THE LARVAL DEVELOPMENT OF LITORIA BREVIPALMATA (ANURA:HYLIDAE) MARION ANSTIS Anstis, M. 1994 12 01; The larval development of Litoria brevipalmata (Anura:Hylidae), Memoirs of the Queensland Museum 37(1): 1-4, Brisbane, ISSN 0079-8835, The tadpole of L.brevipalmata is unique amongst Australian hylid frogs of the pond dwelling nektonic type in possessing a median vent tube. The embryo is one of a small number of species so far known to have three pairs of external gills, (] Anura, Hylidae, Lisoria brevipalmata, embryonic and larval development, M.Anstis, 26 Wideview Rd, Berowra Heights. New South Wales 2082, Australia; 5 January 1994, Litoria brevipalmata Tyler, Martin & Watson, 1972 is the sole member of the Litoria brevipalmaia Species group of Tyler & Davies (1975). A medium- size, ground-dwelling species, it is readily distin- guished from congeners by the combination of a rich brown dorsal colour and pale green in the axilla, groin and posterior surface of the thighs. The type description was based on a series of adult frogs collected at Durimbah Ck near Gosford, NSW, and Byaburra (near Wauchope) NSW, but no information on life history was presented. Mc- Donald (1974) reported its occurrence at Ravensbourne and Crows Nest National Parks in SE Queensland, and the frog was found sub- sequently at Jimna, 80km north of these localities (Czechura, 1978) and in the Kilcoy Shire (McEvoy etal., 1979). Barker & Grigg (1977) stated that large numbers of males were calling afterrain in a flooded paddock near Gosford (late October, 1972), and noted that juveniles were collected during April in wet sclerophyll forest, Two more recent records are now known from near Woogaroo Ck near Wacol, Brisbane (Nattrass & Ingram, 1993). This is the only published coastal locality so far for the species. However, in the summer of 1993/1994 there have been further records from Marsden, Karawatha, Beerwah and Nambour (G. Ingram, pers. comm.) The present paper describes the only ayailable embryo and larval material, MATERIALS AND METHODS The following description is based on an egg mass from onc pair of frogs collected by H.G. Cogger on 29 October, 1972. The pair was taken from beside a permanent pond in a paddock near Ourimbah Ck, Gosford NSW after heavy rain and amplexus oc- curred in a plastic bag, The eggs were laid during the evening of 30 October, Most embryos died in transit, One was preserved at stage 9, five at stage 17, one al stage 20 (just hatched), one tadpole at each of stages 37 and 41 and one newly metamor- phosed at stage 46. Specimens are lodged in the Australian Museum (AMR118482, AMR] 18483), Tadpoles were raised in an outdoor container of pond water expenencing variable water tempera- tures (16-25°C), and were fed on boiled lettuce. Tadpoles were anaesthetised in Chlorbutol solution, then embryos and tadpoles preserved in Tyler's (1972) Fixative, Specimens were measured with a yemier caliper to 0.01mm, or an ocular micrometer allached Lo a stereoscopic microscope. The staging system is that of Gosner (1960). Abbreviations for larval measurements (Table 1) refer to Alig (1970), Anstis (1976) and McDiannid & Altig (1989) as follows: In lateral view: TL = total length; BL = body length; BD = max. body depth; TD = max. tail depth; TM = depth of tai] musculature (in line with TD), BTD = tail depth at body terminus; BTM = depth of tail musculature at body terminus; E=eye diameter; S = diameter of spiracle at opening; SN = snout to naris; SE = snout to eye; SS = snout to spiracular opening; DF = max. depth of dorsal fin; VF = max. depth of ventral fin. In dorsal view: BW = max. body width across abdomen; EBW = width of body at level of eyes; BTMW = max. width of tail musculature at body terminus: IO = inter-orbital span; IN = inter-nanal span; EN =distance from eye ta naris, a In ventral view: MW = transverse width of oral ise. Illustrations were drawn using a drawing tube attached to the stereoscopic microscope. DESCRIPTION OF EMBRYOS The single female laid 556 eggs. It was not pos- sible to make observations on the form of the egg mass, as a result of disturbance during transit. Ova were al stage 9 when firstobserved on | November, 1972, One embryo preserved at this stage had a dark brown animal pole, off-white yegetal pole and measured 2,02mm in diameter, b4 FIG. 1, Embryological stages of L.brevipalmata. A, early slage 17; B, stage 18; C, stage 20. Scale bars = 1 mm. By 2030 on 1 November, the embryos had reached stage 17. The tail bud is bent in this stage, making most measuremenis of embryo length ap- proximate. Embryos preserved at early stage 17 (Fig.1A), have optic bulges, prominent gill plate bulges, a slight pronephric swelling and fine, closely aligned muscular ridges along the neural tube. There are prominent U-shaped ventral suckers joined by the stomodaeal groove. The embryos are pale brown with a pale cream yolk sac. There are a small number of melanophores around the gill plate and neural tube, and the ventral suckers are heavily pigmented. An embryo at stage 18 has very prominent gill plate bulges (Fig. 1B). The stomodaeal groove is deeply furrowed, its rim protruding strongly for- ward. There is a small oral sucker at each corner. The optic bulge is quite indistinct and the tail fins are beginning to develop. The embryo is very pale cream in preservative. The first specimen hatched at stage 20 on 2 November, 3 days after the eggs were laid. The live embryo is dark grey-brown with a light brown yolk sac, The optic region is barely discernable and the oral suckers are prominent, being still joined along the anterior edge of the stomodaeal groove, and divided medially on the posterior edge. The indis- MEMOIRS OF THE QUEENSLAND MUSEUM tinct narial pits are faintly outlined with pigment. There are three pairs of ex- ternal gills; 3 branches on the uppermost pair, 7 on the middle pair and § on the lowest (Fig.1C). The tail musculature ridges arc developing and the fins are translucent. DESCRIPTION OF LARVAE The only larvae available for study were one specimen al stage 37 and one at stage 4]. The former specimen (Fig. 2) is described as fol- lows: Body ovoid in shape, widest across mid-region of abdomen. Snout broad and truncate in dorsal view and truncate in lateral view. Nares small, opening antero- laterally and outlined by a fine border of melanophores. Eyes lateral. A narrow ridge Tins from from each eye to each naris, Spiracle sinistral, ventro-lateral, not visible from above, opening postero-dorsally, with tube diameter decreasing markedly from origin lo opening. Vent tube broad, median in position and opening to a diamond-shape when expanded, When relaxed, the aperture has a ‘<° shape, with point of ‘<’ directing anteriorly (Fig. 2C). Opposite point of diamond-shaped opening (>) attached to edge of ventral fin. Tail fins arched, tapering to a fine point, The dorsal fin extends onto the body up to the mid-point of the abdominal region (Fig. 2B) and in lateral view, fin is deepest just anterior to its mid-point. Ventral fin marginally deepest anterior to its mid- point. Tail musculature deepest at body terminus, then narrows, before broadening slightly near mid- point and tapering to a fine flagellum. Oral disc antero-ventral, bordered by small mar- ginal papillae around all but medial anterior third. A small number of submarginal papillae present. Labial teeth in two complete antenor and three complete posterior rows, each set approximately equal in length (Fig. 3). Keratinised jaw sheaths narrow, with fine serrations along the somewhat irregular inner edges, LARVAL DEVELOPMENT OF LITORIA BREVIPALMATA 3 FIG. 2. Larva of L.brevipalmata at stage 37. A, dorsal view; B, lateral view showing cream internarial patch visible only in life; C, ventral view. Scale bars = Imm. COLOUR IN LIFE Dorso-lateral surface uniform dark brown, with a small cream patch on snout between nares (Fig, 2A). Dorsal surface of tail musculature dark anteriorly, lightening posteriorly. Ventrolateral surface with a grey blue sheen. Fins mostly transparent, with some light sandy pigment along the tail musculature in lateral view. Cream patch on snout barely distinguish- able on specimen at stage 41. COLOUR IN PRESERVATIVE Dorsal surface of specimen at stage 37 dark brown (darkest over the intestines), but cream patch has disappeared. Tail musculature only slightly lighter than body anteriorly, becoming pale cream posteriorly. Hind limbs show some patches of melanophores. Ventro-lateral region over intestinal mass mainly dark blue-grey, with part of intestine and fore-limb visible. Remainder of body dark brown, except for tranlucent region around vent. Fins and musculature have fine dusky pigment diffused over entire surface, increasing slightly towards body terminus. Musculature bordered above and below by fine line of pigment. Specimen at stage 4] has more pigment over tail and body, and body wall is slightly less trans- lucent. Ventral surface dark blue/grey over intestines and anterior half translucent, with fine, diffuse pigment. Metamorphosis was reached on 26 December 1972, 57 days after the eggs were laid. One specimen preserved immediately at stage 46 was 12.7mm in length and shows the uniform brown of the adult over dorsum and limbs. The white labial stripe is present and there is a black canthal stripe from tip of snout to eye. A similar stripe is just beginning in the post-orbital region above the tympanum. At *6 power, numerous scat- tered, fine tubercles cover the dorsum, The ventral surface is white. The green pigment present in the axilla, groin and thighs of the adult, as yet has not developed. DISCUSSION EMBRYOLOGICAL DEVELOPMENT L.brevipalmata is unusual in possessing 3 pairs of external gills. Only one other Australian hylid species has been described as having 3 pairs (L.chloris, see Watson & Martin, 1979). LARVAE The tadpole of L.brevipalmata is the only len- tic, nektonic Australian hylid species yet described as possessing a median vent tube. Prior to about stage 41, live tadpoles may be distin- guished from other sympatric ground-dwelling FIG. 3. Mouthparts of L.brevipalmara larva at stage 37. Scale bar = Imm, 4 