record, and the distribution of the species is extended to the southern Australian coast. The specimen from J. vogleri, with rosethorn- shaped hooks, resembles N. lingualis, N. schmidti, N. sphyrnae, N. strongyla and N. thyrsites. It differs from N. lingualis in having basal hooks with an anterior extension of the basal plate, and from N. thyrsites in having differently shaped metabasal hooks and a larger bulb ratio (see below). N. schmidti differs in having smaller hooks and different scolex proportions. N. sphyrnae differs in having slender, more uncinate hooks in the metabasal armature, while the hooks are more massive rosethorn-shaped in N. strongyla. In the present specimen, the basal hook length is slightly smaller than described by Palm and Walter (2000) for NV. strongyla. However, the metabasal hooks are the same size. Although the scolex proportions were difficult to measure in the present specimen, it is also identified as N. strongyla, representing a new host and locality record. Nybelinia thyrsites Korotaeva, 1971 (Figs 21-24) Material examined From Trachurus declivis Jenyns, 1841: 2 postlarvae from stomach wall, Crayfish Bay, Tasmania, coll. K. B. Sewell, 11.v.1987 (QM G 214194, 212145). From Carcharhinus brachyurus (Giinther, 1870): 1 adult from stomach, Goolwa, South Australia, coll. R. Martin, 28.x.1985 (SAM AHC 28312). From Mustelus antarcticus Giinther, 1870: 1 adult from stomach, Goolwa, South Australia, coll. R. Martin, 28.x.1985 (SAM AHC 28311). Description SL = 2032, 1550 (Fig. 21); SW = 1520, 700; pbo = 1024, 610; py = 640, 500; pb = 368, 520; vel = 992, 490; app = 496; BL = 358 (355-365), 495 (490-500); BW = 117 (114-120), 155 (140- 170); BR=3.1:1, 3.2:1; SP =2.8:1.7:1, 1.2:1:1, Tentacle sheaths spirally coiled; TSW = 51-57, 35-45. Prebulbar organs absent, muscular rings around basal part of tentacle sheaths present. Retractor muscles originate in basal part of bulbs. Tentacles long (TL = 832) and slender, basal tentacular swelling absent; TW basal = 38-44, 25-30; TW metabasal = 57.0, 57.7-62.5; TW apical 32.0, 27.5. Characteristic basal armature present (Fig. 22), consisting of about 7-8 rows of homeomorphous, rosethorn-shaped hooks with slight anterior extension, increasing in size towards metabasal armature (lst row hooks: L = 9.5-11.0, B = 8.0-11.0, L = 7.5-8.7, B = 6.3- 7.5; 8th row: L= 15.8-17.4, L = 16.2-17.5, B = 11.0-12.6, B = 10.0-12.5). Metabasal armature of about 12-14 rows of hooks, with largest hooks in rows 12-17. Metabasal armature with slender falcate to falciform hooks (L = 21.7— 26.9, B = 8.7-11.0; L = 21.3-24.0, B = 11.3- 12.5) (Fig. 23). Apical hooks rosethorn-shaped, with hook form changing from slender rosethorn- shaped with slight anterior extension to rosethorn- shaped with distinct anterior extension (Fig. 24). TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 61 Hooks diminish in size towards apical region (34th row of hooks: L= 14.2-15.8, B= 11.0- 14.2); hsr basal: 6—7; hsr metabasal and apical: 6. Remarks N. thyrsites was redescribed in detail by Beveridge and Campbell (1996). The present material corresponds with the redescription given. In addition, the basal hook form differs from the metabasal and apical hooks; thus, the species belongs to subgroup ‘Ba’ of Palm (1999). The present specimens from Trachurus declivis provide a further example in which the postlarvae can be larger than adult worms, and also show variable scolex measurements depending on the state of contraction. Trachurus declivis and Carcharhinus brachyurus are new hosts for N. thyrsites. Nybelinia victoriae sp. nov. (Figs 25-26) Types Holotype from body cavity of Lepidotrigla modesta Waite, 1899, Port Phillip Bay, Victoria, coll. R. Norman, 17.iii1.1989 (SAM AHC 28343); paratype, same data (SAM AHC 28344). Material examined Types. Description SL = 1030, 1050 (Fig 25); SW=700, 650; Pbo = 570, 450; Pv = 350, 320; Pb =315, 320; BL = 310 (300-315), 310 (305-320); BW = 124 (115-135), 116 (110-125); BR =2.5:1, 2.7:1; SP = 1.8:1.1:1, 1.4:1.0:1. Tentacles elongate; basal tentacular swelling absent. TW = 40.0-50.0, 45.0-S0.0. Tentacle sheaths sinuous to spirally coiled (TSW = 40.0-S0.0, 35.0-40.0); prebulbar organs and muscular rings around basal part of tentacle sheaths not seen. Retractor muscles originate in basal part of bulbs. Armature homeoacanthous, homeomorphous (Fig. 26); distinctive basal armature absent. Hooks falciform, with stout base, elongated handle and strongly recurved tip. Hooks increase in size from basal (L = 7.5-10.0, B = 6.3-8.8) to metabasal region (L = 12.5—15.0, B = 7.0-8.0) and decrease towards apical (L = 11.3-12.5, B = 6.3-8.0) part of tentacle; hsr = 7. Remarks This species, with a tentacular armature consisting of slender falciform hooks, increasing in size towards the metabasal region and decreasing in size towards the apical region, resembles Nybelinia aequidentata Shipley & Hornell, 1906; N. syngenes Pintner, 1927; N. anantaramanorum Reimer, 1980; and N. bengalensis Reimer, 1980. While the former two species differ in having different scolex proportions and distinctly larger hooks (see remarks on N. aequidentata), the present specimens are most similar to N. bengalensis and N. anantaramanorum. N. anantaramanorum has larger hooks and was considered a species inquirenda by Palm and Walter (2000). N. bengalensis differs in having different scolex proportions, including very short bulbs (BR < 2) (Reimer 1980). Thus, the present specimens represent a new species here named Nybelinia victoriae sp. nov., the specific epithet being derived from the Australian state from which samples were collected. With hook size smaller in the basal rather than the metabasal region, the species is considered to belong to subgroup ‘Aa’ of Palm (1999). Genus Heteronybelinia Palm, 1999 Heteronybelinia australis sp. nov. (Figs 27-30) Types Holotype from stomach of Carcharhinus amboinensis (Miiller & Henle, 1839), St Lawrence, Queensland, coll. B. G. Robertson, 29.x.1985 (SAM AHC 28315); paratype, same data (SAM AHC 28316). Material examined From Carcharhinus amboinensis: types. From Carcharhinus brachyurus: 1 specimen, Goolwa, South Australia, coll. R. Martin, 28.x.1985 (SAM AHC 28317). Description SL = 1190, 1210 (Fig. 27); SW = 660, 680; pbo = 650, 700; pv = 530, 650; pb = 350, 310; vel = 220, 190; BL = 313 (305-320), 308 (290— 340); BW = 138 (125-150), 119 (110-130); BR = 2.3:1, 2.6:1; SP = 1.9:1.7:1, 2.3:2.1:1. Tentacle sheaths spirally coiled. TSW = 27.5-— 32.5, 25.0-35.0. Tentacles robust, TL = 255-290, increasing in width towards apex. TW basal and metabasal 30.0—32.5; basal tentacular swelling absent. Prebulbar organs absent; muscular rings 62 H. W. PALM & I. BEVERIDGE FIGURES 25-26. Nybelinia victoriae sp. nov. from Lepidotrigla modesta. Types, SAM AHC 28343-4. 25. Scolex. 26. Basal and metabasal armature. Scale bars: Figure 25, 100 pm; Figure 26, 20 um. FIGURES 27-30. Heteronybelinia australis sp. nov. from Carcharhinus amboinensis. Types, SAM AHC 28315-6. 27. Scolex. 28. Metabasal armature. 29. Basal armature. 30. Mature proglottid. Note the uterine pore. Scale bars: Figure 27, 100 um; Figures 28-29, 10 um; Figure 30, 50 um. TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 63 around basal parts of tentacle sheaths present. Retractor muscles originate at base of bulbs. Tentacular armature homeoacanthous, heteromorphous, with hooks tightly spaced along tentacle (Fig. 28); characteristic basal armature absent (Fig. 29). Hooks massive, rose-thorn shaped, with slight anterior extension of base, increase in size from basal towards metabasal armature. Hook size differs on bothridial/ antibothridial (L = 12.5-15.0, B = 11.3-13.7) and antibothridial/bothridial (L = 11.3-12.5, B = 7.0— 8.7) tentacle surfaces (Fig. 28), with two tentacles having largest hooks on bothridial surfaces and two other tentacles having largest hooks on antibothridial surfaces. Basal hooks L = 5.0-7.5, 7.5-8.7; B =8.7-10.0 (internal) and L = 5.0-6.3, 5.0-7.5; B = 3.8-5.0, 5.0-6.3 (external); hsr = 6— 7. Strobila only very slightly craspedote, velum scalloped; total length 18.0 and 15.0 mm, maximum width 640 and 730, with 125 and 155 segments. In holotype, terminal segment pre- mature (320 x 550), segments wider than long, 231 x 630. Genital pores ventro-submarginal, 60 from anterior margin; alternate irregularly. Cirrus sac of pre-mature segments elongate and slender, 210-230 x 35-50 (n = 5), length:width ratio 5.3:1 (4.6-6.0:1), with distal pole directed anteromedially, reaching anterior end of segment (Fig. 30); cirrus unarmed; internal and external seminal vesicles absent. Vas deferens coiled, extends posteriorly from cirrus sac towards female genital complex. Testes 106-130 in number, smallest testes at margin of medulla, distributed in 1-2 layers; in 2 lateral groups, confluent posterior to ovary; extend between ovarian lobes; 8-10 testes anterior to cirrus sac. Ventral osmoregulatory canal 15 in diameter, internal to sinuous dorsal canal, 5 in diameter; 24 prominent bands of longitudinal muscles on each side of strobila. Segments in paratype mature, wider than long (240 x 700 to 380 x 720). Genital pores 110 from anterior margin. Cirrus sac of mature segments (Fig. 30) elongate and slender, 260-320 x 50-60 (n=5), length:width ratio 5.2:1 (4.3- 5.8:1). Central testes 45-65 in diameter, smallest testes peripheral, 30-50 in diameter, distributed in 1-2 layers; ovarian lobes 110-200 x 80-110, seminal receptacle SO-65 x 25-30. Remarks Heteronybelinia australis sp. nov. has a unique armature. The tentacular hooks are tightly spaced and appear homeomorphous along the tentacle. However, the hook sizes differ on the bothridial and antibothridial tentacular surfaces. In addition, two tentacles have the largest hooks on the bothridial surfaces while the other two tentacles have the largest hooks on the antibothridial surfaces. Whether this armature pattern is unique or whether it occurs in other tentaculariid species remains to be determined. The specific epithet indicates the occurrence of the species in Australian waters. Heteronybelinia estigmena (Dollfus, 1960) (Figs 31-36) Material examined From Carcharhinus limbatus Valenciennes, 1839: 16 adults from spiral valve, Geraldton, Western Australia, coll. B. G. Robertson, 27.xi.1986 (SAM AHC 21352, 28318); 4 adults from spiral valve, Nickol Bay, Western Australia, coll. B. G. Robertson, 11.xi.1986 (SAM AHC 28319); 3 adults from spiral valve, Darwin Harbour, Northern Territory, coll. B. G. Robertson, 28.viii.1986 (SAM AHC 28320); 6 adults from spiral valve, Fog Bay, Northern Territory, coll. B. G. Robertson, 4.x.1986 (SAM AHC 28329). From Carcharhinus amblyrhynchoides (Whitley, 1934): 2 adults from spiral valve, Tommy Cut Mouth, Northern Territory, coll. B. G. Robertson, 10.ix.1986 (SAM AHC 28321). From Carcharhinus sp: 3 adults from spiral valve, Queensland, coll. B. G. Robertson, Oct. 1985 (SAM AHC 18322). From Sarda australis (Macleay, 1881): 9 postlarvae from gill arches, Heron Island, Queensland, coll. M. K. Jones, Jan. 1991 (QM G 218042-218046). Description Adult: SL = 1650, 1500 (Fig. 31); SW = 790, 770; pbo = 820, 820; pv = 690, 610; pb = 480, 430; ppb = 110, 80; vel = 330, 320; BL = 455 (440-480), 425 (420-430); BW = 133 (130- 140), 120 (118-122); BR =3.4:1, 3.6:1; SP = 1.7:1.4:1, 1.9:1.4:1. Tentacles long, slender (TL = 520, 560), TSW = 42.5-47.5, 45.0-55.0; TW basal 45-50, 42.5-45; TW apical 25-30, 27.5-32.5; basal tentacular swelling absent. Prebulbar organs absent, muscular rings around basal part of tentacle sheaths visible in one specimen. Retractor muscles originate at bases of bulbs. Tentacular armature homeoacanthous, heteromorphous, no characteristic basal armature present (Figs 32-33). Hooks rosethorn-shaped, increase in size towards metabasal part of 64 H. W. PALM & I. BEVERIDGE FIGURES 31-34. Heteronybelinia estigmena (Dollfus, 1960) from Carcharhinus limbatus. SAM AHC 28318-20. 31. Scolex. 32. Metabasal armature. 33. Basal armature. 34. Mature proglottid. Scale bars: Figure 31, 150 ym; Figures 32-33, 15 um; Figure 34, 100 um. FIGURES 35-36. H. estigmena from Sarda australis. QM G 218042- 46. 35. Scolex. 36. Basal armature. Scale bars: Figure 35, 150 ym; Figure 36, 15 um. TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 65 tentacle. Hook form varies from compact and rosethorn-shaped (bothridial) to smaller rosethorn-shaped hooks with elongated base (antibothridial). Hook size, metabasal, L = 11.3- 12.5; B = 9.3-10.8 (bothridial) and L = 9.3-10.8; B = 10 (antibothridial), with largest hooks L = 13.8-15.0; B = 10.0-12.5 (bothridial); hooks diminish in size towards apical part of tentacle. Hook size in basal region L = 7.5-8.8; B = 6.3- 10.0 (bothridial) and L =5.0-7.5; B = 6.3-7.5 (antibothridial); hsr = 6-7. Strobila 56 mm long, maximum width 1250, about 280 acraspedote segments (Fig. 34). First segments behind velum short, enlarge in size, mature segments wider than long, from 500 x 840 to 700 x 1030. In mature segments, genital atrium ventro- submarginal, in anterior third of segment; genital pores alternate irregularly. Cirrus sac elongated, 300-450 x 80-110 (n=5), length:width ratio 3.8-4.1:1, directed anteromedially, sac thin- walled; cirrus unarmed and coiled within sac. Ovary median, follicular, ovarian lobes 200—230 (length) x 110-170 (width), Mehlis’ gland 80 in diameter, seminal receptacle 110 x 70; testes of variable shape, 60-80 x 30-60 (n=5) in diameter, arranged in single layer; testes number 106-126 (n=5) per segment, encircle female genital complex and occupy entire medulla except for region of female genital complex and anterior to it; 7-9 testes anterior to cirrus sac. Vitelline follicles 20-40 in diameter; egg diameter 20-25; ventral osmoregulatory canals along margins of cortex, 6 in diameter. Postlarva: SL = 1650, 1500 (Fig. 35); SW = 790, 770; pbo = 820, 820; py = 690, 610; pb = 480, 430; ppb= 110, 80; vel = 330, 320; BL = 460 (430-470), 445 (440-450); BW = 115 (110-120), 120 (118-122); BR =4.0, 3.7; SP = 1.7:1.4:1, 1.9:1.4:1. Tentacles long, slender (TL = 550-580), TSW = 40-45; TW basal = 35— 40, TW metabasal and apical = 30-40; basal tentacular swelling absent. Prebulbar organs absent, muscular rings around basal part of tentacle sheaths present. Retractor muscles originate at base of bulbs. Tentacular armature homeoacanthous, heteromorphous; characteristic basal armature absent. Hook shape varies from compact, rosethorn-shaped (bothridial) to smaller rosethorn-shaped hooks with elongated base (antibothridial); hooks increase in size towards metabasal part of tentacle on bothridial and antibothridial surface (Fig. 36). Hook size, metabasal, L=12.5-15.0, B= 11.8-13.3 (bothridial) and L = 11.3-12.5; B = 7.5-10.0 (antibothridial); basal L = 7.5—-10.0; B = 8.8-11.3 (bothridial) and L = 5.0-7.5; B =6.3-8.8 (antibothridial); hsr = 6-7. Remarks Heteronybelinia estigmena (Dollfus, 1960) is a well-described tentaculariid. Palm (1995) reported this cestode as Nybelinia alloiotica Dollfus, 1960 from Carcharhinus limbatus in the Atlantic, the same host species as in the present study. Palm (1999) and Palm and Walter (2000) provided additional information on the species. H. estigmena resembles H. perideraeus (Shipley & Hornell, 1906) and H. elongata (Shah & Bilquees, 1979) known from the coast of India. However, the hook size increases on both the bothridial and antibothridial tentacle surfaces in H. estigmena. In H. elongata, the hook size increases on the antibothridial tentacle surface only (Palm & Walter 1999), while in H. perideraeus, the basal and metabasal hook sizes are the same (Palm 1999). The present study represents the first record of the cestode from Australian waters. Carcharhinus amblyrhynchoides and Sarda australis represent new hosts for the species. Heteronybelinia pseudorobusta sp. nov. (Figs 37-41) Types Holotype from gills of Lepidotrigla modesta Waite, 1899, Port Phillip Bay, Victoria, coll. R. Norman, 17.iii.1989 (SAM AHC 28341); 3 paratypes, same data (SAM AHC 28342). Material examined Types. Description Small form (holotype and one paratype): SL = 960 (Fig. 37), 730; SW = 660, 550; pbo = 570, 450; pv = 420, 270; pb = 285, 290; app = 220, 160; vel = 70, 60; BL = 280 (275- 285), 298 (275-315); BW = 150 (145-155), 139 (110-165); BR = 1.9:1, 2.1:1; SP = 2.0:1.5:1, 1.6:0.9:1. Tentacle sheaths sinuous to spirally coiled; TSW = 45.0-47.5, 45.0-50.0; TW basal 45.0—-50.0, metabasal 40.0-45.0, basal tentacular swelling absent. Prebulbar organs absent, muscular rings around basal part of tentacle sheaths not seen, retractor muscles originate at base of bulbs. Tentacular armature homeoacanthous, heteromorphous, characteristic basal armature absent (Fig. 38). Hook form changes from compact, rosethorn-shaped 66 H. W. PALM & I. BEVERIDGE FIGURES 37-41. Heteronybelinia pseudorobusta sp. nov. from Lepidotrigla modesta. 37. Scolex of small form SAM AHC 28314. 38. Basal armature. 39. Metabasal armature. 40. Scolex of large form SAM AHC 28342. 41. Metabasal armature. Scale bars: Figure 37, 100 ym; Figures 38-39, 20 um; Figure 40, 100 um; Figure 41, 15 um. FIGURES 42-44. Mixonybelinia cribbi sp. nov. from Platycephalus arenarius. Holotype, QM G 218047. 42. Scolex. 43. Metabasal armature. 