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1, Measurements in mm. A, embryos; B, larvae, C, metamorphosis. jam ee ee ee HI i^ Win 2.7 (2.59-2.88 3.32 ae |" sag | 20 28.9 29.4 Ee ee ee a T] ee n | om | | BD | 54 NA 262 | E | iw č | ig | | s | ome | os | | wN ë | o» | o | | æ | o» | im || | s 'Ŵ' | hylid species such as L. freycineti, L.latopalmata (Anstis, unpubl.), Lnasuta (see Tyler et al., 1983) and L. lesueuri (see Martin et al., 1966) by â combination of a uniform dark brown body colour, a small cream patch on the snout, median vent tube opening and two unbroken anterior rows of labial teeth. ACKNOWLEDGEMENTS Acknowledgement is made to the Australian Museum for the loan of specimens. and to Stephen J. Richards and Jean-Marc Hero of the Department of Zoology, James Cook University for helpful suggestions on the manuscript. LITERATURE CITED ALTIG, R. 1970. A key to the tadpoles of the United States and Canada. Herpetologica 26: 180-207, ANSTIS, M. 1976. Breeding biology and larval development of Litoria verreauxi (Anura:Hylidae). Transactions of the Royal Society of South Australia 100; 193-202, BARKER, J. & GRIGG, G. 1977. ‘A field guide to Australian frogs’. (Rigby: Adelaide), COGGER, H,G. 1975, ‘Reptiles and amphibians of Australia’, (Reed: Sydney), CZECHURA, G.C. 1978, A new locality for Litoria brevipalmata (Anura: Pelodryadidae) from South-Eastem Queensland. Victorian Naturalist 95: 150-151. GOSNER, K.L. 1960. A simplified table for staging anuran embryos and larvae with notes on iden- tification. Herpetologica 16: 183-190. MARTIN, A.A., LITTLEJOHN, MJ. & RAWLIN- SON, P.A. 1966. A key to anuran eggs of the Melboume area, and an addition to the anuran fauna. Victorian Naturalist 83: 312-314. McDIARMID, R.W. ALTIG, R. 1989. Description of a bufonid and two hylid tadpoles from Western Ecuador, Alytes 8: 51-60, McDONALD, K.R. 1974. Litoria brevipalmara. An addition to the Oueensland amphibian list. Her- petofauna 7 (1): 2-4. McEVOY,J,S., McDONALD, K.R. & SEARLE, AK. 1979. Mammals, birds, reptiles and amphibians of the Kilcoy Shire, Queensland. Queensland Jour- gal of Agriculture and Animal Sciences 36; 167- 180. NATTRASS, A.E.O. & INGRAM, G.J. 1993. New records of the rare Green-thighed frog. Memoirs of the Queensland Museum 33: 348. TYLER, M.J., CROOK, G.A. & DAVIES, M. 1983. Reproductive biology of the frogs of the Magela Creek system, Northern Territory, Records of the South Australian Museum 18; 415-440, TYLER, M. J. & DAVIES, M. 1978. Species groups wilhin the Anstralopapuan hylid frog genus Litoria Tschudi, Australian Journal of Zoology, Supplementary Series 63: 1-47. TYLER, M.J., MARTIN, A.A. & WATSON, G.F. 1972. A new species of hylid frog from N.S.W, Proceedings of the Linnaean Society of New South Wales 97: 82-86. WATSON, G.F. & MARTIN, A.A. 1979, Early development of the Australian green hylid frogs Litoria chloris, L.fallax and Lgracilenia. Australian Zoolologist 20 (2): 259-268. A PRELIMINARY ASSESSMENT OF THE SPATIAL DISTRIBUTION OF STONE ARTEFACTS FROM THE SOUTH MOLLE ISLAND ABORIGINAL QUARRY, WHITSUNDAY ISLANDS, CENTRAL QUEENSLAND COAST. BRYCE BARKER AND RICHARD SCHON Barker, B. & Schon, R. 1994 12 01: A preliminary assessment of the spatial distribution of stone artefacts from the South Molle Island aboriginal quarry, Whitsunday Islands, central Queensland coast, Memoirs of the Queensland Museum 37(1):5-12. Brisbane. ISSN 0079- 8835. Distribution of artefacts made from a black tuff raw material type in the Whitsunday region are traced by morphological similarities and petrographic analysis to an Aboriginal quarry source on South Molle Island. Although the petrographic sample size is small and relatively heterogenous there are strong indicators that the source of nearly all the volcanic (off artefactual material found on the Whitsunday Islands and parts of the adjacent mainland are from this source, Explanations for the pattern of distribution include, proximity to raw material source, regional interaction of social networks and the extent of the Ngaro ‘tnbal” system. [_] Black tuff, sourcing, South Molle Aboriginal quarry, petrography, Whitswndey Is. B, Barker, Department of Anthropology und Sociology, University of Queensland; R, Schur, Depurtment of Geology, University of Queensland, Brisbane. Oueensiand 4072, Australia; T5 June 1994. This paper presents preliminary results of a study aimed at sourcing stone artefacts, found in the Whitsunday region, to a large Aboriginal quarry on South Molle Island. South Molle Island is one of the Cumberland Islands (northem Cum- berlands are termed the Whitsunday Islands) lo- cated on the central Queensland coast (Fig. 1). The northern Cumberland group are, according to Tindale (1974); the domain of the Ngaro, whose country extended from Hayman Island in the north down to St Bees Island in the south, including Cape Conway on the mainland coast. Surveys and excavations from islands and the adjacent mainland between Bowen and Midge Point have established an overwhelming homogeneity of artefact raw materials (Barker, 1988, 1989a, 1989b, 1990, 1991a, 1991b, 1992a. 1992b, 1992c, 1993; Hall & Barker, 1989). This material is generally characterised as a black siliceous volcanic tuff and appears identical to the raw material found on the South Molle Island quarry. It was considered that if the black tuff raw material found in the region came from a source or sources other than the quarry on South Molle. that this would be reflected in its pattern of dis- tribution. This paper therefore, examines the dis- tribution of stone artefact raw materials in relation to known quarry sources in the region, and presents the preliminary findings of a petrographic analysis carried out on raw material from the South Molle Island quarry, and on various artefacts from islands and coast as far distant as 100km. It is argued that the majority, if not all, of the black tuff artefacts found on the islands, and on the adjacent mainland coast, are from this source. The rationale for this argument include: 1, the relative intensity of quarry use; 2, the fact that it is the only known culturally ex- ploited source of black tuff material on the is- lands; 3, the quarries central location to all the known sites in the region. The preliminary results from the petrography will be discussed in relation to this model below. THE SOUTH MOLLE ISLAND QUARRY South Molle Island is a small offshore island of 420.5hectares, approximately 2 km from the mainland. The quarry (SMIQ) is located on the south eastem side of the island, on top of a prominent ridge just above Bauer Bay the main resort beach. Quarried raw material covers an area of 300m along a steep ridge, with flakes and other artefacts scattered down the eastern side for over 200m, and down the western side for 500m. The artefacts are up to Im deep in places, Three circular pits, where material has been mined, linc the top of the ridge (Figs 2, 3A,B), The quarry is dominated by extremely large flakes which haye been removed from the substrate and then used as cores (technologically they are retouched flakes). A number of round water rolled cobble ham- merstones from the beach are also present (Fig. 3C). The actual substrate from which the raw material has been struck is almost completely covered by huge densities of artefact discard (Fig. 3D). Seventy five large, backed asymmetrical henn | | | TB Mig Syrup a L2 ” | oe 7 h ay say NS Woe, [Tima t a z3 Me T u have yp oi a Ty l | U un 'n 4 ma Y n NN A AW j Ly a wmi mwN Ps lay’ ~ “e| | t ha pe A ^N Lint ne ee SN a \ AU Matte | Wh - | \ a. ì p n i “Vitaly ji ON E" Praa i" ì - AX l] Ale | Hei» ŷ FIG, 1, The study region, blades (‘Juan Knives’) were collected from the quarry site and the beach below the quarry (2 have also been found on islands) (Barker, unpubl.) in prep).The archaeological excavation of in-situ black tuff artefacts from basal strata in Nara Inlet 1 rockshelter on Hook Island, (approximately 18km east of South Molle) demonstrates the early use of the black tuff material. The near basal date of Nara Inlet 1, indicates that a minimurm age for the use of the South Molle quarry can be placed at sometime prior to 8150BP (Barker, 1989a, 1991a). Analysis of excavations undertaken in 1992 of a rockshelter directly below the quarry, and on the slopes adjacent to the guarry itself, may shed further light on the guarries antiguity (Fig. 2). Although the South Molle Island guarry would appear to represent large scale and intensive ac- tivity, this view must be tempered with the knowledge that it has probably been in use for at least 8000-9000 years. Recent estimates put the total number of artefacts on the quarry at ap- proximately 7 million (Border, 1993) repre- senting an artefact discard of just 2.1 per day or 766.5 per year. From 5 stratified sites excavated by Barker (1989a, 1991a, 1993) on the Whitsun- day Islands, it is clear that discard of black tuff MEMOIRS OF THE OUEENSLAND MUSEUM artefacts occurred in much greater densities in the early period of oc- cupation of the region (sometime before 8150BP until c.2,500BP). After this period, stone artefact dis- card rates for a 50cm x 50cm ex- cavation square decreased from a maximum of 9.5 artefacts per 100 yeurs in the early phase, to as little as 0.5 artefacts per 100 years during the last c.2500 years (Table 1). The first level of analysis, in regard to the use of the quarry and the distribution of artefacts in the region, was on observed mor- phological similarities and dif- ‘vy. | ferences between raw material "| types, and an examination of the historical record relating to use or occupation of South Molle Island. It was considered reasonable to as- sume that if the black tuff raw material came from South Molle Island, that the highest densities of black tuff material would be found in sites within closer proximity to this source (Renfrew, 1977; Sidrys, 1977; McBryde, 1984). The survey work relating to this study was undertaken as a component of Environmental Impact Statements, or as part of the survey strategy designed for a rescarch Phd project in the region (Barker, unpubl.). The Islands and the mainland coast have been extensively surveyed. The area from Southern Cape Conway down to Midgeton has also been extensively surveyed. TABLE 1. Temporal discard of black tiff artefacts from Border Island 1 /2500-peesen) | o, dr ; #2500-present 19 0.7 Nara Inlet LH *8150-2500bp Pn Ŵr | #2500-present 15 | O5 Nara Inlet 1G *8150-2500bp 14 | ong [Nara init AS1_[-2500-present_| 37 | 15 | *These dates represent basal or near basal dates #These dates corralate with major stratigraphic change and greatly increased densitys of other cultural material. DISTRIBUTION OF STONE ARTEFACTS The hinterland area west of Airlie Beach and the Proserpine area have been less extensively sur- veyed and assemblages from these areas are described from private collections, as well as material held in the Queensland Museum, the Anthropology Museum (Depart- ment of Anthropology and Sociol- | — ogy, University of Queensland), | the Bowen Historical Society and the Proserpine Cultural Centre. = HISTORICAL ACCOUNTS OF ABORIGINAL ACTIVITY ON SOUTH MOLLE The most authoritative reference to the South Molle Island quarry is from Roth (1904:19) who men- tions the site in the context of sour- ces of stone for “stone celts” (edge | Rockshottor SM! 17” A, a “ES on y wy wt ee gm; ground axes). He notes: ‘Quarries whence these celts were originally obtained are none too common, there is one on... Molle Island in the Whitsunday Passage and at Culture Creek, or Happy Valley, about 7 miles from Proserpine’, Two other direct historical refer- ences to the use of the South Molle South Molle island —— ani" me oa Yoon (ng rim ne. a walian [ransecihy ngwr wyn es Ol uffegrd acatar Sepa ato cil auair prt Island quarry comes from an ac- count in a popular magazine by Henry Lamond, a former owner of the lease on South Molle Island and from W.E. Bauer, the founder of the South Molle Island resort. Both accounts are relatively recent (1930,s-1950's) and may tend to be apocryphal (Lamond was a regular contributor of stories to magazines and newspapers). Lamond's source is an old Aboriginal man named Percy, who stated that the local Aboriginal population used to obtain flint for axes fram South Molle Island (Lamond, 1960): '....... they went to what we call South Molle for stone with which to make tomahawks. They called that island “Whyrriba” which meant stone axe. I should know. It was my home for exactly ten years'. Bauer's (1958) account is similar, stating that: ‘the native name for South Molle was "Wer- ribee” which translated means stone flints, We have a range on the Island where this flint is very plentiful, and the natives used to make their stone axes here’. Although not mentioning the quarry specifical- ly, there are other historical references relating to FIG. 2. South Molle [sland quarry (adapted from Border, 1993). the use of the Island itself; for example, Dal- rymple (1860) on the Spitfire in 1860, stated that: ‘on Lindeman and Molle Island in September dense clouds of smoke rose from the fires of the natives’. Dalrymple also describes an encounter with two Aboriginal men in a canoe near South Molle Island, which met them soon after leaving the Port Molle anchorage (and, therefore, probably from South Molle Island itself). Dalrymple (1860:3) states: *.....a breeze springing up, the twe aborigines left us to cross over to Hook Island, a distance of 10 miles’. and from an account of the wreck of the schooner ‘Eva’ it is stated that upon landing on Molle Island the survivors were threatened by hostile Aborigines (Loos, 1982). Although there is clear prehistoric and historic evidence of the use of the island, the archaeologi- cal evidence does not support the idea that the quarry was used for axe manufacture. Extensive 8 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. South Molle Island Quarry. surveys of axes from the mainland including private collections, collections from local histori- cal societies, and those housed in the Queensland Museum, have not found a single axe manufac- tured from anything resembling the South Molle Island raw material. Generally, axes are made from rocks with a high degree of plasticity, unlike the South Molle tuff which is highly siliceous and brittle, with excellent flaking characteristics, Ad- ditionally, no axes or blanks or any artefact resembling a stage of axe manufacture has ever been found on the guarry, or in the near vicinity. It may be that Roth (1904) when stating that the guarry was used for the manufacture of stone ‘celis’ was referring to the large backed blades found on the quarry and the beach below it, although his description of stone ‘celts’ as edge ground axes seems unambiguous. Another pos- sibility is that another source of black tuff material exists, from which axes are manufac- tured and it is this source which Roth (1904) mistakenly refers to as the Molle Island quarry (considered unlikely because of the acknow- ledged accuracy of much of Roth's ethnographic accounts, and because of the lack of any evidence that stone axes were made from black tuff raw material). OTHER RAW MATERIAL SOURCES Although relatively extensive geological surveys have been conducted in the Whitsun- days and adjacent mainland (Paine, 1972; Ewart et al., 1992; Bryan, 1991; Parianos, 1992), only one other source of the raw material resembling that of the South Molle lsland material (a siliceous tuff) has been located (see below). It cannot be ruled out that other sources also occur on the islands. However, as the formation of the raw material is described as a ‘pyroclastic surge deposit' which will usually occupy a relative- ly discreet, well defined area, it may not be easily located (Bryan, 1991). The geological conditions which create the raw material are, however, considered a relatively rare com- bination of factors unlikely to be duplicated often (Bryan, 1991). A number of other raw DISTRIBUTION OF STONE ARTEFACTS 9 TABLE 2. Petrographic descriptions. |Type _|Artefact [Location |Description | Sh Mote sw [swe feet - [o [swn Ene gd ld [EM — elie tuff, black to green- [Sth Molle_ | Scalte in colour, with [Sth Molle _|sca red patches of calcite alteration and guartzite/albite. The calcite alteration is characteristic of these implements Artefacts are [Border] | Safacterised by a Border! |fine and even-grained Awe tuff with sparse deformed Whitsunday |fiamme and bedded opagues Repulse I Fine grained, strongl welded: silicihed wr Percy I ign nimbrite with glass ards measuring up tol mim Fine grained even- textured silicified tuff with sparse quartz veins and bedded paques More fine raipat than the o! materials with scattered ‘ood th caste and UNCF {SM5 traversed veins. Thee oles distinguish it from the SM types, to which it jis otherwise similar ideal; sources'are known for the region, These are as follows (Fig. 5): 1, Material that superficially looks similar to the South Molle Island source was observed in a commercial quarry used by the Pioneer Shire Council. The source is located near Kuttabul (north of Mackay), 12km inland and 90km south of South Molle Island. Unlike the South Molle Island quarry, however, there is no evidence that this source was utilised during prehistoric times, although it does confirm the presence of a similar raw material type in the region. 2, A quarry source at Happy Valley on ‘Culture Creek’ (possibly Kelsey Creek) 11km west of Proserpine, is known from the literature (Roth, 1904) but has yet to be rediscovered or charac- terised. However, artefacts from around the Proserpine region are commonly made on stone that is morphologically dissimilar to that found on South Molle Island, making it unlikely that the raw material from this source ts black tuff. # Yul Bessi Eme — | Fiasergie Hinserand iskanje Munland Coast A Maininnd gaea FIG. 4. Raw material sources and distribution within geographical areas, 3, Another raw material source is to be found at Blue Bay on Cape Hillsborough, where beach shingle has been quarried along an exten- sive area of the eastern end of the headland separating Blue Bay from Halliday Bay. This raw material is a blue-grey volcanic rock, mor- phologically dissimilar to the South Molle material. 4, A green volcanic material is found on South Repulse Island. Artefacts from this source have been located on South Repulse Island and Cape Conway. In addition to these sources, the volcanic nature of the region suggests that there would be no shortage of brittle, siliceous raw material sources in the region, a raw material category ideal for stone artefact manufacture. DISTRIBUTION OF BLACK TUFF Most of the black tuff artefacts are found on the Islands (Fig. 5, Table 3). The mainland coast between Bowen and Southern Cape Con- way (Area A Mainland coast) includes a reta- tively high percentage of black tuff artefacts, but also includes a range of other local raw material types. The area west of Airlie Beach, including the Proserpine region (Proserpine hinterland), has little of the material resem- bling the black tuff and is dominated by ar- tefacts made from other materials. The area from Repulse Bay down to Midgeton (Area B, Mainland coast) contains no evidence of black tuff material, and is again dominated by a variety of other raw material types (Fig. 4), On this basis, it can be said that the distribution of the black tuff material, with the exception of the Proserpine hinterland, is aligned to the presence of the quarry source, with greater den- sities appearing nearest to the source, 10 MEMOIRS OF THE QUEENSLAND MUSEUM FROSERFINE Ta “nn HINTERLAND Nn ral pne e (er yg “Y - + a 3 = | eii FIG. 5. Number of sites with different raw matenals per geographcal area. 