44. Basal armature. Scale bars: Figure 42, 200 um; Figures 43-44, 20 pm. TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 67 (bothridial) to more slender hooks with stout base (antibothridial) (Fig. 39). Hook size in metabasal armature ranged between L = 13.8-16.3; B = 11.3-13.8 (bothridial) and L = 16.3-17.5; B = 10,0—11.3 (antibothridial), hooks of basal part of tentacle smaller, between L = 10.0-11.0; B = 11.3-9.0 (bothridial) and L = 11.3-12.5; B = 8.8-10.0 (antibothridial), continuously increasing towards the tip; hsr = 6-7. Large form (2 paratypes): SL = 1310 (Fig. 40); 1310; SW = 600, 620; pbo = 810, 780; pv = 580, 500; pb = 390, 380; app = 330, 330; vel = 280, 310; BL = 376 (360-390), 321 (310-330); BW = 124 (115-140), 121 (105-135); BR =3.0:1; SP = 2.1:1.5:1. Tentacle sheaths sinuous to spirally coiled; TSW = 40-45, 42.5- 47.5; TW basal 45.0-47.5, metabasal 30.0-35.0, 30.0-35.0. Basal tentacular swelling absent, prebulbar organs absent, muscular rings around basal part of tentacle sheaths not seen. Retractor muscles originate at base of bulbs. Tentacular armature homeoacanthous, heteromorphous, characteristic basal armature absent. Hooks change from compact and rosethorn-shaped (bothridial) to more slender hooks with a stout base (antibothridial) (Fig. 41). Hook size in metabasal armature between L = 12.5-14.5; B = 11.3-12.5 (bothridial) and L = 14.5—15.5; B = 8.8-10.0 (antibothridial), hooks of basal part of tentacle small, between L = 8.8-10; B = 7.5— 9.0 (bothridial) and L = 6.3-8.8; B = 6.3-7.5 (antibothridial), increasing in size towards tip; hsr = 6-7. Remarks Heteronybelinia pseudorobusta sp. nov. is characterised by a heteromorphous tentacular armature consisting of small basal hooks increasing in size towards the metabasal region. The hook shape varies from rosethorn-shaped to slender hooks with a stout base on different surfaces of the tentacle. Small basal hooks which gradually increase in size towards the metabasal armature are characteristic of Heteronybelinia robusta (Linton, 1890). However, in contrast to H. pseudorobusta sp. nov., H. robusta has minute basal hooks, less than 5 in length, and the hook form is uniform along the tentacle. In the present specimens, the basal hook size is larger and the hook form is rosethorn-shaped. All other Heteronybelinia species differ in having a different hook shape or arrangement. Thus, the present specimens belong to a new species, H. pseudorobusta sp. nov., the name being derived from the similarity in hook arrangement to H. robusta (Linton, 1890). H. pseudorobusta is a species without a characteristic basal armature and with basal hook sizes smaller than metabasal hook sizes. It therefore belongs to the Heteronybelinia species subgroup ‘Aa’ in Palm (1999). Heteronybelinia pseudorobusta sp. nov. occurred in two different size groups, those smaller than 1 mm and those larger than 1.3 mm, respectively. Thus, the present description is an example of intraspecific morphological variability within species of Heteronybelinia, apparently due to scolex contraction during fixation. Genus Mixonybelinia Palm, 1999 Mixonybelinia beveridgei (Palm, Walter, Schwerdtfeger & Reimer, 1997) Material examined From Macruronus novaezelandiae (Hector, 1871): 1 postlarva, west coast of Tasmania, coll. K. Sewell, June 1986 (QM G 218067). Description SL = 3200; SW = 2750; pbo = 1750; pv = 850; pb = 1300; ppb = 80; vel = 950; app = 730; BL = 1226 (1150-1310); BW = 290 (240-320); BR = 4.2:1; SP = 1.3:0.7:1. Tentacle sheaths sinuous to spirally coiled; TSW = 100-130. Prebulbar organs absent, muscular rings around basal part of tentacle sheaths not seen. Retractor muscles originate in basal parts of bulbs. Basal tentacular swelling absent; TW basal = 110-120, TW metabasal = 115-135. Metabasal armature homeoacanthous, heteromorphous, characteristic homeomorphous basal armature consisting of about 6-7 rows of slender hooks with elongated shaft, stout base and strongly recurved at tip (L = 31.3+40.0, B = 18.8—27.5). Metabasal hooks strongly recurved, rosethorn-shaped with large base on antibothridial surface: L = 50.0-52.5, B = 37.5-42.5; slender falcate hooks with stout base on bothridial surface: L = 55.0-60.0, B = 27.5-30.0; hsr = 6. Remarks The present specimen is characterised by a homeomorphous basal armature of 6-7 rows of hooks and a heteromorphous metabasal armature of massive hooks, thus corresponding in form and size with specimens of M. beveridgei described from African waters by Palm et al. (1997). Other features are the large scolex size as well as TSW and TW. However, due to the contracted nature of 68 H. W. PALM & I. BEVERIDGE the scolex of the current specimen, values for scolex and bulb proportions as well as the tentacle sheaths differed, indicating the variability of these characters due to the degree of contraction of the scolex. The present finding represents a new host and locality record for M. beveridgei. Mixonybelinia cribbi sp. nov. (Figs 42-44) Types Holotype, postlarva from Platycephalus arenarius Ramsay & Ogilby, 1886, Heron Island, Queensland, coll. J. Reddick, Jan. 1997 (QM G 218047). Material examined Types. Description Scolex craspedote, SL = 2920 (Fig. 42); SW = 2200; pbo = 1700; pv = 1020; pb = 730; ppb = 220; vel = 760; app = 780; BL = 615 (600— 630); BW = 232 (220-240); BR = 2.7:1; SP = 2.3:1.4:1. Tentacle sheaths short; TSW = 50-65. Prebulbar organs absent, muscular rings around basal part of tentacle sheaths not seen. Retractor muscles originate in basal part of bulbs. Basal tentacular swelling absent; TW basal = 70-75, TW _ metabasal = 65-75. Homeoacanthous, heteromorphous metabasal armature consisting of rosethorn-shaped hooks on bothnidial and more slender hooks with stout base on antibothridial surface (Fig. 43); characteristic homeomorphous basal armature (Fig. 44) consisting of about 4 rows of slender hooks with enlarged base and strongly recurved at tip (L = 12.5-17.5, B = 10.0-15.0 to 8.8-11.3). Metabasal hooks strongly recurved, rosethorn- shaped with large base on antibothridial surface: L= 20.0—22.5, B = 18.8—21.3; slender falcate with stout base along bothridial surface: L = 21.3—23.7, B = 15.0-17.5; hsr = 6-7. Remarks M. cribbi sp. nov. resembles M. beveridgei in scolex size as well as in tentacular armature. Both species have a homeomorphous basal and a heteromorphous metabasal armature as well as a large scolex about 3 mm in length. However, M. cribbi sp. nov. is clearly distinguishable from M. beveridgei by having only 4 rows of basal hooks compared with 6 in M. beveridgei. In addition, the hooks have a distinctly enlarged base in M. cribbi and all hooks are smaller (20-24 in M. cribbi compared with 46-68 in M. beveridgei, see Palm et al. 1997). The new species was named after Dr T. H. Cribb, Department of Parasitology, University of Queensland, Australia. Mixonybelinia edwinlintoni (Dollfus, 1960) (Figs 45-53) Material examined From Sphyrna lewini (Griffith & Smith, 1843): 10 adults from stomach, Flat Top Island, Queensland, coll. B. G. Robertson, 23.x.1985 (SAM AHC 28324); 1 adult, Geraldton, Western Australia, coll. B. G. Robertson, 27.xi.1986 (SAM AHC 28323). From Rhynchobatus djiddensis (Forsskal, 1775): 9 postlarvae from spiral valve, Moreton Bay, Queensland, coll. S. Butler, 17.iv.1980 (QM G 218048-218060). From Carcharhinus melanopterus (Quoy & Gaimard, 1824): 1 postlarva, (QM G 4813). Description Adults: SL = 1780, 1620 (Fig. 45); SW = 1210, 1160; pbo = 1000, 1000; pv = 760, 690; pb = 620, 530; ppb = 60, 30; vel = 340, 330; BL = 585, 520, 520, 530, 490, within single specimen 580-600; BW = 235 (230-240); BR = 2.5:1; SP = 1.6:1.2:1. Tentacles robust; tentacle sheaths straight, TSW = 50-90. Prebulbar organs absent, muscular rings around basal part of tentacle sheaths visible in some specimens. Retractor muscles originate in basal part of bulbs: basal tentacular swelling absent; TW basal = 60-65; TW metabasal = 70- 75. Metabasal armature homeoacanthous, heteromorphous (Fig. 46); characteristic homeomorphous basal armature consisting of about 10 rows of slender falciform hooks (Fig. 46). Metabasal hooks strongly recurved along antibothridial surface: L = 20.0-25.0; B = 17.8— 21.3; slender falcate hooks along bothridial surface: L = 32.5-35.0; B = 15.0-17.5. Basal hook size: L = 17.5-20.0, B = 10.0-13.8; hsr basal: 7-8; hsr metabasal: 8-9. Largest cestode 53 mm, maximum width 880; 560 segments; strobila craspedote, velum irregularly scalloped, up to 50 wide; pre-mature (no ovary) segments (Fig. 47) wider than long, 100 x 1020 to 120 x 1020. Genital pores submarginal, in first third of segment; alternate irregularly. Cirrus sac elongate, 280-330 x 45-60 (n=5) in mature segments, length:width ratio 5.7:1 (4.7-7.1:1), distal pole directed TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 69 = : ‘j Y “\ \ Sees SLR RUS 4 Coarse aks RSE —_ FIGURES 45-47. Mixonybelinia edwinlintoni (Dollfus, 1960) from Sphyrna lewini. SAM AHC 28324. 45. Scolex. 46. Basal and metabasal armature. 47. Pre-mature proglottid. Scale bars: Figure 45, 150 pm; Figure 46, 25 um; Figure 47, 60 um. anteromedially, nearly reaching anterior end of segment; cirrus unarmed; internal and external seminal vesicles absent. Vas deferens coiled, extends to midline of segment, then posteriorly towards female genital complex. Testes in two groups, not confluent posterior to ovary, 80 (poral) and 116 (antiporal) (estimated), 30-40 in size (central), smallest testes at margin of medulla, external to osmoregulatory canal (15-25), distributed in 2—3 layers, extend anterior to cirrus sac; seminal receptacle present. Small vitelline follicles encircle medulla, between 24 longitudinal muscle bands. Osmoregulatory canals 5 and 20 in diameter. 70 H. W. PALM & I. BEVERIDGE Hor, wee: B GES SRe2277 Ch SD 19 GO, Abs —_ ORS FIGURES 48-52. Mixonybelinia edwinlintoni (Dollfus, 1960) from Rhynchobatus djiddensis. QM G 218048-60. 48. Scolex. 49. Basal armature. 50. Metabasal armature, 20 rows from the base. 51-52. Bulb with tentacle sheath. Scale bars: Figure 48, 150 um; Figures 49-50, 25 um; Figures 51-52, 50 um. FIGURE 53. M. edwinlintoni from Carcharhinus melanopterus. QM G 4813. Scolex. Scale bar: 100 um. FIGURE 54. Kotorella pronosoma (Stossich, 1901) from Dasyatis fluviorum. QM G 218063. Mature proglottid. Scale bar: 50 ym. TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 71 Postlarvae (from R. djiddensis): SL = 1600, 1500 (Fig. 48); SW = 1000, 1180; pbo = 1020, 950; pv = 680, 600; pb = 420, 470; ppb = 140, 120; vel = 280, 250; app= 280, 250; BL = 390 (370-410), 430 (410-440); BW = 188 (180-190), 163 (150-180); BR = 2.1:1, 2.6:1; SP = 2.4:1.6:1; 2.0:1.3:1. Tentacle sheaths straight, TSW = 40- 85. Prebulbar organs absent, muscular rings and thickenings around basal part of tentacle sheaths visible in some specimens (Figs 51, 52). Retractor muscles originate in basal part of bulbs (Fig. 48); basal tentacular swelling absent; TW basal = 80, 80; TW metabasal = 70, 80. Homeomorphous basal armature of about 10 rows of slender falciform hooks (Fig. 49), metabasal armature homeoacanthous, heteromorphous (Fig. 50). Metabasal hooks strongly recurved along antibothridial surface: L = 22.5—25.0; B = 17.5- 20.0; slender falcate hooks along bothridial surface: L = 23.8—26.3; B = 12.5-15.0. Basal hook size: L = 17.5-20.0, B = 12.5—13.8; hsr basal: 7-8; hsr metabasal: 8-9. Postlarva (from C. melanopterus): SL = 1648 (Fig. 53); SW = 848; pbo = 912; pv = 768; pb = 432; ppb = 96; app = 320; vel = 240; BL = 429 (411-443); BW =171 (158-177); BR = 2.5:1; SP = 2.1:1.8:1. Tentacles massive (TL = 630), TSW = 68-70; TW basal 77-80, TW metabasal 85-89, basal tentacular swelling absent. Prebulbar organs absent; muscular rings around basal part of tentacle sheaths not visible; retractor muscles originate at base of bulbs. Tentacular armature homeoacanthous, heteromorphous; characteristic basal armature absent. Hooks diminish in size towards base; hook shape varies from compact rosethorn-shaped (antibothridial) to slender rosethorn-shaped hooks with stout base (bothridial). Hook size, metabasal, rows 16-17, L = 23.7-25.2; B = 14.2-15.8 (antibothridial) and L = 25.2—26.8; B =9.5—-12.0 (bothridial); basal L = 17.4-19.0; B = 14.2-15.8 (antibothridial) and L = 17.4-19.0; B = 12.6-14.2 (bothridial); hsr = 7-8. Remarks Palm and Walter (2000) redescribed Mixonybelinia edwinlintoni (Dollfus, 1960) illustrating a distinctly different basal and metabasal armature within the species. While the metabasal armature has different hooks on the bothridial and antibothridial tentacle surface, the hooks of the basal armature appear to be homeomorphous. The arrangement of the hooks, together with their characteristic shape, and the scolex shape, indicate that the present specimens belong to M edwinlintoni. Rhynchobatus djiddensis and Carcharhinus melanopterus represent two new hosts for the species. Prior to the re-description of M. edwinlintoni, Palm (1999) erected Heteronybelinia heteromorphi, a species which is also very similar to the present material. In H. heteromorphi the basal armature is heteromorphous, while in M. edwinlintoni it is homeomorphous. In addition, the type material of H. heteromorphi was described as being acraspedote while M. edwinlintoni is craspedote (Fig. 47). Interestingly, both species occur within the same host genus, Sphyrna. Other characters such as the massive scolex form, the heteromorphous metabasal armature, and the testis sizes are similar in both species. The type material of H. heteromorphi needs to be re-examined to confirm the differences between the species. Mixonybelinia southwelli (Palm & Walter, 1999) Material examined From Makaira indica (Cuvier, 1832): 1 postlarva, Cape Bowling Green, Queensland, coll. P. Speare, 18.ix.1987 (QM G 218061). From Chaerodon venustus (DeVis, 1885): 1 postlarva, Heron Island, Queensland, coll. R. Bray, 30.1.1991 (QM G 218062). Description SL = 2000, 1550; SW=1050, 690; pbo = 1030, 890; pv = 850, 660; pb = 440, 590; ppb = 40; vel = 450, 220; app = 670, 270; BL = 411 (400-425), 548 (520-580); BW = 153 (140-165), 153 (140-170); BR =2.7:1, 3.6:1; SP = 2.3:1.9:1, 1.5:1.1:1. Tentacle sheaths spirally coiled; TSW = 55-60, 50-60. Prebulbar organs absent, muscular rings around the basal part of tentacle sheaths not seen; retractor muscles originate in basal part of bulbs; tentacular swelling absent; TW basal = 50.0— 55.0, 47.5-55.0, TW metabasal = 35-47, 30-— 35. Armature homeoacanthous, heteromorphous; characteristic basal armature consisting of about 14-16 rows. Antibothridial metabasal hooks massive, rosethorn-shaped, L = 15.0-17.5, 15.0-16.3, B = 13.7-15.0, 15.0- 17.5; bothridial hooks more slender and slightly curved with stout base, L = 18.7-20.0, 17.5— 20.0, B = 11.3-12.5, 11.3-13.8. Basal armature homeomorphous, hooks falciform with stout base and strongly recurved tip, L = 15.0—22.5, 72 H. W. PALM & I. BEVERIDGE 18.8-21.3; B = 7.5-8.7, 7.0-8.0; hsr basal = 6— 7, hsr metabasal = 5-6. Remarks The postlarva from Chaerodon venustus, collected at Heron Island, is similar in scolex size and proportions, armature pattern and hook sizes to those described for M. southwelli by Palm and Walter (1999). Differences in scolex measurements of the specimen from Makaira indica are probably due to the shrunken condition of that specimen. The change in armature pattern from falciform basal hooks to heteromorphous rosethorn-shaped metabasal hooks is characteristic for M. southwelli. The distribution of the species is extended to eastern Australian waters, and Makaira indica and Chaerodon venustus represent new hosts for the species. Genus Kotorella Euzet & Radujkovic, 1989 Kotorella pronosoma (Stossich, 1901) (Fig. 54) Material examined From Dasyatis fluviorum Ogilby, 1908: 2 adults, Moreton Bay, Queensland, coll. S. Butler, 26.ix.1980 (QM G 218063-218064). Description SL = 570, 660; SW = 200, 200; pbo = 400, 390; pv = 385, 510; pb = 80, 85; vel=110, 80; BL=71 (70-73), 78 (71-88); BW = 39.4 (37.5— 40.0), 40.0 (35.3-45.0); BR = 1.8:1, 2.0:1; SP = 5.0:4.8:1, 4.6:6.0:1. TW basal = 17.5, 22.5. Basal tentacular swelling absent. Tentacle sheaths straight; TSW = 11.3—15.0, 5.5—15.0. Prebulbar organs and muscular rings around basal part of tentacle sheaths absent; retractor muscles originate in basal part of bulbs. Tentacular armature homeoacanthous, heteromorphous; basal hooks on bothridial surface L = 6.3-7.5, 6.3-7.5, B =5.0- 6.0, 5.0-6.0, smallest hooks on antibothridial surface L=2, increasing in size towards metabasal region. Small, incomplete worms to 15 mm long, maximum width 570; 90 segments. Strobila acraspedote; pre-mature segments 30 x 110 — 260 x 360, mature segments longer than wide or wider than long, 480 x 430 to 550 x 620 (Fig. 54). In second specimen, mature segments 500 x 440 to 550 x 650, and pre-gravid (with some eggs) segments 320 x 710 to 750 x 620. Genital pores submarginal, 210 from anterior margin; alternate irregularly. Cirrus sac elongate, 150-250 x 30-60 (n=5) in mature segments, length:width ratio 4.4:1 (4.0-5.0:1); cirrus unarmed; internal and external seminal vesicles absent. Vas deferens in large coils, extends to midline of segment, then posteriorly towards female genital complex. Testes 86 (81-100) in number, 45-85 in size (central), smallest testes 20-50 (peripheral), distributed in single layer, confluent posterior and anterior to ovary. Ovary median, ovarian lobes 110-200 x 100-150; Mehlis’ gland small, situated between ovarian lobes. No vitelline follicles in centre of the segment, follicles 33 (25-45) in diameter. No uterus observed, spherical eggs appear in parenchyma, egg diameter 22 (15-40). Remarks The morphology of Kotorella pronosoma (Stossich, 1901) is summarised by Euzet and Radujkovic (1989), Palm and Walter (1999) and Palm and Overstreet (2000). The present specimens from Moreton Bay correspond in scolex size and proportions as well as in the tentacular armature with the other described specimens. The sole difference noted was in testis number which exceeded 80 per segment in the specimens described. The species has previously been found in rays such as Aetobatus narinari (Euphrasen, 1790 ) (see Palm & Overstreet 2000), Himantura imbricata (Bloch & Schneider, 1801), Rhynchobatus djiddensis (see Shipley & Hornell 1906; Palm & Walter 1999) and Dasyatis pastinaca (Linnaeus, 1758) (see Euzet & Radujkovic 1989; Palm & Walter 2000). Dasyatis fluviorum represents a new host record, and the range of distribution is extended to eastern Australian waters. The present finding suggests a cosmopolitan distribution for the species. Previously, the nearest known occurrence of this cestode to Australia was from the Java Sea under the name Nybelinia narinari (MacCallum, 1917), now a synonym of Kotorella pronosoma (see Palm & Overstreet 2000). Kotorelliella gen. nov. Diagnosis: Tentacles elongate, slender; retractor muscle originates at base of bulbs. Metabasal tentacular armature homeoacanthous, heteromorphous; hooks on bothridial surface uncinate becoming slender towards antibothridial surface. Basal hooks heteromorphous; characteristic basal armature present, arranged in a heteroacanthous atypica pattern with TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 73 FIGURES 55-61. Kotorelliella jonesi gen. et sp. nov. from Taeniura lymma. Holotype, QM G 218065. 55. Scolex. 