'The distribution of the material, as outlined above was then further tested by attempting to link the artefactual material to the guarry through petrographic analysis. THE PETROGRAPHIC ANALYSIS A petrographic classification of various black tuff artefacts from the region was undertaken to determine whether or not the stone artefacts found in archaeological sites on the islands, and adjacent mainland, indeed came from the black tuff quarry source on South Molle Island, Eleven artefacts were thin-sectioned to allow classifica- tion by petrography. The aims of this analysis were to iniliate a preliminary geological charac- terisation of the South Molle quarry material; to establish the number of different lithic types of artefacts in the region; and to see if these could be sourced to the South Molle Island quarry. All materials examined were extremely fine-grained and classification was, therefore, made on the basis of alteration of phenocryst assemblages. Two main groups were identified. These are char- actensed here as SM and SR types (Table 2). ŴY ISLANDS SM type. Includes samples SM2 (South Molle guarry), SM4 (South Molle Quarry), WII (Whitsunday _ Island), probably SM1 (South Molle quarry) and possibly SM3 (South | Molle quarry). These artefacts are characterised by a fine-grained silicified tuff, black to green-grey in colour, with scattered patches of cal- cite alteration and quartz/albite. SR type. Includes samples SR1 | (South Repulse Island), BI (Border 2 a Island) and HI1 (Hill Inlet Whitsun- gn day Island). These artefacts are char- acterised by a fine and even-grained silicified tuff, with sparse deformed fiamme and bedded opagues. wy The following samples were un- classified: | MP1 (Middle Percy Island) -a fine ~ grained strongly welded silicified ig- | nimbrite with glass shards measur- ing up to Imm. ER (Earlando Bay) - fine grained even-textured silicified tuff with sparse quartz veins and bedded opa- ques, (except for the quartz, veins, this sample could be grouped in the SR type). SM5 (South Molle Island) - more fine grained than the other materials, with scattered opaques and traversed by quartz veins. These latter distinguish it from the SM type, to which it is otherwise similar. DISCUSSION Although the sample size is small, some generalisations can be inferred from the data (a more detailed geological characterisation of the quarry site is currently being carried out by the Queensland Department of Environment and Heritage). Most of the material supplied from the quarry (SM2, SM4, SM1 and SM3) can be clas- sified as coming from a single source. The dis- tinctive SM5 sample (also from the quarry and resembling the SM type) suggests that the raw material from the South Molle Island quarry ts extremely heterogenous petrographically, and that a large sample of the quarry material will have to be thin sectioned in order to fully classify it geologically. It is considered that once this is done a wider range of material may also be able to be traced to the South Molle quarry source. Most of the remaining samples, (both SR and DISTRIBUTION OF STONE ARTEFACTS {1 TABLE 3. Number and percenlage of sites with different raw materials | mem | omer | Total # of sites with artefacts pêr geographical area. Toif ao PEE o Isiands | 23 | 92 | a | a | | OMBRE | Proserpine | hinterland unclassified types) are superficially similar in appearance to the South Molle raw material. We feel that once further geological analysis is com- pleted on the South Molle Island quarry that these types will be characterised as having come from that source, The SRI sample is a backed implement of ercen tuff similar to a raw material source found on South Repulse Island and thus probably not from the South Molle Island quarry. MP1 from Middle Percy Island 230km south of South Molle Island is the most dissimilar of the raw material types and js thus unlikely to be from the South Molle source. Although the initial petrographic results are largely inconclusive, the preliminary results regarding the distribution of broader raw material types in the region, specifically the black tuff material, show that this is used almost exclusively on the islands from Hayman down to (at least) the Repulse Islands, and on the mainland coast from Bowen to Cape Conway. We feel confident that once a larger sample of the quarry has been attained, most of the artefacts found on the Whit- sunday Islands (northern Cumberlands) and the adjacent mainland coast will be traced to this source. At this stage it cannot be said conclusive- ly from the petrographic evidence that all the black siliceous tuff material found on the islands and adjacent mainland are from the one source, However, it is thought that the geographical dis- tribution of morphologically similar material clusters around the major Aboriginal artefact quarry source on South Molle Island, suggesting that the South Molle quarry is the main, if notonly source of artefactual black tuff material in the Whitsundays (northern Cumberlands) (Fig. 4). refs] + a] Pe [halle ere rae sw CONCLUSION The preliminary results of this ) study show that the distribution of the black tuff raw material is largely | restricted to areas within the Ngaro system. either on the islands of the | 35 _ | northern Cumberlands or the ad- jacent Cape Conway mainland coast. This distribution fits very closely with the boundaries for the Ngaro outlined by Tindale (1974), largely confirmed by research of onc of us (B.B.) demonstrating that ! the quarry on South Molle Island is mainly a raw material source for local use, despite its use over a long period of time. It could be surmised, due to the abundance of suitable raw materials throughout the wider region, that raw material of the quality of the South Molle source is common-place and, there- fore, is not as likely to be used for trade or carried great distances, The highest number of sites con- taining this material, as well as the highest den- sities of this artefactual material, are those within close proximity to the known source on South Molle Island, and are found where distribution by canoe provides the easiest movement (Islands and Area A Mainland coast). It is considered that the absence of the raw material on the mainland coast south west of Cape Conway (Mainland coast B) and the coastal hinterland around Proser- pine (Proserpine hinterland) and its presence at Queens Beach, Bowen over 70km to the north, may delineate the degree of interaction between the island/coastal peoples and groups directly to the south, west and north of the Whitsunday Islands. The preliminary research carried out on the quarry and the artefacts from the archaeologi- ca] sites needs further investigation. This could include technological analyses and ad- ditional geological work in order to more completely characterise the raw material from the quarry and artefacts in order to more conclusively link them to the quarry source. ACKNOWLEDGEMENTS This paper was presented at the Australian Ar- chaeological Association Conference, Darwin, December 1993. I would like to thank Lara Lamb, Bruno David and Harry Lourandos for commenting on versions of this paper; Richard Robins who provided some valuable structural 12 MEMOIRS OF THE OUEENSLAND MUSEUM advice as referee; Andrew Border for access to artefacts from his research area and for use of an unpublished report, J am also indebted to Jim Gaston and John Locke from the Whitsunday Aboriginal Co-op and Arty Jacobson and team from the Department of Environment and Heritage for input during field work. LITERATURE CITED BARKER, B. 1988. An archacological assessment of the proposed Midge Point Resort Complex, Midge Point, Proserpine. (Unpublished report, University of Queensland Archacological Ser- vices Unit), 1989a. Nara Intec 1: A Holocene sequence from the Whitsunday Islands, centra] Queensland coast, Queensland Archaeological Research 6; 53-76. 1989b. 'A preliminary Archaeological Survey of the Proposed Saro's Cove/Endeavour Cove Development, Midge Point, Pioneer Shire, Central Oueensland Coast', (Unpublished report, University of Queensland Archaeological Ser- vices Unit). 1990. ‘Archaeological assessment of the proposed resort complex. development at Midge Point, Midge Mountain, Pioneer Shire, Proserpine’. (Unpublished report, University of Queensland Archaeological Services Unit), 1991a. Nara Inlet l:coaslal resource use and the Holocene marine transgression in the Whitsun- day Islands, central Queensland. Archaeology in Oceania 26; 102-109, 1991h. ‘An Archaeological Assessment of the Proposed Resort and Marina Development at Shute Bay, Shute Harbour’, (Unpublished report prepared for Scotex Pty Ltd, Sydney). 1992a, ‘A Heritage assessment of Dent Island, central Queensland coast’. (Unpublished report prepared as part of an Environmental Impact Statemeni on behalf of Hamilton Island Enterprises Pty Lid), 1992b. *An assessment of the Historic/Prehistoric Cultural Resource Values and formulation of a Heritage Management Plan for the Whitsunday Islands (northern Cumberland group)’. (Un- published report prepared for Conway National Park, Airlie Beach). 1992c. An assessment of the Historic/Prehistorie Cultural Resource Values of the Mainland Whit- sunday Coast. (Report prepared for the Depast- ment of Environment and Heritage, Townsville Queensland), 1993. Barly human exploitation of island environ- ments within the Great Barrier Reef Marine Park, Reef Research 3, BAUER, W.E. 1958. Letter to Mr. Ken J, Monis, Deputy Premier and Minister for Labour and ladusuy, Office of the Minister for Labour and Industry, Brisbane. BORDER, A, 1993, South Malle Island Stone Artefact Quarry Archaeological Survey. (Unpublished report to the Queensland Department of Environ- ment and Heritage). BRYAN, S.E, 1991, Geology and Geochemistry of the Southern Molle group, the Whitsundays, parth- east Queensland. (Unpublished Bsc. Hons. Thesis; Department of Geology, University of Queensland). DALRYMPLE, G.E. 1860. ‘Report of the Proceedings of the Queensland Government Schooner “Spit- fire” in search of the mouth of the River Burdekin’, (T.P. Pugh: Brisbane). EWART. A., SCHON R., CHAPPELL, B.W, 1992, The Cretaceous volcanic-plutonic province of the central Queensland (Australia) coast-a rift related 'calc- alkaline’ province, Transactions of The Royal Society of Edinburgh 83; 327-345. HALL, J. & BARKER, B. 1989, An archaeological assessment of the proposed Woodwark Bay Resort complex, Woodwark Bay, Airlie Beach, (Unpublished report, University of Queensland Archaeological Services Unit), LAMOND, H.G. 1960, An Island Tribe. North Australian Monthly 7; 35-40. LODS, N, 1982, ‘Invasion and Resisiance- Abariginal//European relations on the North Queensland frontier 1861-1897'. (ANU Press: Canberra). PAINE, A.G.L. 1972, Proserpine 1:250,000 geological Series:Explanatory notes, (Australian Govern- ment Publishing Service: Canberra). MCBRYDE, I. 1984. Kulin greenstone guarries:thc so- cial contexts of production and distribution for the Mt William site, World Archaeology 16:267-285, PARIANOS, J, 1992, Geology of the Airlie Block, {Unpublished Msc, Department of Geology: University of Queensland), RENFREW C, 1977, Alternative models for exchange and spatial distribution.Pp. 71-90. In Earle, T.K. & Ericson, J.E. (eds), “Exchange systems in Prehistory’. (Academic Press: New York). ROTH, W.E. 1904, Domestic Implements, arts and manufactures. North Queensland Ethnography Bulletin 7, (Department of Public Lands: Bris- bane). SIDRYS, R. 1977. Mass-distance measures for the Maya obsidian trade. Pp, 91-108, In Earle, TK, & Ericson, J.E. (eds), "Exchange Systems in Prehistory’, (Academic Press; New York). TINDALE, N.B. 1974. ‘Aboriginal Tribes of Australia’. (Uuiversity of California Press: Los Angeles), THE SPECIES OF HAEMOPROTEUS, LEUCOCYTOZOON AND TRYPANOSOMA OF THE AUSTRALIAN HONEYEATER FAMILY MELIPHAGIDAE (AVES: PASSERIFORMES) GORDON F. BENNETT, DEBORAH SQUIRES-PARSONS AND TARMO POLDMAA Bennett, G. F., Squires-Parsons, D. & Poldmaa, T. 1994 12 01: The species of Haemoproteus, Leycacytozoon and Trypanosoma of the Australian honeyeater family Meliphagidae (Aves: Passeriformes). Memoirs of the Queensland Museum 37(1): 13-18. Brisbane. ISSN 0079-8835, The avian haematozoan species of Haemoproteus, Leucocytozoon and Trypanosoma occur- ring in the Australian Honeyeater family Meliphagidae, first described in 1909 and 1910, are re-described using modem criteria. The confusion surrounding the systematic position of these parasites when they were first described is resolved, [_] Meliphagidae, Haemeproteus, Leucacyiozoon, Trypanosoma, Queensland, avian haematuzoa. G.F. Bennett & D.Squires-Parsons, International Reference Centre for Avian Haematozoa and Department of Biology, Memarial University of Newfoundland, St. John's, New- Joundland, Canada, AJB 3X9; T. Paldmaa, Department of Biology, Queen's University, Kingston, Ontario, Canada, KIL 3NG; 28 July 1994. Cleland & Johnston (1909) described Haemoproteus ptilotis, H. philemon and H, meliornis from the meliphagids Prilotis (= Meliphaga) chrysops, Philemon corniculatus and Meliornis (= Phylidonyris) novaehollandiae, which they collected by shooting at Milson Island in the Hawkesbury River and in Sydney. In 1910 they described Trypanosoma anellobiae from Anellabia (= Anthochaera) chrsoptera which was collected near Brisbane, Queensland. They obtained further samples of a number of species of meliphagids from the environs of Eidsvold, Queensland and in 1911 they identified Trypanosoma anellobiae from Myzomela san- guiniolenta, Piilotis (= Meliphaga) fusca, En- tomyzon cyanotis and Myzantha garrula (now Manorina melanocephala). In 1911, they described and illustrated what they called the “intracorpuscular” or “Leucocytezoon” stage of Trypanosoma anellobiae and presenied a discus- sion of how this stage was part of the life cycle of the trypanosome. However, the illustrations clearly indicate a species of Leucocytozoon Johnston (1912) referred to Lewcoryiezoon anel- lobiae in a table and inadvertently established a specific designation. Clearly, there is considerable confusion sur- rounding the identity of the avian haematozoa described by Cleland & Johnston from the Meliphagidae. In 1990-93, a large sample of Noisy Miners (Manorina melanocephala) from Laidley, Wivenhoe Dam and the Toohey Forest of Queensland were examined for blood parasites during the course of a study of their mating and social systems. These birds were infected with Haemoproteus, Leucocytozoon and Trypanosoma and thus gave the opportunity to re-describe these parasites using modem criteria and evaluate their systematic position. MATERIALS AND METHODS Blood samples from 173 Noisy Miners [Manorina melanocephala) in three locations in Queensland (Laidley, Wivenhoe Dam and Toohey forest) were collected via jugular venipuncture during the period 1990-1993. A blood smear from each sample was prepared in the field, air-dned and fixed in ethanol within 12 hours. Smears were then sent to the International Reference Centre for Avian Haematozoa (IRCAH) where they were re-fixed in 100% methanol and stained with Giemsa's stain ata pH 7.2 and examined for parasites. Forty-five birds were infected with species of Haemoproreus, Leucoeyiozaon and Trypanosoma and an addi- tional seven birds were infected with Pias- modium vaughani, which constitutes the first Australian record of this parasite. The blood parasites were drawn with the aid of a camera lucida and the morphometric parameters were determined with the aid of a Zeiss MOP-3 Digital Analyser. The parameters for the haemoproteids were measured by the protocols established by Bennett & Campbell (1972) as modified by Forrester etal. (1977). The parameters used for the leucocytozoids were es- tablished by Bennett et al. (1991). The mor- phomeiric parameters and derived indices for the trypanosomes follow the generally accepted measurements for this group (Woo & Bartlett, 1932). All measurements are presented as the 14 MEMOIRS OF THE OUEENSLAND MUSEUM mean with the standard deviation in parentheses. All photomicrographs were taken with a Zeiss Photoscope II. In the re-descrniptions of the haemoproteids and leucocytozoids, the measure- ments for the males are not presented in the interests of brevity, However, if the dimensions are markedly different from those of the macro- gametocyte, these arc mentioned in text. Through the courtesy of Dr Penny Berents of the Australian Museum in Sydney, the original material used by Cleland & Johnston was made available for study. TAXONOMIC REVIEW Haemoprotens ptilotis (Cleland & Johnston, 1909) emend, Coatney, 1936 TYPE Host Prilotis chrysops (Latham), now Meliphaga clirysops (Latham). ‘TYPE LOCALITY Milson Island, Hawkesbury River, New South Wales, Uninfected erythrocytes. N = 25. Erythrocyte 12.1 (0.8),1m in length, 6.1 (0.6) um in width and 58.1 (7,4)pm7 in area; erythrocyte nucleus 5.2 (0.5)im in length, 1.9 (0.2)am in width and 7.7 (1.2)um? in arca. Immature gametocytes. Youngest forms seen Were usually lateral to Ihe erythrocyte nucleus, but sometimes in a polar position; margin entire. Macrogametocyte (Fig. 1A). N — 35. Infected erythrocyte 12,3 (1.0)lm in length (6% hyper- trophy), 7.1 (0.6)j1m in width (16% h hy) and 72.0 (8.6)um* in area (24% hypertrophy); infected erythrocyte nucleus 5,1 (0.6)um in length (2% atrophy), 2.1 (0.3)pm in width (10% hypertrophy) and 8.5 (1.5)um” in area (10% hy- pertrophy). Parasite halteridial, entire in outline and occupying 58% of the area of the host cell- parasite complex. Parasite somewhat sausage- shaped, 12.1 (1.2)um in length, 3.5 (0.4)um in width at the middle of the parasite and 41.8 (6.4)lLm2 in area; parasite nucleus ovoid to round in outline, median in position, 2.2 (0.1)um in length, 1.4 (0. 1)p1m in width and 2,4 (0.7)um” in area; pigment granules average 10 (1.7) granules per parasile, small and scattered randomly throughout parasite cyteplasm; erythrocyie nucleus only slightly displaced laterally, NDR (Nuclear Displacement Ratio) = 0.6 (0.2) where the NDR represents the degree of lateral displace- ment of the cell through the action of the parasite (Bennett et al., 1990:194), vacuoles not prominent and volutin granules not seen. Microgametocyte (Fig. 1B). N = 10. Microgarmeto- cytes closely similar to macrogametocytes in all dimensions except for the larger parasite nucleus typical of the microgametocytes of all the apicomplexan parasites; parasite nucleus central with volutin granules concentrated at the poles and not occupying the area occupied by the large parasite nucleus. BASIS OF REDESCRIPTION Blood films 115027 and 124757 from Noisy Miners Manorina melanocephala collected by Poldmaa in Queensland, Australia from Toohey Forest on 7 Oc- tober 1990 and Wivenhoe Dam on 6 September 1992 respectively, COMMENTS Haemoprateus ptilotis is a small, halteridial haemoproteid that occupies less than 60% of the host cell-parasite complex. It has few and rather small pigment granules. It is considered to be a distinct species on the basis of its occurrence in the family Meliphagidae, following the ussump- tion that hacmoprotcids arc host family or sub- family specific (Bennett & Peirce, 1988), Haemoproteus ptilonis fits the description presented by Cleland & Johnston (1909) as far as can be followed. The illustrations presented by these authors are essentially the same as those figured including the few, small pigment granules per parasite. The measurements presented by Cleland & Johnston lie well within the range of those presented above, Unfortunately, the hapan- totype (cotype of Cleland & Johnston, 1909) slide of H. ptilatis had degraded beyond use. Not only had the stain faded but the erythrocytes themsel- ves were disintegrating and no parasites could be distinguished at any place on the blood film. In essence, the existing hapantotype slide is worth- less and only the original description and illustra- tions of the species remains to define it. Cleland & Johnston (1909), using the generic designation of Halteridiwm, subsequently emended by Coatney in 1936 to Haemoproteus, also described Haemoproteus philemen trom Philemon corniculatus and their illustrations and dimensions of the parasite in this bird are essen- tially the same as the description for H. ptilotis, including the small parasite with few pigment granules. They mention that although a few of the parasites of H. philemon were larger than those of H. prilotis, *. . little difference could be detected and the identity or otherwise of the two HAEMATOZOA OF AUSTRALIAN HONEYEATERS I5 must await the inyestigation of other stages in their life histories” {Cleland & Johnston, 1909:84). They also described Haemoprojeus meliornis from Meliornis (= Phylidonyris) novaehollandiae, This bird had an intense infec- tion with many erythrocytes having multiple in- vasion of parasites, Their illustrations of this parasite indicate many of the parasiles were im- mature, but the mature forms are