56-57. Metabasal and apical armature, external view. 58. Basal armature, bothridial view. 59. Basal armature, antibothridial view. 60-61. Basal towards metabasal armature, external view. Note the interpolated hooks as in Fig. 61. Scale bars: Figure 55, 150 um; Figures 56-61, 10 um. 74 H. W. PALM & I. BEVERIDGE interspersed hooks on the antibothridial tentacle surface. Hooks solid. Strobila unknown. Type-species: Ko. jonesi sp. nov. Kotorelliella jonesi sp. nov. (Figs 55-61) Types Holotype, 1 postlarva from the spiral valve of Taeniura lymma (Forsskal, 1775), Heron Island, Queensland, coll. I. Beveridge & M. K. Jones, 11.v11.1998 (QM G 218065). Material examined Holotype. Description Scolex compact, 4 elongated bothridia, with hook-like microtriches along bothridial borders. Posterior margins of bothridia free, not fused with peduncle. SL= 1910 (Fig. 55); SW = 390; pbo = 800; pv = 1140; pb = 260; vel = 460; BL = 259 (250-265); BW = 82.5 (80-85); BR = 3.1:1; SP =3.1:4.4:1. Tentacles long and slender; TW basal and metabasal = 21.2—23.8; TW apical = 17.5—20; basal tentacular swelling absent. Tentacle sheaths straight; TSW = 15.0— 17.7. Prebulbar organs and muscular rings around basal part of tentacle sheaths absent; retractor muscles originate in basal part of bulbs. Metabasal (Fig. 56) and apical (Fig. 57) armature homeoacanthous, heteromorphous. Rosethorn- shaped hooks with anterior extension of base cover bothridial tentacle surface (L = 13.5—15.0, B = 7.5—10.0); antibothridial surface with slender spiniform hooks, recurved at tip (L = 15.0-17.5, B = 5.0-6.3); hooks decrease in size towards apical armature; hook shape remains constant (bothridial: L = 11.3-13.7, B = 10.0-11.2; antibothridial: L = 12.5-13.7, B = 4.5-S.5). Basal armature (Figs 58-61) with additional hooks interspersed on antibothridial surface; hook pattern heteroacanthous atypica, consisting of about 5 rows of hooks on bothridial (Fig. 59) and 9-10 rows on antibothridial (Fig. 58) tentacle surface; basal hooks on bothridial surface L = 6.3-10.0, B = 6.3-8.8, on antibothridial surface L = 2.0-10.0, B = 1.5-3.8, continuously increasing in size from base of tentacle towards metabasal armature (Figs 60-61). Remarks The holotype of Kotorelliella jonesi gen. et sp. nov. has a unique form of armature within the Tentaculariidae, changing from a homeoacanthous, heteromorphous hook pattern with rosethorn-shaped hooks in the metabasal armature to a heteroacanthous atypica hook pattern in the basal armature. The size of hooks in the basal and apical regions is smaller than in the metabasal region. On the basis of the scolex morphology, Ko. jonesi sp. nov. appears to be similar to Kotorella pronosoma. Both species have an elongated scolex, four elongated and slender bothridia with free margins, and short oval bulbs. In both species, the metabasal armature is heteromorphous, with hooks changing from uncinate on the bothridial surface to elongate on the antibothridial surface. Both species have a band of hook-like microtriches along bothridial borders, which are clearly visible using light microscopy. Such a microthrix pattern is characteristic for tentaculariid trypanorhynch cestodes. The basal armature of K. pronosoma is homeoacanthous, but that of Ko. jonesi sp. nov. is unusual, having additional hooks interspersed between the oblique spirals of hooks. Thus, the basal tentacular armature is heteroacanthous atypica. The basal region is homeoacanthous in Nybelinia, Heteronybelinia and Mixonybelinia, but in Tentacularia, the bothridial surfaces of the tentacle have extra rows of small hooks (see Beveridge & Campbell 1996). The unique feature of the tentacular armature suggests that the erection of a new genus is justified, even though it is known only from the larval stage. The new species was named after one of the collectors, Dr M. K. Jones, from the Centre for Microscopy and Microanalysis, University of Queensland, Australia. Unidentified material The following additional material, some of which represent new host records, was examined but could not be assigned to a species: Nybelinia sp. from the spiral valve of Rhynchobatus djiddensis, Flat Top Island, Queensland, coll. B. G. Robertson, 25.x.1985 (SAM AHC 28325) Nybelinia sp. from the spiral valve of Notorhynchus cepedianus (Péron, 1807), south coast Kangaroo Island, South Australia, coll. B. G. Robertson, 10.ii.1985 (SAM AHC 28327) Nybelinia sp. from the stomach of Notorhynchus cepedianus, Young Rocks, Kangaroo Island, South Australia, coll. B. G. TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 15 Robertson, 23.v.1985 (SAM AHC 21354, 28328) Heteronybelinia sp. from the spiral valve of Carcharhinus amblyrhynchoides, Tommy Cut Mouth, Northern Territory, coll. B. G. Robertson, 19.ix.1986 (SAM AHC 18326). DISCUSSION The present study provides an overview for the first time of the tentaculariid trypanorhynch fauna from the Australian region. Three new species are added to the genus WNybelinia, two to Heteronybelinia and one to Mixonybelinia, with the latter two genera, as well as Kotorella, reported from the Australian region for the first time. In addition, a new genus Kotorelliella gen. nov., is erected, which is characterised by a homeoacanthous, heteromorphous metabasal armature and a heteroacanthous basal armature. Other features such as the elongated, widely 62. wy AN : {TKN Ne iS ‘ = iar NIZA I-ANICAA TZN \ZZR spaced bothridia and the short bulbs appear similar to Kotorella. The tentaculariid trypanorhynch fauna in Australian waters is relatively species rich with 22 (48 %) of the total of 46 known species occurring in these waters. Several of the new species may be endemic. However, the distribution of many species is still imperfectly known and a number of species previously thought to be endemic to a specific region have subsequently proven to be cosmopolitan (Palm et al. 1997; Palm 1999; Palm & Walter 1999, 2000). Reasons for broad distributions within tentaculariid trypanorhynchs may relate to life-cycle patterns, including widely distributed fish intermediate and elasmobranch definitive hosts. Another reason might be the low host specificity of the parasites, particularly in the intermediate host (Palm et al. 1997; Palm & Walter 2000). It is difficult to clearly distinguish between FIGURES 62-63. Schematic drawing of scolex of Heteronybelinia pseudorobusta sp. nov. from Lepidotrigla modesta, illustrating the arrangement of the different tentacle surfaces within tentaculariid trypanorhynchs. 62. View from bothridial surface, with bothridial borders merging at the apex of the scolex. 63. View from external surface, with bothridial borders widely spaced. Abbreviations, AB = antibothridial view, B = bothridial view, Bo = bothridia, E = external view, I = internal view. 76 H. W. PALM & I. BEVERIDGE some tentaculariid species solely on the basis of the tentacular armature (eg N. strongyla), and additional taxonomic characters are required. To date, the strobilae of only a few tentaculariid species have been described in detail. The present study demonstrates that, apart from the number and size of different genital structures, the cirrus sac length:width ratio might be a useful character as, for example, in H. australis sp. nov. and N. hemipristis sp. nov. The position of the anterior end of the cirrus sac and the distance of the genital pore from the anterior end of the segment, as well as the number of layers of testis, whether the testes are confluent posterior to the ovary, and the number of testes anterior to the cirrus sac appear to be useful taxonomic features. Further species descriptions are needed to show whether or not these characters are variable, as is observed in a number of scolex characters, or whether they can be used more widely in tentaculariid systematics. During this and previous studies, it became evident that the descriptions of the different tentacular surfaces of trypanorhynch cestodes used by Dollfus (1942) and Campbell and Beveridge (1994) may be difficult to apply to larval tentaculariids. In many cases it was hard to identify the various tentacle surfaces. For a better understanding of the position of the four tentacles within tentaculariid trypanorhynchs, the following two schematic illustrations are given (Figs 62, 63) as they may help to standardise the orientation of the different tentacle surfaces within tentaculariid trypanorhynchs. The determination of tentacle orientation is based upon whether the bothridial borders merge at the apex of the scolex (Fig. 62) or whether they are widely spaced (Fig. 63). The former figure shows the bothridial and the latter the external view of the tentacle surfaces. The description of Kotorelliella jonesi gen. et sp. nov. is of particular interest since its armature comprises both homeomorphous and heteromorphous components (Campbell & Beveridge 1994; Palm 1995, 1997). The species seems to be closely related to Kotorella Pronosoma, which has a similar scolex morphology and metabasal armature. Both species have characteristic microtriches along the bothridial borders, which appear similar to those of Tentacularia coryphaenae and species of Nybelinia (see Palm & Overstreet 2000). Although Beveridge et al. (1999) could not align Kotorella with the other tentaculariid genera using cladistic analyses, we consider that Kotorella pronosoma and Kotorelliella jonesi belong to the Tentaculariidae since they possess a homeoacanthous metabasal armature and, more importantly, bands of hook-like microtriches along the bothridial borders. The latter character has as yet only been described for tentaculariid trypanorhynchs. Following the classification of Dollfus (1942), only the metabasal armature is considered in classifying the tentacular armature. However, in more recent classifications, trypanorhynchs with a distinct basal armature such as Mixodigma leptaleum Dailey & Vogelbein, 1974, Paroncomegas araya (Woodland, 1934) and Mixonybelinia beveridgei (Palm, Walter, Schwerdtfeger & Reimer, 1997) have been placed in distinctive genera based in part on the basal armature (Campbell & Beveridge 1994; Campbell et al. 1999; Palm 1999). Kotorelliella jonesi has thus been placed in a new genus. Interestingly, K. jonesi represents a _ transition from homeoacanthous towards heteroacanthous armature as suggested by Campbell and Beveridge (1994, p. 57, lines 7-11). In summary, tentaculariid trypanorhynchs show considerable morphological variability, as can be seen by differences in scolex morphology between the genera Nybelinia, Heteronybelinia and Mixonybelinia, and between Kotorella and Kotorelliella. The tentacular armature also differs between the species, varying from homeoacanthous and homeomorphous (Nybelinia, Mixonybelinia) to homeoacanthous and heteromorphous (Heteronybelinia, Kotorella, Mixonybelinia) patterns. Kotorelliella has a homeoacanthous metabasal and a heteroacanthous basal hook pattern. Additional methods, such as molecular genetic studies, might help to resolve not only species relationships but also the phylogenetic relationships of the species and species groups within this large trypanorhynch family. ACKNOWLEDGMENTS Collecting was supported financially by the Australian Biological Resources Study and the Australian Research Council. Financial support was also provided by the German Research Council DFG PA 664/3-1 and 3-2. We wish to thank R. A. Bray, S. Butler, T. H. Cribb , M. K. Jones, R. Martin, M. O’Callaghan, R. Norman, J. C. Pearson, S. Pichelin, J. Reddich, B. G. Robertson , K. Sewell, P. Speare and J. Stevens for their contributions to this study. TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 77 REFERENCES BEVERIDGE, I. 1990. Taxonomic revision of Australian Eutetrarhynchidae Guiart (Cestoda: Trypanorhyncha). Invertebrate Taxonomy 4: 785- 845. BEVERIDGE, I. & CAMPBELL, R. A. 1996. New records and descriptions of trypanorhynch cestodes from Australian fishes. Records of the South Australian Museum 29: 1-22. BEVERIDGE, I., CAMPBELL, R. A. & PALM, H. W. 1999. Preliminary cladistic analysis of genera of the order Trypanorhyncha Diesing, 1863. Systematic Parasitology 42: 29-49. CAMPBELL, R. A. & BEVERIDGE, I. 1994. Order Trypanorhyncha Diesing, 1863. Pp. 51-82, in ‘Keys to the Cestode Parasites of Vertebrates’. Eds L. F. Khalil, A. Jones & R. A. Bray. CAB International: Wallingford. CAMPBELL, R., MARQUES, F. & IVANOV, V. 1999. Paroncomegas araya (Woodland, 1934) n. gen. et comb, (Cestoda: Trypanorhyncha: Eutetrarhynchidae) from the freshwater stingray Potamotrygon motoro in South America. Journal of Parasitology 85: 313-320. DOLLFUS, R.-P. 1942. Etudes critiques sur les Tétrarhynques du Muséum de Paris. Archives du Muséum national d'Histoire naturelle 19: 1466. DOLLFUS, R.-P. 1960. Sur une collection de Tétrarhynques homéacanthes de la famille des Tentaculariidae recoltées principalement dans la région de Dakar. Bulletin de l'Institut Frangais d ‘Afrique Noire, Série A 22: 788-852. EUZET, L. & RADUJKOVIC, B. M. 1989. Kotorella pronosoma (Stossich, 1901) n. gen., n. comb., type des Kotorellidae, nouvelle famille de Trypanorhyncha (Cestoda), parasite intestinal de Dasyatis pastinaca (L., 1758). Annales de Parasitologie Humaine et Comparée 64: 420-425. FROESE, R. & PAULY, D. 1998. Fish Base 98 CD ROM. International Center for Living Aquatic Resources Management: Manila. HEINZ, M. L. & DAILEY, M. D. 1974. The Trypanorhyncha (Cestoda) of the elasmobranch fishes from southern California and northern Mexico. Proceedings of the Helminthological Society of Washington 41: 161-169. JONES, M. K. & BEVERIDGE, I. 1998. Nybelinia queenslandensis sp. n. (Cestoda: Trypanorhyncha) parasitic in Carcharhinus melanopterus, from Australia, with observation on the fine structure of the scolex including the rhyncheal system. Folia Parasitologica 45: 295-311. KOROTAEVA, V. D. 1971. [Some helminth fauna of the commercial marine fishes of the sub-order Trichiuroidei in the Australia—New Zealand region]. Izvestiya Tikhooneanskogo Nauchno- Issledovatel’skogo Instituta Rybnogo Khozyaistvai Okeanografii 75: 69-84. In Russian. KOROTAEVA, V. D. 1974. [Some helminth fauna of the commercial marine fishes of the sub-order Scomberoidei in the Australian region]. /zvestiya Tikhooneanskogo Nauchno- Issledovatel’skogo Instituta Rybnogo Khozyaistvai Okeanografii 88: 61-66. In Russian. LAST, P. R. & STEVENS, J. D. 1994. ‘Sharks and Rays of Australia’. C.S.I.R.O.: Australia. LESTER, R. J. G., SEWELL, K. B., BARNES, A. & EVANS, K. 1988. Stock discrimination of orange roughy, Hoplostethus atlanticus, by parasite analysis. Marine Biology 99: 137-143. PALM, H. W. 1995. Untersuchungen zur Systematik von Riisselbandwiirmern (Cestoda: Trypanorhyncha) aus atlantischen Fischen. Berichte aus dem Institut fiir Meereskunde, Kiel 275: 1-238. PALM, H. W. 1997 An alternative classification of trypanorhynch cestodes considering the tentacular armature as being of limited importance. Systematic Parasitology 37: 81-92. PALM, H. W. 1999. Nybelinia Poche, 1926, Heteronybelinia gen. nov. and Mixonybelinia gen. nov. (Cestoda: Trypanorhyncha) in the collections of The Natural History Museum, London. Bulletin of the Natural History Museum London (Zoology series) 65: 133-153. PALM, H. W. 2000. Trypanorhynch cestodes from Indonesian coastal waters (East Indian Ocean). Folia Parasitologica 47: 123-134. PALM, H. W. & OVERSTREET, R. 2000. New records of trypanorhynch cestodes from the Gulf of Mexico, including Kotorella pronosoma (Stossich, 1901) and Heteronybelinia palliata (Linton, 1924) comb. n. Folia Parasitologica 47: 293-302. PALM, H. W. & WALTER, T. 1999. Nybelinia southwelli sp. nov. (Cestoda: Trypanorhyncha) with the re-description of N. perideraeus (Shipley & Hornell, 1906) and synonymy of N. herdmani (Shipley & Hornell, 1906) with Kotorella pronosoma (Stossich, 1901). Bulletin of the Natural History Museum London (Zoology series) 65: 123- 131. PALM, H. W. & WALTER, T. 2000. Tentaculariid cestodes (Trypanorhyncha) from the Muséum national d’Histoire naturelle, Paris. Zoosystema 22: 641-666. PALM, H. W., WALTER, T., SCHWERDTFEGER, G. & REIMER, L. W. 1997. Nybelinia Poche, 1926 (Cestoda: Trypanorhyncha) from the Mocambique coast, with description of N. beveridgei sp. nov. and systematic consideration on the genus. South African Journal of Marine Science 18: 273-285. PINTNER, T. 1927. Kritische Beitrage zum System der 78 H. W. PALM & I. BEVERIDGE Tetrarhynchen. Zoologische Jahrbiicher 53: 559- 590. REIMER, L. W. 1980. Larven der Ordnung Trypanorhyncha (Cestoda) aus Teleostiern des Indischen Ozeans. Angewandte Parasitologie 21: 221-231. SEWELL, K. B. & LESTER, R. J. G. 1988. The number of selected parasites in Australian and New Zealand samples of orange roughy Hoplostethus atlanticus, 1983-1986. Technical Report, Department of Sea Fisheries, Tasmania, 26: 1-38. SEWELL, K. B.& LESTER, R. J. G. 1995. Stock composition and movement of gemfish, Rexea solandri, as indicated by parasites. Canadian Journal of Fisheries and Aquatic Sciences 52, Supplement 1: 225-232. SHIPLEY, A. E. & HORNELL, F. L. S. 1906. Report on the cestode and nematode parasites from the marine fishes of Ceylon. Ceylon Pearl Oyster Fisheries Marine Biological Reports, Part 5: 43-96. SPEARE, P. 1999. Parasites from east-coast Australian billfish. Memoirs of the Queensland Museum 43: 837-848. VIJAYALAKSHMI, C., VISAYALAKSHMI, J. & GANGADHARAM, T. 1996. Some trypanorhynch cestodes from the shark Scoliodon palasorrah (Cuvier) with the description of a new species, Tentacularia scoliodoni. Rivista di Parassitologia 13: 83-89. TWO NEW SPECIES OF THE GENUS LEPANUS BALTHASAR FROM SOUTH AUSTRALIA (COLEOPTERA : SCARABAEIDAE) E. G. MATTHEWS & T. A. WEIR Summary Ball-rolling dung beetles of the tribe Scarabaeini are recorded for the first time from South Australia. The new species Lepanus penelopae and L. loftyensis are described from southern Eyre Peninsula and the Mount Lofty Block respectively, and are compared with other species of the genus. There are brief remarks comparing Lepanus Balthasar with the related genera Sauvagesinella Paulian and Aptenocanthon Matthews. TWO NEW SPECIES OF THE GENUS LEPANUS BALTHASAR FROM SOUTH AUSTRALIA (COLEOPTERA: SCARABAEIDAE) E. G. MATTHEWS & T. A. WEIR MATTHEWS, E. G. & WEIR, T. A. 2002. Two new species of the genus Lepanus Balthasar from South Australia (Coleoptera: Scarabaeidae). Records of the South Australian Museum. 35(1): 79-84. Ball-rolling dung beetles of the tribe Scarabaeini are recorded for the first time from South Australia. The new species Lepanus penelopae and L. loftyensis are described from southern Eyre Peninsula and the Mount Lofty Block respectively, and are compared with other species of the genus. There are brief remarks comparing Lepanus Balthasar with the related genera Sauvagesinella Paulian and Aptenocanthon Matthews. E. G. Matthews, South Australian Museum, North Terrace, Adelaide, South Australia S000; and T. A. Weir, CSIRO Division of Entomology, P.O. Box 1700, Canberra, Australian Capital Territory 2601. Manuscript received 8 August 2001. At the time of the revision by Matthews (1974) of the Australian Scarabaeini, generally known as ball-rolling dung beetles, it was believed that this group was absent from South Australia, although it was known from Victoria and Western Australia. However, in 1979 P. Greenslade obtained specimens of an undescribed species of the genus Lepanus Balthasar, which belongs to this tribe, in the Marble Range of southern Eyre Peninsula; and in 1982 another undescribed species of the same genus was collected during extensive pitfall trapping undertaken by J. and P. Greenslade in Kuitpo Kyeema Forest, southern Mount Lofty Ranges, in connection with a project to determine the effects of fire on the forest-floor fauna. Then in 1999 and 2000, more specimens of the second species were picked up by members of the Biological Survey of South Australia in the area of Mount Remarkable at the extreme northern end of the Mount Lofty Block environmental province, as well as near the Barossa Valley, using unbaited pitfall traps. Altogether, the two species are now known from five collection localities (Fig. 1), all situated on low mountain ranges at altitudes from 300 to just over 500 m. Kangaroo Island is part of the Mount Lofty Block but pitfall traps baited with human faeces, set throughout the island by one of the authors (EGM) in November 1990, failed to collect any Lepanus. Lepanus is known from New Guinea, where it was first described, and from densely vegetated mesic and humid habitats along the northern, eastern and southern coasts of Australia as far west as the tingle forest near Walpole, Western Australia (Matthews 1974). The habitat is leaf litter, and food, where known, consists of vertebrate faecal matter. The descriptions below bring the total number of described Australian species of Lepanus to 23, but several undescribed species have been collected in recent years in eastern montane forests (R. Storey, pers. comm.). Specimens are deposited in the South Australian Museum, Adelaide (SAMA); the Australian National Insect Collection, Canberra (ANIC); the Queensland Museum (QMBA); and the Queensland Department of Primary Industries collection at Mareeba (DPIM). All figures except the map are by the senior author. DESCRIPTIONS OF NEw SPECIES Lepanus penelopae sp. nov. (Figs 1, 2, 3, 5, 7, 9, 12) Holotype ‘S. Aust. Eyre Pen. Marble Rge. Dense broombush. Pitfall. 4.10.79. P.J.M. Greenslade’, male, SAMA. Paratypes Same data as holotype, 1 male and 5 females, ANIC, SAMA. Description Body uniformly piceous and nitid, legs rufous. Total length 5.0-6.0 mm. Maximum width across 80 E. G. MATTHEWS & T. A. WEIR FIGURE 2. Lepanus penelopae male, habitus. Scale line 1 mm. elytra 3.4—-3.6 mm. Head — Dorsal surface even, very densely punctate with small shallow punctures not running together, some bearing short, very fine recumbent setae. Dorsal part of eye small, its maximum width contained about 25 times in interocular distance. Prothorax — Anterior angles subquadrate. Sides of pronotum rounded, widest at broadly rounded posterior angles. Dorsal surface very densely and uniformly finely punctate, punctures separated by distances equal to a little more than their diameter, glabrous. Elytra — Striae shallowly impressed with regularly spaced moderately deep punctures separated by 2- 4 diameters. Intervals nearly flat, smooth with very slightly uneven surface, glabrous, with minute punctures. Hind wings — Absent. Sterna — Meso- and metasterna impunctate medially with large shallow cicatricose punctures laterally, glabrous. Legs — Protibia with three large teeth on outer edge, which is serrate between teeth and proximal to them. Claws (Fig. 7) a little expanded basally but not distinctly angulate or dentate. Abdomen — Pygidium (Fig. 9) with a very deep TWO NEW SPECIES OF LEPANUS BALTHASAR 81 3. FIGURES 3-6. Tibiae of males in dorsal view. 3, L. penelopae, left protibia; 4, L. loftyensis, right protibia; 5, L. penelopae, right metatibia; 6, L. loftyensis, right metatibia. b, brush of stiff setae; c, comb of flattened bristles. Scale line 1 mm. basal groove across whole width of disc, groove narrower and shallower in middle, surface inside it transversely striated. Disc very finely punctate, glabrous. Aedeagus as in Fig. 12. Sexual dimorphism — Male with rounded expansion of inner apical end of protibia bearing apical comb of short wide flattened bristles (Fig. 3, c) and on lower surface a brush of long setae which is directed outwardly parallel to surface, extending beyond edge of comb (Fig. 3, b). Metatibia of male (Fig. 5) a little expanded at inner apical edge and bearing small apical tooth there. Remarks In the key to the species of Lepanus published by Matthews (1974), L. penelopae will go to couplet 16 because it has three protibial teeth, and then to couplet 18 because it has a transverse groove on the pygidium and glabrous dorsal surfaces. However, it does not have dentate claws as also required by this couplet. It can then be forced through couplets 19 and 20 to end up with L. illawarrensis Matthews of New South Wales, which appears to be the most closely related species, sharing many aspects of structure and secondary sexual characters. L. penelopae differs from L. illawarrensis in having simple claws, non- geminate elytral striae with much deeper strial punctures, the pygidial groove more strongly narrowed in the middle, the basal ridge of the pygidium straight in the middle, no trace of a median tubercle on the metasternum of the male, male hind tibia not twisted, and somewhat differently shaped aedeagal parameres. Etymology This species is named after Penelope Greenslade, who collected it as well as many of the other specimens of Lepanus described in this paper. Lepanus loftyensis sp. nov. (Figs 1, 4, 6, 8, 10, 11) Holotype ‘S. AUST. Pitfalls Kuitpo Kyeema Forest 15th March, 1982 P. Greenslade’, male, SAMA. Paratypes Same data as holotype, 12 males, 4 females, ANIC, SAMA. ‘S. Aust. Deep Ck Boat Hbr Lane 82 E.G. MATTHEWS & T. A. WEIR 9 FIGURES 7-9. 7, L. penelopae, claw segment of metatarsus; 8, ditto, L. loftyensis. Scale line 0.25 mm; 9, L. penelopae, pygidium of male. Scale line 1 mm. turnoff pitfalls in heath 1-8 Dec 1983 P. Greenslade’, 1 female, SAMA. ‘S. AUST. Mt Remarkable NP 11 km SSW Wilmington 32° 44°56”S 138°04’03”E 20-25 Nov 99 pitfall Flinders Ra. MAMO00401’, 5 females, ANIC, SAMA. ‘S. AUS. 11 km SE Whispering Wall (near Para Wirra) 34°34’54”S 138°55’12”E 27-10-00 SMLR Survey BAR 03201’, 17 males, 31 females, SAMA, QMBA, DPIM. Description Body uniformly fuscous to piceous, legs rufous. Total length 3.4-5.5 mm, maximum width across elytra 2.3-3.6 mm. Head — Surface even, very densely punctate with shallow coarse punctures running together in irregular transverse rows, bearing small recumbent setae. Dorsal part of eye small, slit-like, its maximum width contained about 30 times in interocular distance. Prothorax — Anterior angles obtuse. Sides of pronotum widest about one-quarter of the distance behind anterior angles, thereafter slightly converging or subparallel to broadly rounded posterior angles. Dorsal surface very densely and uniformly punctate, punctures separated by distances equal to less than their diameter, bearing very fine, short, fully recumbent setae. Elytra — Striae very superficial, geminate with slightly crenulate edges, impunctate. Discal intervals flat, in same plane as striae, with more or less undulate surface, finely shagreened, with a row of punctures bearing recumbent setae along edges of each interval, an additional row usually present in middle of intervals near base. Hind wings — Atrophied, represented by short stubs. Sterna — Mesosternum with shallow punctures on posterior half. Metasternum entirely densely punctate with shallow punctures laterally bearing short, fine recumbent setae. Legs — Protibia (Fig. 4) with 3 large teeth on outer edge, which is serrate between teeth and proximal to them. Claws (Fig. 8) strongly dentate. Abdomen — Pygidial disc simple, without groove, uniformly densely and shallowly FIGURE 10. Lepanus loftyensis male, habitus. Scale line 1 mm. TWO NEW SPECIES OF LEPANUS BALTHASAR 83 punctate, punctures bearing very small recumbent setae usually worn off. Aedeagus as in Fig. 11. Sexual dimorphism — Male with rounded expansion of inner apical end of protibia bearing apical comb of short, wide, flattened bristles (Fig. 4, c) and a brush of long setae underneath which is directed downward perpendicular to surface. Metatibia of male (Fig. 6) with strongly recurved apical end bearing acute inner tooth. Prothorax of male broader anteriorly, subquadrate; that of female slightly narrower anteriorly. Clypeal teeth of male slightly smaller than those of female. Remarks In the key to species of Lepanus in Matthews (1974) L. loftyensis will first go to couplet 16 because it has three teeth on the outer edge of the protibia, then to couplet 17 because it has a simple pygidium and setose dorsal surfaces, then to L. villosus Matthews of north Queensland because of its crenulate elytral striae and densely punctate meso- and metasterna. It differs from L. villosus in having superficial elytra striae, reduced hind wings, very different parameres of the aedeagus, and being of much larger size. Further, L. loftyensis has the setae on the pronotum only about half the length of those on the elytra, the first elytral interval with a double row of seta- bearing punctures, and the upper edges of the epipleural carina not visible from directly above due to the curvature of the elytra. The designation MAM 00401 on labels of the Mount Remarkable specimens refers to a locality 11 which is described in Brandle (2001) as mountainous, on a hill crest, with loam/sand soil and an overstorey of Allocasuarina verticillata at a cover density of 25-50%, and at an altitude above sea level of 430m. For the Parra Wirra specimens the designation BAR 03201 refers to a ridge top covered with Eucalyptus fasciculosa / Acacia paradoxa woodland and an understorey of Calytrix, Hakea and Xanthorrhoea, at an altitude of 515 m (L. Queale, pers. comm.). Etymology The name refers to the Mount Lofty Block environmental province in which all four collection localities of this species are contained. The block is Province number 3 as delineated in the Biological Survey of South Australia, and extends from Kangaroo Island northward to Mount Remarkable. The biota of this province is characterised as fully Bassian (Brandle 2001). DISCUSSION The existence of the species here described as Lepanus loftyensis was briefly noted by Matthews (1984: 6, footnote) erroneously as an undescribed species of Sauvagesinella Paulian. Lepanus and Sauvagesinella are closely related genera which are not separable in the female sex, but in the male there are several secondary sexual characters which will distinguish them. Unlike Lepanus, Sauvagesinella males have a prominent median 12 FIGURES 11 & 12. Aedeagi in right and left views. 11, L. loftyensis; 12, L. penelopae. Scale line 1 mm. 84 E. G. MATTHEWS & T. A. WEIR tubercle on the metasternum (only a trace of a tubercle in some Lepanus), a row of close-set tubercles along the inner edge of the metatibia, and a more or less compressed metafemur with a longitudinal fold or ridge on its ventral surface. Sauvagesinella as understood here does not occur outside the extreme southern part of Western Australia, where there are three species (Matthews 1974). Storey and Monteith (2000) mention that males of all species of Aptenocanthon Matthews, where known, have a similar form of the fore tibial apex with its inner angle expanded and bearing a short brush of stiff setae bent downwards at right angles to the upper tibial surface. This is a similar situation to that found in the two species of Lepanus described above (eg Fig. 3, b) and is also known to occur in several other species of Lepanus and all three species of Sauvagesinella. As well as this brush, there is a comb of flattened bristles at the protibial apex (Figs 3 and 4, c) which occurs, to varying degrees, in all known males of Aptenocanthon and Sauvagesinella as well as some species of Lepanus. Clearly, these male characters are not limited to Aptenocanthon but their value in classification must await further studies on the interrelationships of these genera of Scarabaeini, as also pointed out by Storey and Monteith (2000). In South Australia the only other known representatives of the dung beetle subfamily Scarabaeinae belong to the tribes Onthophagini (native and introduced Onthophagus Latreille), Onitini (introduced Onitis Fabricius) and Oniticellini (introduced Euoniticellus Janssens), none of which make or roll food balls. All these groups are differentiated from the tribe Scarabaeini by their middle and hind tibiae, which are strongly expanded apically. In Scarabaeini these tibiae are slender and only a little expanded, an adaptation for rolling the food material. REFERENCES BRANDLE, K. 2001. A biological survey of the Flinders Ranges, South Australia 1997-1999, Biodiversity Survey and Monitoring, National Parks and Wildlife, South Australia, Department for Environment and Heritage, pp. i-xviii, 1-455. MATTHEWS, E. G. 1974. A revision of the scarabaeine dung beetles of Australia. II. Tribe Scarabaeini. Australian Journal of Zoology, Supplementary Series 24: 1-211. MATTHEWS, E. G. 1984. A guide to the genera of beetles of South Australia. Part 3. South Australian Museum, Special Educational Bulletin Series 6: 1- 60. STOREY, R. I. & MONTEITH, G. B. 2000. Five new species of Aptenocanthon Matthews (Coleoptera: Scarabaeidae: Scarabaeinae) from tropical Australia, with notes on distribution. Memoirs of the Queensland Museum 46(1): 349-358. A CHECKLIST OF CISSEIS (SENSU STRICTO) GORY & LAPORTE, 1839 (COLEOPTERA : BUPRESTIDAE : AGRILINAE) SHELLEY BARKER Summary A checklist of Cisseis (s.s.) Gory & Laporte, 1839 is presented and discussed, summarising the information available on the genus at the present time. A CHECKLIST OF CISSEIS (SENSU STRICTO) GORY & LAPORTE, 1839 (COLEOPTERA: BUPRESTIDAE: AGRILINAE). SHELLEY BARKER BARKER, S. 2002. A checklist of Cisseis (sensu stricto) Gory & Laporte, 1839 (Coleoptera: Buprestidae: Agrilinae). Records of the South Australian Museum 35(1): 85-90. A checklist of Cisseis (s.s.) Gory & Laporte, 1839 is presented and discussed, summarising the information available on the genus at the present time. S. Barker, Department of Entomology, South Australian Museum, North Terrace, Adelaide, South Australia 5000. Manuscript received 3 December 2001. MATERIAL Specimens examined came from the following institutions: AMSA - Australian Museum, Sydney ANIC — Australian National Insect Collection, CSIRO, Canberra BMNH - The Natural History Museum, London BPBM - B. P. Bishop Museum, Honolulu HMOE - Hope Museum, Oxford HUMB -— Humboldt University Museum, Berlin MCSNG —- Museo Civico di Storia Naturale Genoa, Italy MNHN -— Museum National d’ Histoire Naturelle, Paris NMVA — National Museum of Victoria, Melbourne PMCE — National Museum of Czech Republic, Prague QMBA — Queensland Museum, Brisbane RMBB - Royal Museum, Brussels SAMA -— South Australian Museum, Adelaide. Abbreviations: WA — Western Australia NWA - Kimberly area, Western Australia Q — Queensland NSW — New South Wales V — Victoria SA — South Australia T — Tasmania NT — Northern Territory. INTRODUCTION Many of the Australian buprestid genera have not been reviewed since the death of H. J. Carter in 1941. Because of the large amount of subsequent collecting and confusion with species identification, the genera are in a parlous state of order. One of worst has been Cisseis (Coleoptera: Buprestidae: Agrilinae), last researched by Carter (1923, 1929). With the intention of revising Cisseis, the author commenced work on the group in 1997. All of the readily available types, more than half of which are housed in European museums, have now been examined. Some of the difficulties have been resolved and a number of new species described (Barker 1998, 1999a, 1999b, 2001). Some species clearly did not belong in Cisseis and these, together with nine new species, were placed in a new genus Stanwatkinsius Barker & Bellamy (2001). Carter (1923) did not examine male genitalia, which the author has found to be highly diagnostic. As a result, many of his specimens located in various museum collections are incorrectly identified. In order to summarise the knowledge available on the genus, a checklist has been constructed of valid species together with their distributions by state or country. Types and their locations are also listed. The following abbreviations are used: HT = holotype; ST = syntype; LT = lectotype; AT = allotype. CHECKLIST OF CISSEIS (SENSU STRICTO) GORY & LAPORTE, 1839. aberrans Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 98. (6 HT, Woodridge, WA, H. Demarz, SAMA I 21504). WA. 86 S. BARKER acuducta (Kirby, 1836)(Trachys). Faun. Bor. Amer.: 162. (6 HT, N. Scotia, BMNH). Q, NSW, V, SA, T. marmorea (Gory & Laporte, 1839)(Ethon). Mon. Bupr. ii: 4. (HT, MNHN not seen) cuprifrons Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 157. (6 HT, Australia, Standing, BMNH) laeta Kerremans, 1903. in Wytsman Genera Ins. fas. 12: 227. (o HT, Australia, Moffarts, BMNH) adusta Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 101. (¢ HT, Lucindale, SA, Feueheerdt, SAMA I 21505). SA. albertisi Gestro, 1877. Ann. Mus. Civ. Stor. Nat. Genova. 9: 357. (3 ST sex unknown, Somerset, Australia, D’Albertis, 1/75 MCSNG; @ ST, Somerset, Australia, RMBB). Q. albosparsa Gory & Laporte, 1839. Monograph ii: 3. (HT, MNHN not seen). NT, Q. cupriventris Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 161. (& HT, Australia, Chevrolat, BMNH). inflammata Carter, 1923. Proc. Linn. Soc. N.S.W. 48: 167. (S HT, S. Johnstone R., Q, H. W. Brown AMSA K67240) (new syn.). aquilonia Bellamy, 1991. Tijdschr. v. Ent. 134: 171. (6 HT, BPBM not seen). Philippine Arch. armstrongi Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 101. (¢ HT, Bogan R., NSW, J. Armstrong, NMVA). NSW. augustgoerlingi Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 102. (6 HT, Marloo Stn., Wurarga, WA, A. Goerling, ANIC). WA. aurocyanea Carter, 1934. Proc. Linn. Soc. N.S.W. 59: 258. (2 HT, Fletcher, Q, E. Sutton, AMSA K67296). NSW, Q. bedfordi Obenberger, 1923. Casopis Cs. Spol. Entom. 32: 9. (2 HT, Cape Bedford, Q, NMPC). Q. brooksi Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 102. (6 HT, Mareeba, Q, J. G. Brooks, ANIC). Q. broomensis Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 103. (6 HT, Broome, WA, H. W. Brown, SAMA I 21509). NWA. browni Carter, 1934. Proc. Linn. Soc. N.S.W. 59. 258. (6 HT, Dedari, WA, H. W. Brown, AMSA K67297). WA. carterella Obenberger, 1935. Col. Catalogus 12: 846. (2 LT, Herberton, Q, C. J. Wild, QMBA)(replac. name). Q. elliptica var. frontalis Carter, 1923. Proc. Linn. Soc. N.S.W. 48: 170. (homonym). carteri Obenberger, 1924. Archiv. fiir Naturg. 90: 109. (3 LT, Yilgarn, WA, NMPC). WA. chalcophora Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 103. (¢ HT, Kapunda, SA, S. Barker, SAMA I 21507). SA. clermonti Théry, 1945. Bull. Soc. Ent. Fr. 50: 46. (HT, MNHN not seen). Bougainville. corpulenta Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 104. (6 HT, Tallering stn., Pindar, WA, S. Barker, SAMA I 21511). WA. cupreicollis (Hope, 1846)(Ethon). Ann. Mag. Nat. Hist. 17: 64. (6 HT, Moriatta (sic), SA, HMOE no. 893). SA aenicollis (Hope, 1846)(Ethon). Ann. Mag. Nat. Hist. 17: 65. (2 HT, Adelaide, HMOE no. 892 2/2). cupreola Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 10S. (o HT, Mt Spec, Q, J. G. B., ANIC). SA, NSW, Q. cupripennis (Guerin, 1836)(Buprestis). Voy. Coquille: 65. (HT, MNHN not seen). NSW, V. semiscabrosa Thomson, 1879. Typi Bupr. Mus. Thomson. App. 1A: 53. (3d HT, Sydney)(new syn.). nitidicollis Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 162. (6 HT, Australie, Bates, BMNH)(new syn.). cyanea Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 105. (¢ HT, Wialki, WA, S. Barker, SAMA I 21512). WA. cyaneopyga Carter, 1923. Proc. Linn. Soc. N.S.W. 48: 170. (2 LT, Lake Austin, WA, H. W. Brown, AMSA K67292). WA. goerlingi Carter, 1936. Proc. Linn. Soc. N.S.W. 61: 104. (2 HT, Marloo stn., Wurarga, WA, A. Goerling, AMSA K67466)(new syn.). cyanura Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 163.