unquestionably the same as H. prilotis and the dimensions of the cells are within the range of those cited in the redescription above, Regrettably, the hapan- totype slide of H. meliornis ts in equally poor shape as that of H, priloris and nothing could be seen on the smear except disintegrating cells that had lost their stain and the slide is cssentially worthless. The blood film of H. philemon was fortunate to have been cover-slipped with Canada balsam and the cells on the smear were intact although badly faded. As described originally, this smear contained cells with up tò four imma- ture parasites. However, only two mature parasites were seen and these, as far as could be determined given the lack of staining, appeared to be similar to their illustrations in 1909 and to those used in the re-description above, Although the hapantotype slides of these parasites are use- less for taxonomic purposes, on the basis of the original descriptions and line drawings presented by Cleland & Johnston (1909), all three species described from the meliphagids by Cleland & Johnston are the same species. By page priority (ICZN, Section 69B (11)), therefore, Haemoproteus ptilotis is the name of the haemoproteid in the Meliphagidae and Haemoproteus philemon and Haemoproteus meliornis fall as synonyms. Leucocytazoon anellobiae (Cleland & Johnston, 1911) emend. Johnston, 1912 TYPE HOST The little wattle bird, Anrhochaera chrysopïera (Latham). TYPE LOCALITY Brisbane, Queensland, Australia. Macrogametocyte (Fig. 1C). N — 51. Parasite with round morphs only. Parasite broadly ovoid 1o round, with a maximum diameter of 11,3 (l.m, minimum diameter of 9.5 (0.9)um, a periphery of 33.1 (3.5)im and an area of 85,4 (15.6)nm”, occupying 80% of the area of the host cell-parasite complex; parasite nucleus round to broadly ovoid and sometimes elliptical, 4.0 (0.5)um in length, 2.8 (0.5)jlm in width and 8.2 (Pym? in area, without a marked karyosome, occupying 9.7% of the area of the parasite; vacuoles small and not prominent; volutin granules not seen; nucleus of host cell-parasite complex usually as a nbbon but sometimes as a cap, 23.) (8,7}um° in area and covering 14,9 (4.8)um of the periphery of the parasite (44%) and occupying 21% of the area of the host cell- parasite complex; host cell-parasite complex 108.5 (22,6)um” jn area. Microganwiocy (Fig. ID). N = 14. Microgameto- cyte similar to thc macrogametocyte in most respects except for the usual larger nucleus and pale staining that occurs in the apicomplexan parasites. The microgametocyte on average is 5-10% larger in most dimensions than the macro. gametocyte and the host cell-parasite nucleus is larger and covers a greater amount of the penphery of the parasite (66% compared to 44% for the macrogametocyte), BASIS OF DESCRIPTION HAPANTOTYPE: Blood film no. 115021 from Manorina melanocephalu collected by Poldmaa at Toohey Forest, Queensland on 7 October 1990, PARAHAPANTOTYPES: Blood film no. 8872 from che Noisy Miner Manorina melanocephala collected by Bennett at Kenmore, Queensland on 7 September 1968. blood filmno, 124710 from the same species collected by Poldmaa ac Wivenhoe Dam, Queensland on 15July | 992. COMMENTS This is a small round leucocytozoid, one of the smallest of the species described. It is considered to be a distinct species on the basis of the presumed familial/subfamilial specificity demonstrated for a number of species of Leucocytozeon (Bennett et al., 1991). Bennett & de Swardt (1989) believed that Leucocytozoen anellobiae also occurred in the South African Gurney's sugarbird (Promerops gurneyi) which was at that time classified as a meliphagid. How- ever, this genus is now believed to have little or norelationship with the Australian Meliphagidae and has been placed in its own family, the Promeropidae (although some authorities con- sider them to be in the subfamily Promeropinae of the Meliphagidae), On comparison of the sugar bird material with that from the Australian noisy miner, it was evident that the South African Species Was much larger and that Bennett and de Swardt (1989) were in error. Therefore, the South African leucocytozoid was described as 16 MEMOIRS OF THE QUEENSLAND MUSEUM Leucocytozoon deswardti by Bennett et al. (1992). Cleland & Johnston (1910) WE described Trypanosoma anellobiae from Anellobia (=Anthochaera) chrysoptera and in 1911, elaborated on this parasite with remarks based on finding this trypanosome in several other species of the Meliphagidae. They were also convinced that the “Leucocytozoon” stage was the intracorpuscular stage of the trypanosome life cycle, a commonly held view at the time. This view may have been prompted by observa- tion of the highly fusiform (almost trypanosome-like) appearance of Leucocytozoon gemanni of owls, birds which are frequently concurrently infected with both parasites. Johnston (1912) referred in a table to Leucacytozoon anellobiae. The footnote to this specific name reads “The name Leucocytozoon anellobiae is here given to a blood parasite found by Dr. Cleland and myself in several species of birds, We believe it to be a phase in the life history of Trypanosoma anellobiae (Cleland & Johnston). [have used the above name as possess- ing specific value, Should our opinion as to the specific identity of the two forms be correct, then the name L. anellobiae becomes a synonym, or, to be more exact, it refers toa particular phase of T. anellobiae.” Thus Johnston inadvertently created Leucocytozoon anellobiae as a valid species. Whether inadvertent or not, the name stands as the valid designation of the leucocytozoid of the Australian meliphagids and is herein so recognised. Cleland & Johnston (1910) did not indicate the disposition of the material used to define the “Leucocytozoon” stage of Trypanosoma anellobiae. However, when Johnston (1912) created Leucocytozoon anellobiae, he was in the Department nf Biology of the University of Queensland. There is no trace of this material at the Oueensland Museum or on record at other Australian institutions as far as is known (Lester Cannon, pers. comm.). In as much as that no “type” material was designated for L. anellobiae, we are designating hapantotype and parahapantotype slides from Manorina melanocephala, one of the hosts from which the =. = ven mi icrogametocyte; C, Leucocytozoon anellobiae, anellobiae, microgameiocyte; E, Trypanosoma anellobiae. D Ne — -— FIG. 1. A, Haemoproteus prilotis, two macrogameiocytes; B, Haemoproteus ptilotis, cyte; D, Leucocytozeon “Leucocytozoon” stage was described and are birds from the type locality. Trypanosoma anellobiae Cleland & Johnston, 1910. TYPE HOST The little wattle bird, Anthochaera chrysoptera (Latham). TYPE LOCALITY Brisbane, Queensland, Australia. Trypomastigote (Fig, 1E). N = 6, Trypomas- tigote small and slender, averaging 25.6 (2.3)um in length and 5.7 (0.9)um in width at the position of the nucleus. Kinetoplast 2.7 (0.8)um from posterior end and 9.7 (0.6)1m from the nucleus. Nucleus 12.3 (0.8)wm from the anterior end which has a long free flagellum averaging 10.8um (only two free flagellae measured), Trypomastigote 82,0 (12. 7)um? in area, nucleus 15.6 (2.5)um? in area, the nucleus representing 19% of the area of the parasite. The distance from the posterior end to the kinetoplast represents 10% of the length of the trypanosome, while the distance of the centre of the nucleus from the posterior end is 48% of the length of the parasite, the nucleus approximately at the mid-point of the irypomastigote. The width of the trypanosome at the centre of the nucleus is 22% of the length of the organism. HAEMATOZOA OF AUSTRALIAN HONEYEATERS 17 BASIS OF DESCRIPTION Blood film No. 115082 from Manorina meélanocepiala collected by Poldmaa at Toohey Forest, Queensland on 18 October 1990. COMMENTS Cleland & Johnston (1910) described Trypanosoma anellobiae from _ Anellobia chrysoptera (now Anthochaera chrysapteraì from a bird shot at Brisbane, Queensland. The infection was light and their description indicates that the trypomastigote was about 0,035mm in length with a maximum breadth of 0.002mm, The “kinetonucleus” was situated 0,003mm from the posterior end. They could not see the nucleus of the organism and they did not detect a free flagel- lum. They concluded that the undulating membrane was very narrow and believed it to be short. Their illustrations (Plate xxxiv, figs 6, 11) are clearly those of the trypanosome illustrated in Fig. 1E of this study. The measurements they presented are closely similar to those presented herein, especially with respect to the position of the kinetoplast. It is clear that the trypanosome in the Noisy Miner is the same as that described by Cleland & Johnston. The hapantotype slide of Trypanosoma anellobiae was examined and found to be in the same condition as described for the hapantotype slides of Haemoproteus priloris and H. meliornis. Only a few of the crythrocytes were intact and the stain had faded to the extent that the blood smear was a monocolour, The blood smear was reported to also contain two species of microfilanae, However, no trace of these parasites could he found. The blood smear i$ unacceptable as the basis for definition of a taxon. On the basis of the original description, Trypanosoma anellobiae is a small trypanosome that Jacks the striated appearance and larger size of the T. avium group; it is also easily separated from T. paddae, T. corvi and T. hannai on the basis of its much smaller size and distinctive derived ratios. It ts smaller than T. beuffardi but shares the same slender appearance of this trypanosome and also differs in that the kinetoplast is much closer to the posterior end. The length of T. anellobiae is within the same range as T. everen, and the placement of the kinetoplast is similar in both of these species. However, T, everest is a broader trypanosome (which gives the organism a “stumpy” ap- pearance) with a nucleus that occupies about 26% of the area of the trypomastigote and cannot be confused with T. anellobiae. On the other hand, T. anellobiae is remarkably similar to T. on- tarioensis Woo and Bartlett, 1982. Both trypanosomes are small, slender and with the kinetoplast located close