(2 dd ST & 2 ST, BMNH). Q. derbyensis Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 106. (¢ HT, Derby, H. W. Brown, SAMA I 21520). NWA. duodecimmaculata (Fabricius, 1801)(Buprestis). Sys. Eleuth.: 191. (HT sex unknown, BMNH). All mainland states. duodecimguttata (Boisduval, 1835) (Buprestis). Voy. Astrolabe, Entom. 2: 93. (HT, MNHN not seen). quaturodecimnotata Hope, 1846. Ann. Mag. Nat. Hist. 17: 64. (HT not seen, HMOE). pustulata Thomson, 1879. Typi Bupr. Mus. CHECKLIST OF CISSEIS (S.S.) GORY & LAPORTE, 1839 87 Thoms. App 1A: 51. (HT sex unknown, MNHN). biologia Froggatt, 1910. Australian Insects: 165. (HT not seen, AMSA?)(new syn.). fallaciosula Obenberger, 1935. Acta. Soc. Ent. Csl. 32: 36. (HT sex unknown, PMCE)(new syn.). elongatula Blackburn, 1888. Proc. Linn. Soc. N.S.W. 3: 892. (6 HT, NT, SAMA). NT. elliptica Carter, 1923. Proc. Linn. Soc. N.S.W. 48: 170. (2 LT, Cue, WA, H. W. Brown, NMVA). WA. excelsior Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 107. (d HT, Mt Carbine, Q, T. M. S. Hanlon & M. Powell, SAMA I 21514). Q. ernestadamsi Barker, 1999. Rec. S. Aust. Mus. 32: 46. (6 HT, Edungalba, Q, E. E. Adams, ANIC). Q. fascigera Obenberger, 1919. Ent. Mitteilungen 8: 20. (6 HT, New Guinea, PMCE no. 23772). New Guinea. fraterna Kerremans, 1900. Mem. Soc. Ent. Belg. 7: 80. (2 HT, Germ. N. G., Webster, BMNH). New Guinea. frontalis Kerremans, 1898, Ann. Soc. Ent. Belg. 42: 157. (S HT, New Guinée, Standing, BMNH). New Guinea. fulgidicollis Macleay, 1888. Proc. Linn. Soc. N.S.W. 3: 1231. (2 do ST, Derby, NWA, ANIC). NWA, Q. gibbera (Carter, 1937)(Neospades). Trans. Roy. Soc. S. Aust. 61: 121. (2 HT, Mullaly, NSW, H. J. Carter, AMSA K111950). NSW. gouldi (Hope, 1846)(Ethon). Ann. Mag. Nat. Hist. 17: 65. (2 HT, Port Essington, HMOE no. 889). NT. heroni Carter, 1934. Proc. Linn. Soc. N.S.W. 59: 259. (6 HT, Dorrigo, NSW, W. Herron, AMSA K67298). NSW. impressicollis Macleay, 1878. Ent. Soc. N.S.W. 2: 248. (2 HT, Gayndah, AMSA K32713). Q. viridiaurea Macleay, 1878. Ent. Soc. N.S.W. 2: 248. (2 2 2 ST, Gayndah, AMSA no. K32712)(new syn.). viridicuprea Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 160. (2 HT, Cocktown (sic), Standing, BMNH)(new syn.). nitida Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 164. (6 HT, Cocktown (sic), BMNH)(new syn.). inops Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 168. (2 HT, Australie, Standing, BMNH). NSW. kohouti Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 107. (6 HT, Canberra, K. Pullen, ANIC). NSW, Q. laticollis Carter, 1923. Proc. Linn. Soc. N.S.W. 48: 169. (2 HT, BMNH). Q. leucosticta (Kirby, 1818)(Buprestis). Trans. Ent. Soc. Lond. 12: 382. (2 HT, BMNH). All mainland states. aurulenta Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 159. (@ HT, Australia, Chevrolat, BMNH)(new syn.). fulgidifrons Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 161. (2 HT, Australia, Chevrolat, BMNH). macmillani Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 108. (6 HT, Wanneroo, WA, R. P. McMillan, SAMA I 21516). WA. macqueeni Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 108. (6 HT, Milmerran, Q, J. Macqueen, ANIC). Q. maculata (Gory & Laporte, 1839)(Ethon). Mon. Bupr. ii: 5. (HT, MNHN not seen). NSW. marmorata Gory & Laporte, 1839. Mon. Bupr. ii: 4. (HT, MNHN not seen). NSW, Q. viridicollis Thomson, 1879. Typi Bupr. Mus. Thomson. App. 1A: 50. (6 HT, MNHN). aena Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 158. (3 22 ST, Sydney, Bates, BMNH). minutissima Thomson, 1879. Typi Bupr. Mus. Thomson. App. 1A: 54. (HT, MNHN not seen). SA, V. simplex Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 171. (2 dd ST & Q ST, Australie, Standing, BMNH)(new syn.). modesta Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 170. (S HT, Australie, Stark, Chevrolat collection, BMNH). WA, SA, V. myallae Carter, 1934. Proc. Linn. Soc. N.S.W. 59: 260. (2 HT, Bogan R., NSW, J. Armstrong, AMSA K67299). NSW. nigrita Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 168. (d HT, Australie, Fairmaire, BMNBH). nigripennis Macleay, 1888. Proc. Linn. Soc. N.S.W. 3: 1231. (¢ HT, Barrior (sic) Range, NWA, Froggett, ANIC). NT, WA. ignicollis Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 164. (od ST, Australie, Standing, BMNH; o ST, Australie, Chevrolat, BMNH; 2 doo ST, Port Denison BMNH)(new syn.). nigromaculata Kerremans, 1895. Ann. Soc. Ent. Belg. 39: 218. (HT not in BMNH). Malacca. 88 S. BARKER nitidiventris Carter, 1934. Proc. Linn. Soc. N.S.W. 59: 260. (2 HT, Gosford, NSW, N. MacGregor, AMSA K67300). NSW, Q. niveosparsa (Carter, 1927)(Neospades). Proc. Linn. Soc. N.S.W. 52: 228. (6 HT, Bogan R., NSW, J. Armstrong, AMSA K67288). NSW, SA. notulata (Germar, 1848)(Ethon). Linn. Ent.: 178. (3d HT, Adelaide, HUMB). SA, NSW, T. atroviolacea Thomson, 1879. Typi. Bupr. Mus. Thoms. App. 1A: 52. (6 ST & @ ST, Austr, MNHN)(new syn.). morosa Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 158. (2 d6¢ ST, Australie, Stark, BMNH)(new syn.). nubeculosa (Germar, 1848)(Ethon). Linn. Ent.: 176. (2 LT, Adelaide, HUMB). SA, V. chalcoptera (Germar, 1848)(Ethon). Linn. Ent.: 177. (2 36 3 ST, Adelaide, HUMB). similis Saunders, 1871. Cat. Bupr.: 103. (2 HT, Adelaide, BMNH)(new syn.). oblonga Kerremans, 1903. in Wytsman Genera Ins. fas. 12: 229. (2 HT, Australie, Fairm., BMNH). V, NSW, Q, T. obscura Blackburn, 1887. Trans. R. Soc. S. Aust. 10: 252. (2 HT, T 320, type locality unknown, BMNH)(unique). opima Thomson, 1879. Typi Bupr. Mus. Thomson. App. 1A: 50. (@ HT, Champion Bay, MNHN). WA. ovalis Carter, 1923. Proc, Linn. Soc. N.S.W. 48: 170. (2 HT, Nov. Holl. occid., Fry Coll. 1905.100, BMNH 37811)(unique). WA. parva Blackburn, 1887. Trans. R. Soc. S. Aust. 10: 253. (6 HT, BMNH). WA, SA, V, NSW. pygmaea Blackburn, 1891. Trans. R. Soc. S. Aust. 14: 299. (2 HT, Victoria, C. French, NMVVA)(new syn.). patricia Carter, 1935. Proc. Linn. Soc. N.S.W. 55: 180. (d6 HT, Bunbury, WA, F. L. Whitlock, AMSA K67302). WA, V, NSW. pauperula Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 167. (do HT, Australie, A. Deyr., Chevrolat collection, BMNH). T. prasina Carter, 1923. Proc. Linn. Soc. N.S.W. 48: 168. (6 LT, Australia, Blackburn’s collection, 3267, SAMA). SA, Q. puella Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 170. (2 66 ST & @ ST, Australie, Fairm., BMNH). Q. curta Kerremans, 1903. in Wytsman Genera Ins. fas. 12: 229. (6 HT, Peak Downs, Fairmaire, Mus. Godeffroy no. 11142, BMNB). pulchella Carter, 1923. Proc. Linn Soc. N.S.W. 48: 171. (HT sex unknown, French collection, NMVA). Q. pulleni Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 109. (6 HT, Edungalba, Q, S. Barker, SAMA I 21518). ACT, Q. regalis Thomson, 1879. Typi Bupr. Mus. Thomson. App. 1A: 50. (6 HT, Queensland, MNHN). NT, Q. robertfisheri Barker, 1999. Rec. S. Aust. Mus. 32: 47. (d HT, Melrose, SA, R. H. Fisher, SAMA I 21407). SA. roseocuprea (Hope, 1846)(Ethon). Ann. Mag. Nat. Hist. 17: 64. (2 HT, Moriatta (sic), SA, HMOE no. 889). WA, SA, V, NSW. dispar Blackburn, 1891. Trans. R. Soc. S. Aust. 14: 297. (2 HT, 3832, BMNH). fairmairei Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 169. (2 HT, Australie, Fairmaire, BMNH). cuprea Kerremans, 1903. in Wytsman Genera Ins. fas. 12: 228. (2 HT, NSW, Standing, BMNB). rubicunda Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 169. (¢ HT, Australie, Fairm., BMNH). WA, SA, V, NSW, Q. undulata Kerremans, 1903. in Wytsman Genera Ins. fas. 12: 227. (2 HT, NSW, Standing, BMNH)(new syn.). purpurea Kerremans, 1903. in Wytsman Genera Ins. fas. 12: 228. (2 HT, Sidney (sic), Fairm, Mus. Godeffroy no. 11108)(mew syn.). scabiosa (Boisduval, 1835)(Buprestis). Voy. Astrolobe Entom. 2: 96. (HT, MNHN not seen). scabrosula Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 167. (3 63d ST & @ ST, Australie, Fairm., BMNH). NSW, Q. semiobscura Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 170. (o ST, Australie, Fairm., BMNH; 2 ¢@ ST, Australie, Deyr., BMNH). SA, V. septuosa Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 109. (6 HT, Charity Ck Bridge, Manning R., NSW, S. Watkins, SAMA I 21521). SA, NSW, Q. sexnotata Fauvel, 1891. Rev. d’Ent. 10: 181. (HT, MNHN not seen). New Caledonia. signaticollis (Hope, 1846)(Ethon). Ann. Mag. Nat. Hist. 17: 64. (d6 HT, Port Essington, HMOE no. 891). WA, NT, Q. speciosa Barker, 2001. Trans. Roy. Soc. S. Aust. CHECKLIST OF CISSEIS (S.S.) GORY & LAPORTE, 1839 89 125: 110. (6 HT, Midland Jctn, WA, R. P. McMillan, SAMA I 21523). WA. stellata Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 111. (6 HT, Cairns, SAMA I 21524). Q. stigmata Gory & Laporte, 1839. Mon. Bupr. ii: 3. (HT, MNHN not seen). WA. subbifascialis Carter, 1927. Proc. Linn. Soc. N.S.W. 52: 229. (2 HT, Bogan R., NSW. J. A., AMSA K67291). NSW. tasmanica Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 165. (2 ST, Tasmanie, Oberthur, BMNH; ST, Tasmanie, Chevrolat, BMNH). T, V, NSW. trimentula Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 111. (6 HT, Pilliga East SF, NSW, S. Watkins, SAMA I 21526). V, NSW, Q. tyrrhena Carter, 1923. Proc. Linn. Soc. N.S.W. 48: 168. (2 HT, Kalamunda, WA, H. M. Giles, AMSA K67293). WA vicina Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 164. (6 HT, Australie, Standing, BMNH). V, NSW, Q. collaris Kerremans, 1903. in Wytsman, Genera Ins. fas. 12: 229. (2 36 ST, Gayndah, Fairmaire, BMNH). ornata Kerremans, 1903. in Wytsman Genera Ins. fas. 12: 229. (d HT, NSW, Standing, BMNH). violacea Kerremans, 1903. in Wytsman Gen. Ins. fas. 12: 228. (6 ST & 2 ST, Gayndah, Fairmaire, BMNH). NSW, Q. viridiceps Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 168. (6 HT, Australie, Standing, BMNH). NSW. viridipurpurea Carter, 1924. Proc. Linn. Soc. N.S.W. 49: 27. (2 36 ST, Geraldton, WA, J. Clark, AMSA K67294). WA. sapphira Carter, 1933. Proc. Linn. Soc. N.S.W. 58: 163. (2 HT, Moore R., WA, H. W. B., AMSA K67295)(new syn.). watkinsi Barker, 2001. Trans. Roy. Soc. S. Aust. 125: 112. (¢ HT, Tinonee Rd, Manning R., NSW, S. Watkins, SAMA I 21528). NSW. westwoodi (Gory & Laporte, 1839)(Coraebus). Mon. Bupr. ii: 15. (6 ST & 2 2 ST, MNHN ). SA, V, T. verna Blackburn, 1891. Trans. R. Soc. S. Aust. 14: 299. (HT damaged sex unknown, Warburton District, Victoria, French collection, NMVA). viridana Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 165. (2 HT, Austral., Standing, BMNH). theryi Kerremans, 1903. in Wytsman Genera Ins. fas. 12: 229. (2 6d ST & 2 22 ST, Victoria, Théry, BMNH). DISCUSSION C. adspersa Kerremans (HT not in BMNH) hab. India belongs in another genus. C. albertisi Gestro is not a synonym of C. acuducta (Carter 1923) which does not occur at the tip of Cape York in the type locality. C. cornuta Gestro (6 ST, MCSNG) = Hypocisseis latipennis Macleay. C. duodecimguttata Guérin-Méneville, 1830 (Buprestis). Type species by subsequent designation (Duponchel, 1843). C. fossicollis Kerremans (HT BMNH) = Alcinous fossicollis (Kerremans). C. fulgidicollis Macleay is very uncommon in collections and has been confused with other species, particularly with C. stellata Barker. C. gestroi Kerremans 1892, Mem. Soc. Ent. Belg. 1: 225 is unknown to me; the holotype is not in the BMNH and the species is not listed under Cisseis in Kerremans (1903). C. maculata Gory & Laporte was misidentified by Carter (1929) who listed C. tasmanica Kerremans and C. pauperula Kerremans as its synonyms; both are good species. C. modesta Kerremans was listed as a synonym of C. rubicunda Kerremans Carter (1929). I find they are separate species. C. nigrita Kerremans was listed as a synonym of C. notulata (Germar) by Carter (1929). Carter (1923) followed Blackburn’s concept of C. notulata, which was incorrect. C. nigrita is a separate species. C. nigripennis Macleay was listed by Carter (1923) as belonging in Neospades. I find it belongs in Cisseis (s.s.). C. obscura Blackburn 1s a species that I have been unable to find in any available collection. C. ovalis Carter could be a synonym of C. opima Thomson. The unique holotype is a discoloured female specimen. . rugiceps Thomson belongs in Neospades. . scabiosa (Boisduval) is unknown to me. . semiobscura Kerremans was listed as a synonym of C. notulata by Carter (1929). It is a separate species, much smaller than C. notulata and common in South Australia. C. spilota Hope, 1846 (Ann. Mag. Nat. Hist. 17: 64) (HT, HMOE not seen) is unknown to me. Hope indicated that it belonged in another AANA 90 S. BARKER genus. Carter (1929) incorrectly listed it as a synonym of C. signaticollis Hope, 1846; spilota has precedence over signaticollis. C. stigmata Gory & Laporte, 1839 selected as type species by Bellamy (1998) (invalid). C. violacea Kerremans was listed as a synonym of C. notulata by Carter (1929). It is a separate, smaller species occurring in the vicinity of Brisbane, Queensland. ACKNOWLEDGMENTS My thanks for assistance to: C. L. Bellamy, Sacramento; M. Moulds, AMSA; T. A. Weir, ANIC; M. Kerley, BMNH; A. Samuelson, BPBM; I. Lansbury, HMOE; M. Uhlig, HUMB; R. Poggi, MCSNG; M. Hanlon, Sydney; J. Menier, MNHN; K. Walker & C. McPhee, NMVA; S. Bily, PMCE; G. B. Monteith, QMBA,; J. Cools, RMBB; E. G. Matthews, SAMA; J. A. Gardner, Waite Agricultural Research Institute, Adelaide. REFERENCES BARKER, S. 1998. Selection of lectotypes and redescriptions of three Cisseis (Coleoptera: Buprestidae) species. Records of the South Australian Museum 31: 21-23. BARKER, S. 1999a. Designation of a lectotype and descriptions of four new species of Australian Buprestidae (Coleoptera). Records of the South Australian Museum 32: 45-49. BARKER, S. 1999b. Designation of lectotypes of three species of Cisseis (Coleoptera: Buprestidae). Transactions of the Royal Society of South Australia 123: 155. BARKER, S. 2001. Descriptions of twenty one new species of Cisseis (sensu stricto) Gory & Laporte 1839 (Coleoptera: Buprestidae: Agrilinae). Transactions of the Royal Society of South Australia 125: 97-113. BARKER, S. & BELLAMY, C. L. 2001. Stanwatkinsius, a new genus of Australian jewel beetles (Coleoptera: Buprestidae: Agrilinae) with a key to known species. Transactions of the Royal Society of South Australia 125: 1-14. BELLAMY, C. L. 1998. Type species designations in the family Buprestidae (Coleoptera). Deutsche Entomologische Zeitschrift 45: 9-15. CARTER, H. J. 1923. Revision of the Genera Ethon, Cisseis and their allies. (Buprestidae.). Proceedings of the Linnean Society of N.S.W. 48: 159-176. CARTER H. J. 1929. A check list of the Australian Buprestidae. Australian Zoologist 5: 265-304. DUPONCHEL, P. A. J. 1843 (1841-1848). In ‘Dictionaire universal d’histoire naturelle’. Vols 1-6. Ed. C. D. D’Orbigny. Bureau Principal de l’Editeur & Renard, Martinet & Cie: Paris. GORY, H. L. & LAPORTE DE CASTELNAU, F. L. 1839. ‘Histoire naturelle et iconographie des insectes Coléoptéres. Monographie des buprestides’. Volume 2, livraisons 25—35. P. Duménil: Paris. KERREMANS C. 1903. Genera Insectorum 12: 1-338. THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS COMMITTEE 10 MARCH 1921 - 30 OCTOBER 2001 WOLFGANG ZEIDLER Summary The Flora and Fauna of South Australia Handbooks Committee has been operating for 80 years producing a series of handbooks on the major elements of the South Australian natural environment. The main role of the Committee was to encourage and persuade specialists to write books on the flora and fauna for no financial reward. The completed manuscripts were then edited by members of the Committee and, in the past, were printed by State Print or its equivalent and published by the State Government Printer at the State Government’s expense. Several years ago this part of the Government’s operations ceased, with no provision made for the further printing of handbooks. THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS COMMITTEE 10 MARCH 1921 —- 30 OCTOBER 2001 WOLFGANG ZEIDLER ZEIDLER, W. 2002. The Flora and Fauna of South Australia Handbooks Committee 10 March 1921 — 30 October 2001. Records of the South Australian Museum 35(1) 91-95. Wolfgang Zeidler, South Australian Museum, North Terrace, Adelaide, South Australia 5000; and Visiting Research Fellow, Adelaide University, Adelaide, South Australia 5005. Manuscript received 13 December 2001. The Flora and Fauna of South Australia Handbooks Committee has been operating for 80 years producing a series of handbooks on the major elements of the South Australian natural environment. The main role of the Committee was to encourage and persuade specialists to write books on the flora and fauna for no financial reward. The completed manuscripts were then edited by members of the Committee and, in the past, were printed by State Print or its equivalent and published by the State Government Printer at the State Government’s expense. Several years ago this part of the Government’s operations ceased, with no provision made for the further printing of handbooks. Since the privatisation of the Government Printer the Committee has sought other means to publish its books and the last two handbooks have been printed with the help of the Board of the Botanic Gardens (‘Fungi’) and the SA Research & Development Institute (Aquatic Sciences) (‘Marine Invertebrates LI’). However, it became very clear to the Committee that the only way it could continue was to establish a publications fund. Initially, the State Government was approached because of its traditional support for the Committee, but this proved fruitless, although the Department of Environment & Heritage and the South Australian Museum did offer some support if funds could also be found from other sources. Various philanthropic trusts were approached and, although some were sympathetic, no funding was forthcoming. Similarly, all three of the State’s universities, whose students and staff are amongst the main users of the handbooks, declined to assist. The formation of a partnership with a publisher was also considered, but rejected as it would potentially compromise the original intention of the Committee because handbooks with commercial potential would be favoured. Left with no alternative, the Handbooks Committee reluctantly concluded that it could not continue to function as originally intended and should be disbanded. The last meeting of the Flora and Fauna of South Australia Handbooks Committee was accordingly held on 30 October 2001. With the demise of the Handbooks Committee, which was once the envy of all other states, it is important to record its history and major achievements. The Committee began in 1921 when Prof E. Wood-Jones, Prof T. B. G. Osborn and Prof J. B. Cleland presented the idea to the then Premier Mr H. Barwell. The concept was for authoritative handbooks to be written by experts in their field without remuneration, to be published by the Government Printer at its convenience, and to be sold at a low price so as to be affordable to the wider community. Here follows an extract recording the foundation of the handbooks series, prefacing Eichler’s (1965) ‘Supplement to J. M. Black’s Flora of South Australia’. After the first World War, the South Australian Branch of the British Science Guild was very active in devising ways for increasing scientific knowledge and its applications. At the time an up-to-date account of the Flora of our State was a pressing need. With this view, Professor J. B. Cleland suggested that the Science Guild might undertake to arrange for the preparation of a series of Handbooks dealing with the Fauna and Flora of South Australia. If these were prepared in an honorary capacity by leading authorities, the Government might well be asked to publish them, as it were, on a pound for pound basis. The Branch accepted the suggestion and it was arranged that a deputation of Professors F. Wood Jones, T. G. B. Osborn and J. B. Cleland should wait on the Hon. The Premier Mr., afterwards Sir Henry Barwell, with the proposal. 92 W. ZEIDLER Fortunately Professor Cleland had prepared a memorandum on the subject in the following terms and this document was handed to the Premier on 25" February, 1921. PROPOSED SCHEME FoR THE PREPARATION OF HANDBOOKS ON THE FAUNA AND FLORA OF SOUTH AUSTRALIA L There is an almost complete lack in South Australia of any handbooks dealing with the Fauna and Flora of the State. Il. In consequence, many individuals with scientific tastes are unable to develop these. This is a great loss to the State, as these persons, with suitable handbooks available, might be led on to contribute voluntarily to the State additions to scientific knowledge of great value to our pastoral, agricultural and other interests. II. | A scheme under the auspices of the S.A. Branch of the British Science Guild is proposed for the preparation of a series of such handbooks to be edited by a sub- committee of experienced workers, the individual parts to be prepared by specialists (in an honorary capacity). IX. The scheme would be spread over a period of 10-15 years. X. Parts would be issued as completed by the authors. None could be available during this financial year. One or two may be so for 1921-1922. XI. — The work involved will be highly skilled and laborious. In the interests of science the authors will be prepared, we believe, to devote their time and knowledge to this work without fee, if a means of publication can be obtained. XII. The value to the community of the works of these authors, thus offered as a gift to the State and world of science, can be put at a very high figure. Would the Government be prepared to accept this offer of gratuitous service and as a return — on the “pound for pound subsidy” principle — arrange for the publication of the Handbooks by the Government Printer? XII. As the work will be spread over many years, this should not necessitate any additions to the staff of the Government Printer, the printing of the parts being proceeded with by him as opportunity offered. The actual out-of-pocket expense to the State would consist, under these circumstances, of the paper only. The parts as completed might be submitted to the Premier or to the Minister of Education, and laid on the table of the House and ordered to be printed and thus be dealt with as Parliamentary Papers. IX. Many of the parts will require to be illustrated. In many cases, blocks are already prepared which doubtless could be used and so reduce expense. X. The educational value of these handbooks will be great. A copy of each might be placed with advantage in every public school in the State, when it could be made available not only to teachers and scholars but to the public of the district as well. To meet this demand and to enable distribution to be made to private individuals (at a nominal price to induce the wide use of these handbooks) about 1,500 copies of each should be printed. XI. Probably from 20 to 30 handbooks would eventually be published (over the period of 10-15 years), the sizes varying from about 20 to 150 pages according to subject matter. Professor CLELAND, Adelaide University. Professor WOOD JONES, Adelaide University. The Premier read through the memorandum and said, “A very generous offer, gentlemen. I shall lay it before Cabinet.” Professor Cleland received from the secretary to the Premier, a letter dated 10° March, 1921 in the following terms accepting the offer. “With reference to the proposed scheme under the auspices of the South Australian Branch of the British Science Guild for the preparation of a series of handbooks on the Fauna and Flora of South Australia which was submitted by yourself and Professor Wood Jones to the Premier on the 25" ultimo, I am directed by the Premier to inform you that in consideration of the contributors in the compilation of the handbooks undertaking the work in an honorary capacity the Government is prepared to undertake the printing of the publication at the Government Printing Office at the expense of the State.” In recent years the Chief Secretary of South Australia decided that the Government Printer should publish the handbooks on a commercial basis, rather than that the Handbooks Committee should apply each year for a government grant for publication. Consequently there is now no delay in printing when the authors offer their completed manuscripts. The British Science Guild has long since ceased to exist but the work of the Handbooks Committee survived until now. The main aim of the THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS COMMITTEE 93 Committee has always been to produce books that document, in a rigorously scientific way, the biological heritage of the State, although in recent years the Committee encouraged authors to broaden their focus to southern Australia in general. Perhaps the most outstanding work published by the Committee is the ‘Flora of South Australia’. The remarkable J. M. Black produced the four-volume work single-handedly and revised the first three volumes, working until he was 96. Revision of the fourth volume was completed after his death. For South Australia’s sesquicentenary a completely revised and rewritten new edition was produced, making this State the only one at the time to have a comprehensive up-to-date record of State flora. There are also other monumental works for which other states have no equivalent (see list of handbooks) and, while some volumes are out of date, many have been updated or reprinted to meet the demand of scientists, naturalists and students. It seems remarkable that Hale’s “The Crustaceans of South Australia’ (1927, 1929), reprinted in 1976, is still used as a basic text, although very much out of date and only providing a limited coverage of the group. It is clear that the handbooks provide invaluable data for research workers, not only in South Australia, or even nationally, but internationally as well. Indeed, the handbooks series has been the envy of scientists in other states. Their contribution to secondary and tertiary education, as significant textbooks, is well recognised. For example, the four-part ‘Flora’, ‘Fishes’ and the three-part ‘Marine Invertebrates of Southern Australia’ series are essential texts for tertiary students. Unfortunately, the later volumes produced by the Government Printer were managed on a commercial basis and were remaindered within two years, so that they were only available for a relatively short period. Future plans for new editions of ‘Fishes’, Amphibians’, ‘The Vegetation of South Australia’, ‘Crustacea’ (as part of the ‘Marine Invertebrates’ series) and ‘Flora’ have been abandoned, although they may be taken up by others. In particular, the State Herbarium is in the process of producing an electronic version of the ‘Flora’, and the South Australian Research & Development Institute (Aquatic Sciences) intends to proceed with future volumes of the ‘Marine Invertebrates’ series as well as revising earlier volumes. As part of the winding up process, the Committee has transferred copyright of the botanical texts to the Board of the Botanic Gardens and State Herbarium, and of the zoological and ecological texts to the Board of the South Australian Museum. Copies of correspondence and minutes of the Handbooks Committee have been deposited with State Archives. TitLes IssueD By THE FLorA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS COMMITTEE 1922 ‘Flora of South Australia’. Part I. Cyathaceae — Orchidaceae. J. M. Black. Second edition 1943. Reprinted 1948. Reprinted (facsimile) 1960, 1972. Third edition, Lycopodiaceae — Orchidaceae (revised by J. P. Jessop) 1978. ‘The Fishes of South Australia’. E. R. Waite. Reprinted 1962. ‘The Mammals of South Australia’. Part I. The Monotremes and the Carnivorous Marsupials. F. Wood-Jones. Reprinted in one volume with parts II (1924) and III (1925), with an Introduction by P. Crowcroft and a Selected Bibliography by J. H. Calaby 1968. ‘Flora of South Australia’. Part II. Casuarinaceae — Euphorbiaceae. J. M. Black. Second edition 1948. Reprinted 1963. Reprinted (facsimile) 1977. ‘The Mammals of South Australia’. Part II. The Bandicoots and the Herbivorous Marsupials. F. Wood-Jones. Reprinted in one volume with parts I (1923) and III (1925), as above, 1968. ‘The Building of Australia and the Succession of Life’ with special reference to South Australia. Part I. W. Howchin. ‘The Mammals of South Australia’. Part Il. The Monodelphia. F. Wood-Jones. Reprinted in one volume with parts I (1923) and II (1924), as above, 1968. ‘Flora of South Australia’. Part III. Meliaceae — Scrophulariaceae. J. M. Black. Second edition, Callitrichaceae — Plumbaginaceae 1952. Reprinted 1964. Reprinted (facsimile) 1977. ‘The Crustaceans of South Australia’. Part J. H. M. Hale. Reprinted in one volume with part II (1929) 1976. 1923 1923 1924 1924 1925 1925 1926 1927 94 1928 1929 1929 1929 1930 1934 1935 1936 1937 1938 1939 1940 1947 W. ZEIDLER ‘The Building of Australia and the Succession of Life’ with special reference to South Australia. Part II. Mesozoic and Cainozoic. W. Howchin. ‘Flora of South Australia’. Part IV. Bignoniaceae — Compositae. J. M. Black. Second edition, Oleaceae — Compositae (revised by E. L. Robertson) 1957. Reprinted (facsimile) 1980. ‘The Crustaceans of South Australia’. Part Il. H. M. Hale. Reprinted in one volume with part I (1927) 1976. ‘The Reptiles and Amphibians of South Australia’. E. R. Waite. ‘The Building of Australia and the Succession of Life’ with special reference to South Australia. Part III. Pleistocene. W. Howchin. “Toadstools and Mushrooms and other Larger Fungi of South Australia.’ General introduction; Toadstools and Mushrooms. J. B. Cleland. Reprinted in one volume with part II (1935), including ‘Notes on some Edible and Poisonous Fungi’ by P. H. B. Talbot, 1976. ‘Toadstools and Mushrooms and other Larger Fungi of South Australia’. Polypores, Coral Fungi and remaining Hymenomycetes; Puff-balls, Jelly-like fungi, the larger Ascomycetes and Myxomycetes. J. B. Cleland. Reprinted in one volume with part I (1934), as above. ‘The Seaweeds of South Australia’. Part I. Introduction and The Green and Brown Seaweeds. A. H. S. Lucas. ‘The Vegetation of South Australia’. J. G. Wood. ‘The Molluscs of South Australia’. Part I. The Pelecypoda. B. C. Cotton & F. K. Godfrey. ‘Primitive Insects of South Australia’. Silverfish, Springtails and their allies. H. Womersley. ‘The Molluscs of South Australia’. Part II. Scaphopoda, Cephalopoda, Aplacophora and Crepipoda. B. C. Cotton & F. K. Godfrey. ‘The Seaweeds of South Australia’. Part 1959 1961 1962 1964 1965 1966 1972 1974 1975 1976 1978 1978 1979 1980 1980 1982 1984 1984 1986 II. The Red Seaweeds. A. H. S. Lucas & F. Perrin. ‘South Australian Mollusca’. Archaeogastropoda. B. C. Cotton. ‘South Australian Mollusca’. Pelecypoda. B. C. Cotton. ‘The Marine and Freshwater Fishes of South Australia’. T. D. Scott. ‘South Australian Mollusca’. Chitons. B. C. Cotton. ‘Supplement to J. M. Black’s Flora of South Australia’ (second edition 1943-— 1957). H. J. Eichler. ‘Aboriginal Man in South and Central Australia’. Part I. Edited by B. C. Cotton. Only one part was published. ‘The Vegetation of South Australia’. R. L. Specht. Second edition. ‘The Marine and Freshwater Fishes of South Australia’. T. D. Scott, C. J. M. Glover & R. V. Southcott. Second edition. Reprinted (facsimile, with 16 pp of colour plates) 1980. ‘Plant Feeding and other Bugs (Hemiptera) of South Australia’. Heteroptera Part I. G. F. Gross. ‘Plant Feeding and other Bugs (Hemiptera) of South Australia’. Heteroptera Part II. G. F. Gross. ‘Butterflies of South Australia’. R. H. Fisher. ‘Amphibians of South Australia’. M. J. Tyler. ‘Lichens of South Australia’. R. B. Filson & R. W. Rogers. ‘Mosses of South Australia’. D. G. Catcheside. ‘Acacias of South Australia’. D. J. E. Whibley. ‘Marine Invertebrates of Southern Australia’. Part I. Edited by S. A. Shepherd & I. M. Thomas. ‘Psylloidea of South Australia’. F. D. Morgan. ‘The Marine Benthic Flora of Southern Australia’. Part I. H. B. S. Womersley. ‘The Dynamic Partnership: Birds and Plants in Southern Australia’. Edited by H. A. Ford & D. Paton. 1986 1986 1987 1989 THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS COMMITTEE 95 ‘Flora of South Australia. Volumes I-IV’. Edited by J. P. Jessop & H. R. Toelken. Fourth edition (third edition consisted of part I only). ‘The Ecology of Forests and Woodlands of South Australia’. Edited by H. R. Wallace. ‘The Marine Benthic Flora of Southern Australia’. Part II. H. B. S. Womersley. ‘Marine Invertebrates of Southern Australia’. Part II. Edited by S. A. Shepherd & I. M. Thomas. 1990 1992 1994 1997 1997 ‘Orchids of South Australia’. R. Bates & J. Z. Weber. ‘Acacias of South Australia’. D. J. E. Whibley & D. E. Symon. Second edition. ‘The Fishes of Australia’s South Coast’. Edited by M. F. Gomon, J. C. M. Glover & R. Kuiter. ‘Larger Fungi of South Australia’. C. A. Grgurinovic. ‘Marine Invertebrates of Southern Australia’. Part III. Edited by S. A. Shepherd & M. Davies. Dr Wolfgang Zeidler (Hon. Secretary), on behalf of the Flora and Fauna of South Australia Handbooks Committee — Em Prof William D. Williams (Chairman), Dr Sue Barker (Botanical Editor), Dr Margaret Davies (Zoological Editor), Dr Shelley Barker, Ms Robyn Barker, Dr John G. Conran, Mr Peter Copley and Dr Brian D. Morley. OBITUARY GRAEME LLOYD PRETTY 1940 -— 2000 HELEN TOLCHER Summary Graeme Lloyd Pretty was born in Melbourne on 25 June 1940. His father was in the RAAF and the family soon moved to Richmond NSW, where Graeme spent his childhood and went to school. After graduating from Sydney University BA (Hons) in Classical Archaeology and Dip Ed, he contemplated undertaking a PhD with Jack Golson at the Australian National University. However, he began working at the South Australian Museum in January 1962 as Assistant Curator of Anthropology under Norman Tindale, from whom he learned much and with whom he kept in touch long after the latter’s retirement. His first interest was in Melanesian prehistory — he might well have made this his major field of work. OBITUARY GRAEME LLOYD PRETTY 1940-2000 ~ PCE ~ WES oe FD Graeme Pretty and Henry Pollach (left) at Roonka, 8 August 1971. Roonka photos R83: 2. South Australian Museum. Graeme Lloyd Pretty was born in Melbourne on 25 June 1940. His father was in the RAAF and the family soon moved to Richmond NSW, where Graeme spent his childhood and went to school. After graduating from Sydney University with a BA (Hons) in Classical Archaeology and Dip Ed, he contemplated undertaking a PhD with Jack Golson at the Australian National University. However, he began working at the South Australian Museum in January 1962 as Assistant Curator of Anthropology under Norman Tindale, from whom he learned much and with whom he kept in touch long after the latter’s retirement. His first interest was in Melanesian prehistory—he might well have made this his major field of work. In 1964 he was appointed Assistant Curator of Archaeology, and archaeological fieldwork was added to his duties. He excavated or assisted at a number of sites on the lower River Murray, seeking the advice of John Mulvaney who had recently excavated Fromm’s landing. At the same time he continued research into Melanesian culture, including six weeks fieldwork in the Southern Highlands of Papua New Guinea (PNG) in 1968-69. Subsequently, he was invited to review the functions of the PNG Museum and Art Gallery. Later, he also advised the Australian National Gallery on its Melanesian collections. In 1968 Pretty began a project that was to become the most important one of his working life. A salvage operation to remove a surface campsite and scattered human remains from a sandhill beside the River Murray at Roonka, near Blanchetown, developed into a major excavation that would attract worldwide interest. At that time he was unique in having consulted with the relevant Aboriginal group and obtained approval before beginning work—he continued this contact throughout and after the conclusion of the dig. This rapport with the Aboriginal people undoubtedly contributed to making possible the excavation on the scale and significance that was achieved. 