to the postenor end; measurements cited herein lie within the ranges guoted by Woo & Bartlett for their species. Both species have a long free flagellum that is about one-half the body length and the derived ratios are essentially the same, Tt would be essentially impossible to separate the 1wo species on the basis of their morphometrics and appearance. Trypanosoma ontarioensis was originally described from a corvid in Ontario, Canada but its appearance is similar to the numerous small trypanosomes that have been inadequately described from South American birds and many of these South American trypanosomes can un- doubtedly be assigned to this species. In addition, T. ontarioensis has been recorded from Sweden and appears to have a broad distribution. Trypanosoma onfarioensis is a readily cultured trypanosome, doing particularly well on diphasic blood-agar medium, producing infective cultured forms in two wecks. While the isolation of Australia would suggest that T. anellobiag is a distinct species and no attempt will be made to synonymize the two species al this time, the close similarity of the two species toeach other requires expenmental confirmation of their identity. This study has also highlighted what will be- come a major problem for museums and repositories of hapantotype material of the Protozoa. The hapantotype material of the Cleland & Johnston species were 84-85 years of age and had deteriorated to the extent they were of no value as the basis for the definition of the taxons they were supposed to represent. It is almost 90 years since the first edition of the International Code for Zoological Nomenclature was published and the practice of establishing type material became mandatory although type material had frequently been designated decades before. Stained protozoal and similar material does age with time and as indicated in this study, in 85 years had deteriorated beyond use, While the use of coverslips and mounting media does aid in the preservation of the cells, the problem of stain fading over a long period still occurs, The preservation of hapantotype material will become a serious problem that will have to be addressed by curators of this type of material and should he addressed as a priority by the Intemational Com- mission for Zoological Nomenclature. 18 MEMOIRS OF THE QUEENSLAND MUSEUM ACKNOWLEDGEMENTS The financial support of the Natural Sciences and Engineering Research Council of Canada to both the first and last authors is gratefully ac- knowledged. LITERATURE CITED BENNETT, G, F. & CAMPBELL, A. G. 1972. Avian Haemoproteidae, I. Description of Haemoproteus fallisi n, sp. and a review of the haemoproteids of the family Turdidae. Canadian Journal of Zoology 50: 1269-1275. | BENNETT, G. F., EARLE, R. A. & PEIRCE, M. A. 1992. The leucocytozoidae of South African birds: the Passeriforms. Onderstepoort Journal of Veterinary Research 59: 235-247. BENNETT, G. F., EARLE, R. A., PEIRCE, M, A., HUCHZERMEYER, F. W. & SQUIRES-PAR- SONS, D. 1991, Avian Leucocytozoidae: the leucocytozoids of the Phasianidae sensu lato. Journal of Natural History 25: 1407-1428. BENNETT, G.F. & PEIRCE, M. A. 1988. Morphologi- cal form in the avian Haemoproteidae and an annotated checklist of the genus Haemoproteus Kruse, 1890. Journal of Natural History 22: 1683- 1696. BENNETT, G.F., PEIRCE, M. A. & EARLE, R. A. 1990, The haemoproteids of the shrikes of the the avian family Laniidae (Passeriformes). South African Journal of Zoology 25: 194-198. BENNETT, G.F. & DE SWARDT, D. H. 1989, First African record of Leucocytozoon anellobiae (Apicomplexa: Leucocytozoidae) in Gurney's sugarbird, Promerops gurneyi. Ostrich 60: 171. CLELAND, J. B. & JOHNSTON, T. H. 1909. Descrip- tions of new haemoprotozoa from birds in New South Wales, with a note on the resemblance between the spermatazoa of certain honeyeaters (Fam. Meliphagidac) and spirochaete- trypanosomes, Journal of the Proceedings of the Royal Society of New South Wales 43: 75-96. CLELAND, J. B. & JOHNSTON, T. H. 1910. The haematozoa of Australian birds. - No. I. Transac- tions of the Royal Society of Australia 34: 100- 114. CLELAND, J. B & JOHNSTON, T. H. 1911. The haematozoa of Australian birds, No. II. Journal of the Proceedings of the Royal Society of New South Wales 45: 415-444, COATNEY, G. R. 1936. A check-list and host-index of the genus Haemoproteus. Journal of Parasitology 22: 88-105, FORRESTER, D. J., GREINER, E. C., BENNETT, G, F & KIGA YE, M. M. 1977. Avian Haemoproteidae, 7. A review of the haemoproteids of the family Ciconiidae (storks) and descriptions of Haemoproteus brodkorbi sp. nov. and H. peircei sp. noy: Canadian Journal of Zoology 55: 1268- 1274. INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE. 1985, (International Trust for Zoological Nomenclature in association with British Museum (Natural History) London). JOHNSTON, T. H. 1912. Internal parasites recorded from Australian birds. Emu 12: 105-112. WOO, P. T. K. & BARTLETT, C. M. 1982. Trypanosoma ontarioensis n. sp. and T. paddae from Corvus brachyrhynchos in Ontario, Canada with notes on the biology of T. ontarioensis n. sp. Canadian Journal of Zoology 60: 2107-2115. A REVIEW OF HETEROPORODRILUS FROM SOUTH-EAST QUEENSLAND (ANNELIDA:OLIGOCHAETA) ROB J.BLAKEMORE Blakemore, R.J. 1904 12 01: A review of Heteroporodrilus from south-east Queensland CRY aaa Memoirs of the Queensland Museum 37(1):19-39. Brisbane. ISSN 79-8835. The endemic earthworm Heteroparvdrilus Jamieson, 1970, is revised to acconimodate four new species, plus new combinations mainly from Plurellus s. lat., that raise the generic total from ten to nineteen species. Previous emendments to these two genera, both of which display nephropore alternation, had reduced primary differentiation to absence or presence of stalks on thc calciferous glands. In the current revision, however, the form of prostatic glands is considered more definitive. Possession of tubular prostates qualifies Plurellus, now restricted to three known species, while in Heteroporadrilus there is development trom this plesiomor- phic state 1o the more derived racemose or tubuloracemose form of prostate. Herero- porodrilus has an extensive distribution in the eastern subregion of Australia yet it appears especially diverse in south-east Queensland, compared to Phwrellus s. strict. that is maint restricted Io central coastal New South Wales.Redescription of the type species of bot genera, H. Iryoni and P. hereroporus, confirms the basis of this revision, The distributions and ecology of these and other described species are considered and new keys ta genera are provided, [_] Heteroporodrilus, Plutellus, calciferous glands, taxonomy, earthworm ecology. R.J. Blakemore, University of Queensland, St Lucia, Queensland 4072, Australiu; cartent address; c/o CSIRO Division of Entomology, PO Box 1700, Canberra, Australian Capital Territory 2601, Australia; 13 March 1993. The current study resulted from part of a project investigating the distribution and ecology of earthworms in managed soils and their effects on soil fertility in south-east Queensland, Surveys located 44 exotic species that were often abun- dant and widely distributed, as well as 41 native taxa thal were generally more restricted in their distributions. This paper concerns the indigenous Heteroporodrilus Jamieson, 1970 that has not been extensively revised since its designation. This genus is frequently encountered in the wider Brisbane region. Fletcher (1889) was the first to recognise a well-marked group of species (which he never- theless referred to Cryptodrilus, Fletcher, 1886) sharing, amongst other characters, nephropore alternation, It was left to Jamieson (1970) to formally define Heteroporodrilus for ten (plus one dubious) 'heteropore' species, many of which were by then assignable to Woodwurdiella Stephenson, 1925. Inthe same paper, a restriction of the heterogeneous assemblage attributed to Plutellus Perrier, 1873 was suggested (as en- visaged by Gates, 1961), as only P. manifestus (Fletcher, 1889) from central New South Wales appeared to be congenenc with P. heferoporus, the type-species. Re-description of type material confirmed the basis for a restriction of Plutellus lo species with tubular prostates, bul this was deferred (Jamieson, 1971a,b). A numerical analysis by Wallace (1972) confirmed these groupings even though prostate morphology was withheld as an attribute. In order to accommodate two new heteropore species from Queensland that bore racemose prostates it was necessary for Jamieson & Nash (1976) to emend the generic definition of Plurel- lus to include both tubular and racemose prostates thus reducing the distinction from Heteroporodrilus. Such ‘infragenenc variation’ was used by these latter authors to ‘strongly refute’ lhe use of racemose prostates to distin- guish a family Megascolecidae from a family Acanthodrilidae with tubular prostates (and holonephridia) as advocated by Gates (1959) anu rejected by Jamieson (1971), However, a primi- tive 'plutelloid” from Lord Howe Island (Jamieson, 1977) secmed to have more in com- mon with its congeners in New South Wales. The definition of Plutellus was further modified by Dyne (1981) to receive three new species from the periphery of the Morion Bay Region, bringing the peneric total to eight, and the ‘tenuous’ dis- tinction from Hetereporodrilus was reduced principally to the presence or absence of stalks to the calciferous glands. 