98 H. TOLCHER, J. PRESCOTT & J. MULVANEY Despite the disadvantages and difficulties of working on a surface fully exposed to gale-force winds and summer heat rising to the 50°C range, he retrieved a wealth of archaeological material and information from a site which proved to have been occupied for some 18 000 years. In all, 216 complete or fragmentary burials were uncovered. The grave goods were unprecedented at that time. In addition to almost every variant of human mortuary practice, it also gave evidence for hitherto undocumented customs, of which there were well over 150 examples. Roonka looms large for its deep significance. Here is a statistical sample from one place, across some thousands of years, permitting health, injury and dietary studies of the population. Pretty correctly saw that aspects of the rituals involved— the mode of burial, including grave goods, and the antiquity of the rituals—presented a major window to past belief and behavioural systems. No other burial ground has produced so much evidence. The almost exclusively amateur workforce of volunteers became, over the eight years of active fieldwork, a team of patient, dedicated, highly skilled excavators. Under Pretty’s direction, the site was strictly controlled, meticulously recorded and kept secure from outside interference. The standard and scope of the work attracted substantial financial support as well as visitors from abroad, including osteo-archaeologists and a delegation from the People’s Republic of China. One group will remember the visit not only for the archaeology but because their bus became bogged to the axles on the Roonka Flat. Pretty was appointed to the position of Senior Curator of Anthropology and Archaeology in 1975. The section of Pretty’s work that has been best documented is probably that dealing with chronology. He had a critical appreciation of the role of physical dating methods in archaeology and sought to create a temporal framework for the finds at Roonka. The chronology of the site was determined by a larger number of different dating methods than any other archaeological site in Australia, including thermoluminescence, palaeomagnetism, uranium fluorimetry and C-14 dating, both conventional and AMS. It is not generally known that he was responsible for planting the idea for developing luminescence dating in the Physics Department at the University of Adelaide in conversation with Professor John Prescott. This was in 1972, at the very infancy of the thermoluminescence technique. The earliest Adelaide work on thermoluminescence dating per se was based on samples from Roonka. Pretty extended his fieldwork research to Island Melanesia in 1971 and 1973, finding additional support from sources outside Australia. However, the demands of attempting to elucidate the intricacies of the Roonka information, together with constraints on finance and staffing, resulted in the Melanesian work being put aside—apart from ongoing work, under his direction, on the sorting of the Foreign Ethnology reserve collections and their eventual rehousing at Kent Town in 1986. As Senior Curator of Archaeology, Pretty’s work was directed almost exclusively to organising the mass of information from the Roonka site in order to arrive at what he considered an adequate assessment on which to base his final report. During the 1980s a master index of the Roonka data was prepared. The Third Australasian Archaeometry Conference was held in Adelaide in 1988. Pretty devised the theme of the Symposium, ‘Early Man in the Southern Hemisphere’, which was designed to give a specifically archaeological balance to the program. He arranged invitations and finance for the international contributors: G. Delibrias, G. J. Bartstraa and H. J. Deacon; and organised the conference excursion and wrote the field notes. In his own contribution to the symposium, a paper analysing the cultural chronology of Roonka, it is possible to see a foretaste of his overall interpretation of Roonka in its wider context. An early indication is also found in his exciting and innovative 1977 paper, “The cultural chronology of Roonka Flat’. In it he remarked ‘This present paper therefore celebrates the author’s satisfaction that the archaeology is now sufficiently clear to commit preliminary notice of it to paper’. Yet, during the following 23 years, he produced no paper with the promised detail. It is difficult for those of us who knew him to avoid reflection on his apparent reluctance to put his ideas on paper where they could be exposed to the critical appraisal of his peers. He did publish sections of the work with his collaborators, but the overall synthesis was yet to come. Perhaps he was sensitive to possible criticism; perhaps the very volume of his material was an obstacle. It seems likely that he laboured over the precise language with which to present a polished final monograph. In 1982 restructuring of the South Australian Museum’s divisions and staffing made Pretty one of a number of Curators of Anthropology. He OBITUARY —- GRAEME LLOYD PRETTY 99 accepted a severance package in 1994, continuing his work on the Roonka report until 1996, when he was stricken with viral encephalitis, from which he never fully recovered. He died on 6 November 2000. In his work Pretty was patient, a seeker of precision in language that was baffling to many, and single-minded to a degree that ignored workplace relationships and, at times, made it difficult for people to work with him. His memory was exceptional. The breadth of his reading was remarkable and was reflected in his large personal library. He left the Museum’s Foreign Ethnology collection well organised and totally accessible, and the Roonka material fully recorded and stored. He gained the respect of his field workers, instructing them in aspects of Aboriginal culture and inspiring in them a dedication and loyalty to the project that will ensure the publication of the final report. Graeme married Dana in 1973 and they had one daughter. To those who knew him well he was a kind and considerate friend. At home he was a gracious and amusing host, an erudite and entertaining conversationalist, a devoted husband and a proud and loving father. AUTHORED AND COLLABORATIVE WorK To 1995, LisTED By FIELD Archaeology PRETTY, G. L. 1964. Stone objects excavated in New Guinea. Man 64, 138: 17. PRETTY, G. L. 1964. A stone figure of a crested cockatoo from Melanesia. Man 64, 220: 183-184. PRETTY, G. L. 1965. Two stone pestles from Western Papua and their relationship to prehistoric pestles and mortars from New Guinea. Records of the South Australian Museum 15, 1: 120-130. PRETTY, G. L. (with M. J. TYLER). 1966. A bizarre refuge for some Australian leptodactylid frogs. Records of the South Australian Museum 15, 2: 360- 361. PRETTY, G. L. 1967. Excavations at the Anglo-Saxon site of Wallingford, Berkshire, England, 1966. Journal of the Anthropological Society of South Australia 5, 2: 3-5. PRETTY, G. L. 1967. Rescue excavation of an aboriginal grave, Tailem Bend. South Australian Naturalist 41, 4: 9-11. PRETTY, G. L. (with S. A. GALLUS). 1967. The anthropology and archaeology of the Nullarbor Plain, South Australia. Pp. 47-49 in ‘Caves of the Nullarbor’. Eds. J. R. DUNKLEY & T. M. L. WIGLEY. Adelaide. PRETTY, G. L. 1968. Excavation of Aboriginal graves at Gidgealpa, South Australia. Records of the South Australian Museum 15, 4: 671-677. PRETTY, G. L. 1968. Two more prehistoric stone artifacts from Western Papua. Records of the South Australian Museum 15, 4: 693-697. PRETTY, G. L. 1969. Prehistoric background to Australia. Tradition: Journal of the History Teachers’ Association (S.A.) May. Pp. 13-23. PRETTY, G. L. 1969. Excavation of an Aboriginal cemetery near Blanchetown, River Murray, South Australia. Paper read to the 41st Congress of the Australian and New Zealand Association for the Advancement of Science, Adelaide, August 1969. 15 pp., 10 illus. PRETTY, G. L. 1970. Excavation of an Aboriginal cemetery on Roonka Station, River Murray. Kalori, Journal of the Museums Association of Australia 38: 17. PRETTY, G. L. 1971. Excavations at Roonka Station, Lower River Murray, South Australia, 1968-1970. Journal of the Anthropological Society of South Australia 19, 9: 6-15. PRETTY, G. L. 1971. Further investigations into Melanesian culture-history: South Australian Museum field research in Island Melanesia, 1971. Kalori: Journal of the Museums Association of Australia 42: 89-95. PRETTY, G. L. 1972. ‘Report of an Inspection of Archaeological Sites and Field Monuments in the Territory of Papua New Guinea’. Papua New Guinea Public Museum and Art Gallery: Port Moresby. vii + SO pp., illus. PRETTY, G. L. 1975. Archaeology in South Australia: a report on recent work. Australian Archaeology: Australian Archaeological Association Newsletter 3: 32-39. PRETTY, G. L. 1975. ‘Social Change in Prehistoric Australia; the Evidence from Roonka’. Paper read to the 46th Congress of the Australian and New Zealand Association for the Advancement of Science, Canberra. 12 pp., 16 illus. PRETTY, G. L. 1976. The excavations at Roonka Flat, South Australia: an insight into ancient Australian society. Actes et Proc. IX Congres, Union Internationale des Sciences Prehistoriques et Protohistoriques, Nice. Section 6, 2: 100-112. PRETTY, G. L. 1977. Archaeology in South Australia. Pp. 40-54 in ‘South Australian Yearbook’. Australian Bureau of Statistics: Adelaide. PRETTY, G. L. 1977. The cultural chronology of the Roonka Flat: a preliminary consideration. Pp. 288- 331 in ‘Stone Tools as Cultural Markers: Change, Evolution, Complexity’. Ed. R. V. S. WRIGHT. Australian Institute of Aboriginal Studies: Canberra. PRETTY, G. L. 1978. Review of V. D. Watson & J. D. 100 Cole, ‘Prehistory of the Eastern Highlands of New Guinea’. The Artefact 5: 98-102. PRETTY, G. L. 1980. Prehistory. Pp. 9-17 in ‘Proceedings of the Fiftieth Anniversary Seminar of the Anthropological Society of South Australia’. Adelaide. PRETTY, G. L. 1981. Trial excavation of an Aboriginal camp site and site survey, Kinchega National Park. Pp. 53-63 in ‘Darling Surveys, I’. Ed. J. M. HOPE. Occasional Papers in Prehistory 3. Australian National University: Canberra. PRETTY, G. L. 1982. ‘The Archaeology of Roonka: Examining the Human and Environmental History of an Australian Place’. Paper read to the Nature Conservation Society of South Australia, Adelaide. 23 pp., 20 illus. PRETTY, G. L. 1982. Prehistory of the Australians: the view from Roonka. University of Adelaide, Department of Classics, Lecture Series-Archaeology TH: 1-11. PRETTY, G. L. (with R. C. PATON & R. D. J. WEATHERSBEE). 1983. Tribal man. Pp. 115-126 in ‘Natural history of the South East of South Australia’. Eds. M. J. TYLER, C. R. TWIDALE, J. K. LING & J. W. HOLMES. Royal Society of South Australia: Adelaide. PRETTY, G. L (with T. J. GARA). 1983. Scientific background. in ‘Roonka Conservation Park Management Plan’. Ed. P. J. HUGHES. ANUTECH Services: Canberra. PRETTY, G. L. 1984. Review of John Aubrey (ed.), ‘Monumenta Britannica or a Miscellany of British Antiquities (parts 1 & 2)’. Mankind 14, 3: 254-6. PRETTY, G. L. 1986. The prehistory of South Australia. Pp. 3-62 in ‘The Flinders History of South Australia: Social History. 3’. Ed. E. RICHARDS. Wakefield Press: Adelaide. PRETTY, G. L. (with R. S. MERRILLEES & C. A. HOPE). 1990. ‘Living with Egypt’s Past in Australia’. Museum of Victoria: Melbourne. viii + 78 pp., illus. PRETTY, G. L. 1990. The significance of the tombs of evolved prehistoric hunters and gatherers; Southern Australia and Southern France compared. (Invited address, Royal Society of South Australia. May, 1990.) Newsletter, Royal Society of South Australia, June. Pp. 4-7. The following are in manuscript form and it is intended that they will be published in due course. PRETTY, G. L. ‘Archaeological Investigations at Roonka, Part I - Description of the evidence gathered from the Karmakalingbila Ngaiawang territory: Geological and geochronological background; cultural landscape and stratification; ecological, demographic and cultural finds’. Text, tables, illus. H. TOLCHER, J. PRESCOTT & J. MULVANEY PRETTY, G. L. (with G. K. WARD) (eds). ‘Archaeological Investigations at Roonka, Part 2 - Ethnoanalogical and comparative analysis: Conjectures about territory, society and ideas in the Karmakalingbila Ngaiawang territory in the light of its Murray Darling Basin context’. (Accepted for publication in British Archaeological Reports: International Series.) PRETTY, G. L. ‘Archaeological Investigations at Roonka, Part 3 - Ethnodeductive and systemic analysis: Conjectures about the mechanics and dynamics of cultural change in_ the Karmakalingbila Ngaiawang territory in the light of its Central Southern Australian context’. Text, tables, illus. PRETTY, G. L. ‘Archaeological Investigations at Roonka, Part 4 - Decoding the symbol inventory of the Karmakalingbila Ngaiawang sequence: Restoring immediate intelligibility and ultimate causal identity to culturally unfamiliar modes of action through appeals to phyletic behavioural repertoires’. Text, tables, illus. PRETTY, G. L. ‘Archaeological Investigations at Roonka, Part 5 - The explanation of prehistoric hunter-gatherer culture change in the Karmakalingbila Ngaiawang territory during the Holocene; tests of confirmation and concluding synthesis’. Text, tables, illus. Art PRETTY, G. L. 1963. A turtle shell mask of Torres Strait type in the Macleay Museum, University of Sydney. Records of the South Australian Museum 14, 3: 421-425. PRETTY, G. L. 1971. The National Collection of Primitive Art, Canberra. Art and Australia 8, 4: 324- 329. PRETTY, G. L. 1971. Comment on R. McGhee, ‘Differential artistic productivity in the Eskimo cultural tradition’. Current Anthropology 17, 2: 216. PRETTY, G. L. 1976. ‘Arts of Melanesia: Adelaide Festival of Arts 1976’. (Exhibition Catalogue.) David Jones Gallery: Adelaide. Cultural heritage and public policy PRETTY, G. L. 1970. ‘Report on the Commonwealth Collection of Primitive Art from New Guinea and the Pacific Islands’. Commonwealth Art Advisory Board: Canberra. iv + 78 pp., illus. PRETTY, G. L. 1971. Museums and controls on the export of cultural property. Kalori: Journal of the Museums Association of Australia 42: 89-95. PRETTY, G. L. (General Editor). 1977. ‘National Folk Province Feasibility Study Papers’. Commonwealth Department for the Environment, Housing and OBITUARY — GRAEME LLOYD PRETTY 101 Community Development and South Australian Department for the Environment: Adelaide, as follows: No. 1 PRETTY, G. L. ‘The Ngaiawang Folk Province: A Proposal’. No. 2 PRETTY, G. L. ‘The National Folk Province Concept - Specification for a feasibility study’. No. 3 ROWNEY, B. G. ‘Kapunda - Historical development of its town plan’. No. 4 PRETTY, G. L. ‘The National Folk Province Concept and the National Estate’. No. 5 KALIBATAS, E. ‘Craigie’s Plains: Construction and materials analysis of a historic vernacular sheep station’. No. 8 SPIERS, G. K. ‘Survey of Heritage Agencies in South Australia’, No 9 ROWNEY, B. G. ‘Surviving cottages in Mine Square, Kapunda’. No. 10 PRETTY, G. L. (with B. G. ROWNEY & P. F. DONOVAN). ‘Ngaiawang Folk Province: Preliminary inventory of the cultural landscape’. No. 11 SPIERS, G. K. ‘The administration of heritage properties through Local Government: I, The South Australian Planning and Development Act 1996- 1975’. No. 12 DONOVAN, P. F. & SCHEPPERS, K. H. ‘Report on the Kapunda Colonial Festival, 1976’. No. 13 SPIERS, G. K ‘The administration of heritage properties through Local Government: II, The Municipal Heritage Trust’. No. 15 DONOVAN, P. F. ‘Woakwine Folk Province: Outline sketch of cultural landscape’. No. 16 DONOVAN, P. F. & SCHEPPERS, K. H. ‘The administration of heritage property through Local Government: III, Report from the Heritage Officer, District Council of Kapunda’. No. 19 ROWNEY, B. G. & DONOVAN, P. F. ‘Old Legislative Council Building, Adelaide : An architectural assessment’. No. 20 ROWNEY, B. G. & DONOVAN, P. F. ‘Establishing conservation priorities for the built environment: The example of Kapunda’. Ethnography PRETTY, G. L. 1969. The Macleay Museum mummy from Torres Straits; a postscript to Elliot Smith and the diffusion controversy. Man: the Journal of the Royal Anthropological Institute NS 4, 1: 24-43. PRETTY, G. L. 1969. ‘Salvage Ethnography in New Guinea: the South Australian Museum Expedition to the Southern Highlands District, Papua, 1968-1969’. Report to the Wenner-Gren Foundation for Anthropological Research, New York. iv + 85 pp., illus. PRETTY, G. L. 1970. Aboriginal Studies. (Review of ‘Man, Land and Myth in North Australia: the Gunwinggu people’ by R. M. & C. H. Berndt ; ‘Yiwara-Foragers of the Australian Desert’ by Richard A Gould.) Australian Book Reviews 9, 12: 332-334. PRETTY, G. L. 1974. Comment on Francis L. Utley, ‘The migration of folk tales: four channels to the Americas’. Current Anthropology 15, 1: 19-20. PRETTY, G. L. 1980. The Aboriginal cultural landscape of the Lower Murray Valley. Pp. 43-52 in ‘Preserving Indigenous Cultures; a New Role for Museums’. Eds. R. Edwards & J. Stewart. UNESCO and Australia Council: Canberra. PRETTY, G. L. (with A. CALDER). 1980. Mummification in Australian and Melanesia. Pp. 194-210 in ‘Mummies, Disease and Ancient Cultures’. Eds. A. & E. Cockburn. Cambridge University Press: New York. PRETTY, G. L. and course participants, Visual Arts Discipline, Flinders University for the Gerard Aboriginal Community. 1983. ‘Artifacts of the Ngaiawang Tribe, Lower Murray Valley, South Australia, obtained by E. J. Eyre (1841-1844)’. (Document basis for the renewal of the Gerard Community artifact industry.) PRETTY, G. L. 1993. The Aborigines of the South East of South Australia and the Western District of Victoria before white contact. Community History (Adelaide) 3, 4: 2-5. Evolutionary biology PRETTY, G. L. (with P. SMITH & M. PROKOPEC). 1988. Dentition of a prehistoric population from Roonka Flat, South Australia. Archaeology in Oceania 23: 31-36. PRETTY, G. L. (with M. PROKOPEC & P. SMITH). 1994. Prehistoric South Australian Aboriginals: New evolutionary perspectives. Variability and Evolution (Adam Mickiewicz University, Poznan) 4: 21-56. Forensic anthropology PRETTY, G. L. 1975. The recovery of human remains for forensic purposes. Proceedings Australian Forensic Science Society 1, 3: 68-74. PRETTY, G. L. (with D. J. POUNDER & M. PROKOPEC). 1983. A probable case of euthanasia amongst prehistoric Aborigines at Roonka, South Australia. Forensic Science International 23: 99- 108. PRETTY, G. L. (with M. PROKOPEC, D. SIMPSON & L. MORRIS). 1984. Craniosynostosis in a prehistoric Aboriginal skull: A case report. Ossa. International Journal of Skeletal Research 9-11: 111-118. 102 Heritage preservation PRETTY, G. L. 1967. ‘Preliminary catalogue of archaeological monuments in the Territory of Papua New Guinea’. Papua New Guinea Public Museum and Art Gallery: Port Moresby. iv + 95 pp., map. PRETTY, G. L. 1968. Surveying archaeological monuments in Papua New Guinea. Journal of the Anthropological Society of South Australia 6, 2: 5-8. PRETTY, G. L. 1970. Survey of prehistoric monuments in South Australia. Pp. 39-50 in ‘Aboriginal antiquities in Australia: their nature and preservation’. Ed. F. D. McCarthy. Australian Institute of Aboriginal Studies 22. PRETTY, G. L. 1976. The Ngaiawang Folk Province. Journal of the Anthropological Society of South Australia 14, 9: 13-22. PRETTY, G. L. 1978. The Ngaiawang Folk Province. Pp. 28-45 in ‘The Tide of Australian Settlement — Conservation of the Physical Evidence’. Australia ICOMOS Proceedings, Beechworth, 14-16 April 1978. Australia/ICOMOS: Hawthorn, Vic. History PRETTY, G. L. 1967. Edward Gibbon Wakefield. Pp. 559-562 in ‘Australian Dictionary of Biography’. Ed. D. Pike. Melbourne. PRETTY, G. L. 1975. The historical collections in the South Australian Museum. Pp. 173-176 in ‘Proceedings of the National Seminar on the Conservation of Cultural Material, Perth, August 1973’. Eds C. Pearson & G. L. Pretty). Institute for the Conservation of Cultural Material: Perth. PRETTY, G. L. 1986. Australian history at Roonka. Journal of the Historical Society of South Australia, 14: 107-122. PRETTY, G. L. 1986. Foreword to ‘People, Places and Buildings’. J. Faull & G. Young. South Australian Centre for Settlement Studies: Adelaide. Materials conservation PRETTY, G. L. (with C. PEARSON). (Eds). 1976. ‘Proceedings of the National Seminar on the Conservation of Cultural Material, Perth, August 1973’. Institute for the Conservation of Cultural Material: Perth. 297 pp., illus. Museology PRETTY, G. L. 1967. The British Museum. Friends of the South Australian Museum Newsletter 4: 9-11. PRETTY, G. L. 1968. ‘Report on the State of the Papua New Guinea Museum: the Collections of Archaeology and Ethnology’. Trustees, Papua New H. TOLCHER, J. PRESCOTT & J. MULVANEY Guinea Public Museum and Art Gallery: Port Moresby. iv + 95 pp., illus. PRETTY, G. L. 1971. Oceanic museum resources in Australian museums. Pp. 179-186 in ‘Meeting on Studies of Oceanic Cultures’. Australian National Advisory Committee for UNESCO: Canberra. PRETTY, G. L. 1976. The Ethnographic Collections in the South Australian Museum. Pp. 107-119 in ‘Proceedings, National Seminar on Conservation of Cultural Material’. Eds. C. Pearson & G. L. Pretty. Institute for the Conservation of Cultural Material: Perth. Palaeoecology PRETTY, G. L. (with W. E. BOYD). 1989. Some prospects for archaeological palaeobotany in Australia; an example from South Australia. Australian Archaeology 28: 40-52. PRETTY, G. L. (with W. E. BOYD). (in press). From the sensible to the spiritual — Cultural signals from palaeobotanic evidence at Roonka, South Australia. Archaeological Dialogues. Palaeopathology PRETTY, G. L. (with M. E. KRICUN). 1989. Health status of the prehistoric Roonka population. World Archaeology 21, 2: 198-224. PRETTY, G. L. (with M. PROKOPEC). 1990. Skeletal aging. Rate of tooth attrition in hunter- gatherer populations, prehistoric Roonka, South Australia. Colloquiae Anthropologicae 14, 2: 331- 334. PRETTY, G. L. (with M. PROKOPEC). 1991. Observations on health, genetics and culture from analysis of prehistoric population data from Roonka, South Australia. Pp. 151-158 in ‘Human Palaeopathology: Current Syntheses and Future Options’. Eds. D. J. Ortner & A. C. Aufderheide. Smithsonian Press: Washington, DC. PRETTY, G. L. (with T. BROWN & M. E. KRICUN). 1992. Extensive compensatory remodeling of craniofacial structures; a case from prehistoric South Australia. Pp. 139-143 in ‘Craniofacial Variation in Pacific Populations. Papers presented at the XVII Pacific Science Congress Symposium, Honolulu, Hawaii, May 30, 1991’. University of Adelaide: Adelaide. PROKOPEC, M. and PRETTY, G. L. 1994. Praehistorische Bewohner der Roonka-Siedlung (Sudaustralien). Ethnographisch-Archaeologische Zeitschrift 35, 1: 136-148. SIMPSON, D., PROKOPEC, M., MORRIS, L., & PRETTY, G. L. 1984. Prehistoric craniosynostosis. A case report. Records of the Adelaide Children’s Hospital 3, 2: 163-168. OBITUARY —- GRAEME LLOYD PRETTY 103 Radiometric dating PRETTY, G. L. (with J. R. PRESCOTT, H. POLACH & B. W. SMITH). 1983. Some comparisons of C-14 and thermoluminescent dates from Australia. (Proceedings of the 1981 Groningen C-14 Conference). Council of Europe PACT Journal 8: 205-211. PRETTY, G. L. 1988. Radiometric chronology and significance of the fossil hominid sequence from Roonka, South Australia. Pp. 32-52 in ‘Early Man in the Southern Hemisphere’. Ed. J. R. Prescott. Supplement to Archaeometry: Australasian Studies. University of Adelaide Department of Physics and Mathematical Physics: Adelaide. Technology PRETTY, G. L. 1970. Review of ‘Excavations of the prehistoric iron industry in West Borneo’ by T. Harrisson & S. J. O'Connor. Man: The Journal of the Royal Anthropological Institute, London. NS. 5: 710-711. Trace element chemistry PRETTY, G. L. 1991. (with F. D. PATE, J. T. HUTTON & R. A. GOULD). Alterations of in vivo elementary dietary signatures in archaeological bone. Evidence from the Roonka Flat Dune, South Australia. Archaeology in Oceania 26: 58-69. Helen Tolcher, c/- South Australian Museum, North Terrace, Adelaide, South Australia 5000; Professor Emeritus John Prescott, Physics Department, Adelaide University, South Australia 5005; and Professor Emeritus John Mulvaney, Centre for Archaeological Research, Australian National University, ACT 0200. Records of the South Australian Museum 35(1): 97-103. IRNIECOIRIDS OJF plpteiils SOUTH AUSTRALIAN MUSEUM VOLUME 35 PART 1. MAY 2002 ISSN 0376-2750 CONTENTS: 33 49 Te 85 91 oF. ARTICLES I. BEVERIDGE New species and new records of Cloacina von Linstow, 1898 (Nematoda: Strongyloidea) parasitic in macropodid marsupials from Papua New Guinea. B. P. KEAR Darwin Formation (Early Cretaceous, Northern Territory) marine reptile remains in the South Australian Museum. H. W. PALM & I. BEVERIDGE Tentaculariid cestodes of the order Trypanorhyncha (Platyhelminthes) from the Australian region. E.G. MATTHEWS & T. A. WEIR Two new species of the genus Lepanus Balthasar from South Australia (Coleoptera: Scarabaeidae). S. BARKER A checklist of Cisseis (sensu stricto) Gory & Laporte, 1839 (Coleoptera: Buprestidae: Agrilinae). W. ZEIDLER The Flora and Fauna of South Australia Handbooks Committee 10 March 1921 — 30 October 2001. NOTE H. TOLCHER, J. PRESCOTT & J. MULVANEY Obituary — Graeme Lloyd Pretty Published by the South Australian Museum, North Terrace, Adelaide, South Australia 5000. INE CORDS Ole Ie0e, SOUTH ~ AUSTRALIAN MUSEUM VOLUME 35 PART 2 OCTOBER 2002 THE LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE (COLEOPTRA : INSECTA) C. H. S. WATTS Summary The larvae of the aquatic Hydrophilid genera Allocotocerus Kraatz, Hybrogralius Orchymont and Regimbartia Zaitzev are described and figured for the first time.The larvae of the following Australian species are also described and figured, most for the first time: Allocotocerus punctatus (Blackburn); Amphiops aueenslandicus Balfour-Browne; Berosus australiae Mulsant; Enochrus eyrensis (Blackburn), E. mastersi (W. MacLeay); Helochares tristis (W. MacLeay), H. clypeatus (Blackburn), H. tenuistriatus Regimbart, H. foveicollis (Montrouzier); Hydrobiomorpha sp. ; Hydrophilus bilineatus (MacLeay); Hybrogralius hartmeyeri (Regimbart); Laccobius decipiens Gentili; Limnoxenus zealandicus (Broun); Regimbartia attenuata (Fabricius); and Sternolophus margincollis (Hope). THE LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE (COLEOPTERA: INSECTA) CHS WATTS WATTS, CHS. 2002. The larvae of some Australian aquatic Hydrophilidae (Coleoptera: Insecta). Records of the South Australian Museum 35(2): 105-138. The larvae of the aquatic Hydrophilid genera Allocotocerus Kraatz, Hybogralius Orchymont and Regimbartia Zaitzev are described and figured for the first time. The larvae of the following Australian species are also described and figured, most for the first time: Allocotocerus punctatus (Blackburn); Amphiops queenslandicus Balfour-Browne; Berosus australiae Mulsant; Enochrus eyrensis (Blackburn), E. mastersi (W. MacLeay); Helochares tristis (W. MacLeay), H. luridus (W. MacLeay), H. clypeatus (Blackburn), H. tenuistriatus Regimbart, H. foveicollis (Montrouzier); Hydrobiomorpha sp.; Hydrophilus bilineatus (MacLeay); Hybogralius hartmeyeri (Regimbart); Laccobius decipiens Gentili; Limnoxenus zealandicus (Broun); Regimbartia attenuata (Fabricius); and Sternolophus marginicollis (Hope). The larvae were identified by rearing larvae collected in the field, or occasionally by association and elimination, or by using biochemical methods. The pupae of Amphiops queenslandicus and Hybogralius hartmeyeri are described and figured. Unlike most other Hydrophilids the larvae of Amphiops are thought to pupate exposed on the stems of emergent vegetation. A key is provided to the larvae of Australian genera of aquatic Hydrophilids other than Notohydrus Balfour-Browne, Paranacaena Blackburn, Phelea Hanson and Agraphydrus Regimbart, whose larvae remain unknown. CHS Watts, South Australian Museum, North Terrace, Adelaide, South Australia 5000. Manuscript received 23 April 2001. The Hydrophilidae (sensu Hansen 1991) comprise a major part of the Australian aquatic beetle fauna, occurring in all types of fresh water and most commonly among vegetation at the edge of standing water. Adults and larvae occur together. The taxonomy of the adults is now relatively well known thanks to the work of Hansen (1991) at the generic level and above; and that of Gentili (1980, 1992, 1993, 2000) and Watts (1987, 1988, 1989, 1990, 1995, 1998a, 1998b) on the aquatic fauna at the species level. No recent work has been done on the species level taxonomy of the extensive portion of the family in which both adults and larvae are terrestrial. In contrast to the situation with adults, the only work specifically on Australian larvae is that of Anderson (1976) who described the larvae of Helochares tristis (W. MacLeay), Enochrus maculiceps (W. MacLeay) and Chasmogenus nitescens (Fauvel) together with details of their life histories. One feature of the Australian aquatic Hydrophilid fauna is its low endemicity at the generic level. Consequently, descriptions of a number of Australian genera are available in scattered Northern Hemisphere publications. The work on the New World genera was summarised and added to by Archangelsky (1997). This work included a number of genera which also occur in Australia. For a number of years I have been rearing field- collected larvae of Australian aquatic Hydrophilids with the aim of discovering and describing the larvae of all genera and, for those already known from non-Australian species, to extend the descriptions to incorporate Australian species. The basic motivation was to produce a key to enable the larvae of all Australian aquatic Hydrophilidae to be identified at least to genus. As a result I have reared 12 species in 9 genera. Two additional genera, Allocotocerus and Regimbartia, were identified by associating adults and larvae by the biochemical technique of alloenzyme electrophoresis. Others were identified using the descriptions in Archangelsky (1997), eg Hydrobiomorpha;, or by association and elimination, eg Enochrus mastersi. I have not attempted any phylogenetic analysis 106 since, with such a high proportion of genera also occurring widely outside Australia, the only sensible approach would have been to include an examination of all known genera, which was beyond the scope of this study. Some comments on relationships are given under most generic descriptions. In preparing this paper I have drawn heavily on the work of Archangelsky (1997) on the New World fauna. The descriptions follow his format and a number of the figures used to illustrate the key come from his important publication. Although their larvae are known from other places, I have not seen Australian examples of the following genera: Chaetarthria Stephens, Chasmogenus Sharp, Crenitis Bedel, Coelostoma Brulle or Paracymus Thomson, and since I have nothing to add have not included descriptions. Detailed descriptions can be found in Archangelsky (1997). In addition, the larvae of Paranacaena Blackburn, Notohydrus Balfour- Browne, Phelea Hanson and Agraphydrus Regimbart are unknown. MATERIALS AND METHODS Larvae were collected in the field and reared in small aquaria formed by placing the base of a small petrie-dish inside a larger petrie-dish and filling the space between them with damp sand (Watts 1963). Mosquito and chironomid larvae were the main food items provided. These were accepted by all larvae other than those of Hydrophilus (see under Hydrophilus). All species successfully reared pupated in cells constructed in the wet sand with the exception of those of Amphiops which pupated on the surface or attached to the sides of the container. Larvae were preserved in 75% ethanol and examined under a stereomicroscope. Permanent microscope slides were prepared for detailed examination. Drawings were made with the aid of a camera lucida. Habitus drawings were only prepared when no previous illustration was available or when Australian species differed from those previously illustrated. Unless otherwise mentioned the descriptions are based on third instar larvae. Earlier instars usually differ somewhat, most noticeably in the stouter cephalic appendages and in fewer spines on the mentum. The key is constructed to accommodate all instars. Measurements of the head capsule were made under a stereoscopic microscope with a graduated eyepiece. Measurements of total CHS WATTS length were made with dial callipers on both mounted and alcohol preserved material, but in this case the measurements can only be indicative as factors such as larval age and preservation effects can alter the length considerably. Measurements of the head capsule, particularly the width, are a more reliable indicator of size. Unless stated otherwise, the specimens were collected by myself and are in the collection of the South Australian Museum. SYSTEMATICS KEY TO THE GENERA OF AUSTRALIAN AQUATIC HYDROPHILIDAE BASED ON LARVAL CHARACTERS 1) Not including the genera Notohydrus, Paranacaena, Phelea and Agraphydrus, whose larvae are not known. 2) Including Spercheus which is in the family Spercheidae but which has larvae that can be confused with Hydrophilidae. 1 — _ Tip of mandible bifid (Fig. 2g) ............. yderkeyedaapeigegi ok Spercheidae (Spercheus) — _ Tip of mandible not bifid ..................... ceuetesieeeten Ae gas Hydrophilidae. ..2 Hypopharyngeal lobe well developed, like a pubescent tongue, originating at the base of the labium on the left side (Figs 2a, 2b) Subfamily Sphaeridiinae* .......0..00..... bupsisatosdyestosissssparsorsee Coelostoma Brulle — Hypopharyngeal lobe reduced or absent paths tinahabet Subfamily Hydrophilinae...3 Abdominal segments with long filamentous gills (Fig. Sa), or with multiple setose lateral projections on abdominal and thoracic segments (Fig. 15a) .......... pesdboodEodnsedaacds sthabharaee Tribe Berosini...4 — Lacking long filamentous abdominal gills or complex lateral projections, at most with simple short fleshy projections (Figs 4a, 14a, 16a) oo. eecccseeeesesseeseessenes 6 Gills long, without setae (Fig. 5a); labroclypeus asymmetric (Fig. 5b); mandibles asymmetric (Fig. Sf); prementum small and squat, without ligula (Fig. 5c), basal segment of antenna without lateral projection (Fig. Se) 0... eee eee SinacaSeaap evleny HHitepavier abun 23 Berosus Leach — Abdominal and thoracic segments each with two or more setose, lateral projections 3 = LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 107 (Fig. 15a); mandibles nearly symmetrical (Fig. 15h); prementum long and thin, with ligula (Fig. 15d); basal segment of antenna with lateral projection (Fig. 15g) ........ 5 Body spines predominantly blunt, weakly bifid at tips (Fig. 3b); lateral projection on basal segment of antenna near apex, sharp, spine-like (Fig. 3f); apex of second segment of antenna with lateral extension (Fig. 3d); maxillary stipe without spines (Fig. 3e); ligula relatively short, prementum stouter (Fig) 36) <....0-1.. Allocotocerus Kraatz Body spines predominantly pointed (Fig. 15c); lateral projection on basal segment of antenna thumb-like, further from apex (Fig. 15g), second segment of antenna without lateral extension (Fig. 15e); maxillary stipe with spines on inside (Fig. 15f); ligula relatively long, prementum more elongate (Fig. 15d)... ee shes ebdgebibetnadsatentcbedeas Regimbartia Zaitzev Apical segment of antenna about same length as penultimate (Figs 12d, 16e); femora with fringe of swimming-hairs (Fig. 2h); with prostyles (two retractable, fleshy appendages on last abdominal segment) se Dred aL Subtribe Hydrophilina. ..7 Apical segment of antenna much shorter than penultimate (Figs 9e, 14e); femora lacking, or virtually lacking, swimming- hairs; without prostyles ...........-2eee 9 Labroclypeus without teeth (Fig. 12a); mandibles asymmetrical, left one very robust, right one more slender (Fig. 12e); ligula shorter than first segment of labial palpus (Fig. 12b); up to 40 mm long .... RENE are cet ae Hydrophilus Muller Labroclypeus with weak to moderate teeth (old specimens may lack teeth due to wear) (Figs lle, 16f); mandibles symmetrical or nearly so; ligula longer than first segment of labial palpus (Figs 11b, 16c); up to 20am long. nag oe 8 Prementum longer than mentum (Fig. 16c); first segment of antennae with few spines (Fig. 16f); mandibles each with two large distal teeth and one small proximal denticle (Fig. 16f) 0... Sternolophus Solier Prementum slightly shorter than mentum (Fig. 11b); first segment of antenna with numerous spines on inside (Fig. 11c); mandibles each with one large distal tooth which is pick-shaped at the tip and one or 10 11 12 13 14 two small proximal denticles (Fig. 1 le) wacedeaeseetatins Hydrobiomorpha Blackburn Left expansion of epistoma much more prominent than the right and with a row of stout setae on front edge (Fig. 13b); left mandible with group of stout setae at base of middle tooth which is lacking on the right mandible (Fig. 13f) .......... 10 Lateral expansions of epistoma similar, without a row of stout setae on front edge (Fig. 14b); mandibles without sucha group C0) Bo] Fo 11 Frontal sulci parallel and not uniting to form a coronal sulcus (Fig. 13a); nasale with three teeth (Fig. 13b); prementum wider than long (Fig 13c); ligula absent or Virtually so (Fig. 13C) oo. sceeseeeeeeeeeeeee pabbadeanetpragaresiitealaryeas Laccobius Erichson Frontal sulci meet just before occipital foramen to form a coronal sulcus (Fig. 10a); nasale with five teeth (Fig. 10b); prementum longer than wide (Fig. 10c); ligula present though small (Fig. 10c) .. ebagagasneaterpetast ete Hybogralius Orchymont Legs very short, without claws, prementum with a large round ligula (Fig. 2e); length up to 6 mm......... Chaetarthria Stephens Legs usually longer, with claws. Prementum with or without ligula, but if present never large and round; length up to USS TO peisk easy oa sca egeeesecbectveeebabeate 12 Mentum large, wider than long, prementum small, lacking ligula (Fig. 4b) .............. Be ceadpomedd st tshe heen dteg’ Amphiops Erichson Mentum square or longer than wide, prementum well developed, ligula present (Figs 6b, 8b, 14c) .... eee ecceeeeeeeeee 13 All abdominal segments with dorsal sclerites, although often small; without coronal sulcus (Fig. 2c); lengthupto6 mm; antennal appendage as long as apical segment; mandibles symmetrical with three teeth. stone Tribe Anacaenini....14 Abdominal segments, other than 1 and 8, lacking dorsal sclerites; with short coronal sulcus (Fig. 1a); antennal appendage half length of apical segment (Fig. 8d); mandibles often asymmetrical and with fewer than three teeth; length up to 15 mm wigan Fes 5 aaa sean pate sev pstegesttupedinateeg’ 15 With lateral abdominal flaps (Fig. 2c); without cervical sclerites Crenitis Bedel 108 CHS WATTS FIGURE 1. a, dorsal side of head capsule of Hybogralius hartmeyeri. 1, maxillary palpus; 2, stipes of maxillary palpus; 3, mandible; 4, mandibular teeth; 5, labium; 6, antenna; 7, right lobe of epistome; 8, teeth of nasale; 9, nasale (7 & 9 make up the labroclypeus); 10, stemmata; 11, frontal sulci; 12, coronal sulcus; 13, cervical sclerites; 14, occipital foramen; b, details of dorsal side of labium of Hybogralius hartmeyeri. 1, labial palpus; 2, ligula; 3, prementum; 4, mentum. LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 109 S