20 MEMOIRS OF THE QUEENSLAND MUSEUM The wide distribution of Heteroporodrilus from Queensland to Victoria corresponds in part with various riverine systems (Jamieson, 1970). It appears especially aggregated, or rather the reports are more frequent, in the Brisbane region of south-east Queensland. In contrast, Plutellus s. lat., Was recorded further south in coastal New South Wales and just to the south and north of Brisbane. Dyne (1981, 1984) found this distribu- tion pattern enigmatic because no appreciable biogeographic barners intervened, He also iñ- timated that should intermediate forms with short-stalked calciferous glands be found, it would be doubtful that the (wo genera could remain discrete. The present study contends that just such inter- mediate forms already existed in Plurellus s. lat., moreover 'short-stalked' calciferous glands had also been confirmed in Heteraporodrilus, but thc consequences are here interpreted in a different manner. The apparent discontinuity of the dis- tributions is largely resolved by revision of the definitions of both genera, based on new material and re-examination of previous accounts, result- ing in an amalgamation of the ‘northern’ heteropores while maintaining the ‘central’ plutelloids as a discrele zoogeographic and phyletic entity. CONVENTIONS Onginal illustrations, drawn using a Wild M5 microscope with camera lucida, have shaded clitella. All scale bars represent I mm. Abbreviations used are: i ii etc., segments num- bered from the peristomium, ii/ iii/ etc., on ii, on iii etc.; 1/2 2/3 etc. intersegmental furrows, i.e., between i and ii etc.; a b etc., individual setae from the ventralmost on each side; A B ctc.; longitudinal series between setae a setae b ete. gmn. genilal marking; n.p. nephropore; NSW, New South Wales, Old., Oueensland, OMG, Oueensland Museum registration number; $p.p., spermathecal pore; ; U. (convention from Ger- man) circumference. Setal ratios, of debatable systematic importance, follow the formula (xii/aa:ab;cd'dd;u). SYSTEMATICS Heteroporadrilus is emended to accommodate, the new species, and to receive several species that were previously placed in Plutellus (sensu Jumieson & Nash, 1976; Dyne, 1981). The revision proceeds in three stages. First, at least two species (H. jamïesoni sp. nov. and H. ox- leyensis) belonging to the established Hetereporodrilus, are found to have short-stalked calciferous glands thereby expanding the generic definition to species with sessile or short-stalked calciferous glands. Hence the distinction from Heteraporodrilus of the only “Plutellus' species reported from Queensland, P. incommodus, P. raveni and P. notatus, which are all described with similar short-stalks, is removed. Secondly, expanding the definition to include Dyne's (1981) ‘intermediate’ species with short-stalked glands and racemose ortubuloracemose prostates absorbs the two species from northem New South Wales, P. minyoni and P. clarkei. Thirdly, Plutel- lus is restricted to only those species with tubular prostates and long-stalked calciferous glands, This division is especially strong as no genetic correlation between prostate form and calciferous nexus is implied. Tubular prostates are generally considered the plesiomorphic condition, thus Heteroporodrilus can be viewed as a derived, apomorphic sibling group of a more primitive Plutellus, The above split has the added con- venience of making the geographic distributions disjunct with Plutellus confined to the coastal region of central New South Wales (two species) and Lord Howe Island (one species) and the remainder in Heteroporodrilus, chiefly in south- cast Queensland and northern New South Wales. 'The guestion of the phyletic relationships of rhe Lae Howe Island plutelloids, is not ventured Heteroporodrilus Jamieson, 1970 emend. TYPE SPECIES Heteroporodrilus tryoni (Fletcher, 1890) from Milton, Brisbane. DIAGNOSIS Moderate to large size terrestrial worms (52- 580mm long) generally with less than 200 seg- ments (range 90-387). Sometimes dorsum canaliculate and with or without brown-grey pig- mentation. Prostomium variable from pro- epilobous to tanylobous, often grooved. Seiae & per segment (only occasionally a and/or b retained on xviii), Dorsal pore from 5/6 or beyond, Male pores and pores of one pair of racemose to tubuloracemose prostates in xviii, Spermathecal pores 2-5 pairs, the last in 8/9 (H. notatus and H. clarkei near mid-ix). Nephropores conspicuous at the anterior border of their seg- ments in D lines in JIi-iv or v; in C lines, or REVIEW OF HETEROPORODRILUS 2 alternating between D and C (or mid-BC) for a few segments; then from x where alternation between B and D lines commences for the remainder of the body (may be asymmetrical). Gizzard in v. Calciferous glands, 3-5 ventro- lateral pairs sessile or on short-stalks (i.e. about as long as broad) on the oesophagus, the last pair always in xiii. Spermathecae have one or more, discrete or composite, diverticula. Holonephnc with (adiverticulate but sometimes bilobed) ter- minal bladders. Holandric, testis and funnels free or in unpaired (pericardiac) testis sacs, DISTRIBUTION Murray-Darling River basins in NSW and South Australia; the Wimmera River, Victoria; Tweed River basin in northem NSW; river catch- ments in south-east Old. SPECIES OF HETEROPORODRILUS The numbers and conditions of specimens in- spected in previous definitions (in brackets) indi- cates the stability of those descriptions, Only species with an asterisk are (re)described in this paper. 1. *H. bongeen sp. nov. from Bongeen, Old. 2. H. canaliculatus (Fletcher, 1889) from Lachlan River, Forbes, NSW. Crypiodrilus canaliculanis Fletcher, 1889: 1534- 1536 (twelve specimens, poorly preserved); Fletcher, 1890: 996. Plutellus canaliculatus; Michaelsen, 1900. Heteroporodrilus canaliculatus; Jamieson, 1970: 111-112 (types no longer traceable). 3. H. clarkei (Dyne, 1981) comb. nov. from Whian Whian State Forest. northern NSW. Plutellusclarkei Dyne, 1981:97, figs 1a; 2c, e; table 1 (three specimens) 4. H, cooraniensis (Spencer, 1900) from Cooran, d Id. Cryptodrilus cooraniensis Spencer, 1900: 42-43, figs 34, 35, 36 (spirit specimens); Sweet, 1900; 114; Jensz & Smith, 1969: 86 (only one guestionable syntype remains in reasonable condition). Woodwardia cooraniensis; Michaelsen, 1907: 162; Bage, 1910; 234-236, figs 18-21. Heteroperodrilus cooraniensis; Jamieson: 1970, 112-113 (material not available). 5. *H. dioecius (Stephenson, 1933) from Toowoomba, Petrie, Brookfield and Samford (new record), all in Old. Woodwardiella dioecia Stephenson. 1933; 910- 912, figs 9; 10 (four clitellate specimens). Heteroporodrilus dioecius: Jamieson, 1970: 113- 114, figs 5d-f; 9b: 10b. c (four clitellate and onc aclitellate specimens). *H. doubei sp, nov. from Lismore, NSW. H. mcommodus (Jamieson & Nash, 1976) comb. nov. from Eudlo Creek and Forest Glen, Old. Plutellus incommodus Jamieson & Nash. 1976: 47- 49, figs 1: a, b; 2a-g: table 1 (14 specimens). . *H. jamiesoni sp. nov, from Mt Glorious, Old. Heteroporodrilus ashworthi (Stephenson, 1933) syn. nov. (part, Mt Glorious specimen non Stephen: son, 1933 specimen) sensu Jamieson, 1970: 109- 111, figs 2; 3a; Sa,b; 9a; 10a (one clitellate specimen). Note: H. ashworthi sensu Jamieson (1970) is not Use same as W. ashworthi Stephenson, 1933, but repre- sents a new species. Since W. ashworthi is a junior synonym of H. oxleyensis (Fletcher, 1889) the namc ashworthi is unavailable and H. ashworthi sensu Jamieson (1970) is described herein as H. jamiesoni Sp. nov. 9. H. lamingtonensis (Jamieson, 1970) comb, nov. from O’Reilly’s, Lamington Nat. Park, id. Pw lamingtonensis Jamieson, 1970; 115-117, figs 5a; 9c; 10d (a single specimen). 10. H. mediterreus (Fletcher, 1887b) from banks of the Darling River between Bourke and Brewarrina NSW, Gragin and Warialda, NSW, Cryprodrilus mediterreus Pletcher, 1887b; 614-615 {len specimens - Jost). Heteroporodrilus mediterreus Jamieson, 1970: 117-120, figs 4a,b,e,f,g; 9d; 10e (six poùr specimens, only two clitellate). Note: The records by Jamieson (1970: 119) for this species in Victoria are incorrect. 11. H. minyoni (Dyne, 1981) comb. nov. from Whian Whian State Forest, northern NSW. Plusellus minyoni Dyne, 1981; 100-102, figs 1b; 2a, b, d; table 1 (a single intact specimen and several anterior amputees). 12. H. notatus (Dyne, 1981) comb. nov. from Cooroy, Old. Plutellus notatus Dyne, 1981: 102-104, fig 3 a-c; table 1 (three specimens). 13. *H. oxleyensis (Fletcher, 1889) from Oxley; Brookfield (new record), Samford (new record) and Mt Cotton (new record), all near Brisbane, Old.; ?Marrickville, NSW. Crypiodrilus Oxleyensis Fletcher, 1889: 1537- 1538, (onc complete but aclitellate specimen and two other “more of less incomplete specimens”). eu EN oo 22 MEMOIRS OF THE QUEENSLAND MUSEUM Plutellus flescheri (party; Michaelsen, 1900; 173 ( excluding Cryptodrilus fletcheri Beddard, 1887). Woodwardia oxleyensis; Michaelsen, 1907: 162. Heteroporodrilus oxleyensis Jamieson, 1970; 120- 122, figs 3b-e; 5g; 9e; 10l,g (two clitellate specimens). Woodwardiella ashworthi Stephenson, 1933: 912- 914. figs 11-13, syn. nov. (a single clitellate specimen). Heieroporodrilus ashworthi; Jamieson, 1970: 109- 111 (part excluding Mt Glorious specimen}. 14. H. raveni Jamieson and Nash, 1976) comb. nov. from Forest Glen, Qld. Plutellus raveni Jamieson & Nash, 1976: 50-52, figs 2h-j; 3, table 1 (six specimens). 15. H. shephardi shephardi (Spencer, 1900) from Horsham, (and Dimboola?) Victoria (Wimmera River). Cryptodrilus shephardi Spencer, 1900; 40-41, figs 28, 29, 30, (“spirit specimens"), Jensz & Smith, 1969: 91 (three lectotypes remain), Woodwardia shephardi, Michaelsen, 1907: 162, Heteronorodrilus shephardi shephardi Jamieson, 1970; 122-124, figs 4c,d; 9f; 10i (a paralectotype specimen), 16. H. shephardi armatus Jamieson, 1974 from Penola and Naracoorte, Victoria (about half a dozen specimens), Heteroperadrilus shepharadi armatis Janicson, 1874; 85-87, figs 2b; 10b; 11a; 13, table 3 (four or five specimens). 17, H. sloanei (Fletcher, 1889) from Connabarabran, NSW. Crypradrilus Sloanei Fletcher, 1889: 1526-1537, (four aclitellate specimens and one doubtful juvenile). Pluteilus sloanei; Michaelsen, 1900. Heteroporodrilus sloanei; Jamieson, 1970; 124, (three desiccated specimens). Note: Jamicson (1970) considered synonymy with H. canaliculatus and H. lamingtonensis dependent on ayailability of new material, 18. *H. thompsoni sp, nov. from Brookfield, Old. 19. *H. iryoni (Fletcher, 1890) from Milton, Sherwood, Brookfield and Toowong sub- urbs of Brisbane; Binna Burra, Lamington National Park; Mr Glorious; Mt Mee and Nambour, all in Old. Cryptodrilus Tryoni Fletcher, 1890: 994-996, (ane poorly preserved specimen). Plutellus tryoni; Michaelsen, 1900; 171. Woodwardia (?) tryoni; Michaelsen, 1916: 62, Woodwariella tryoni; Boardman, 1932: 127-128 (a single, poorly preserved specimen). Heteroporodrilus ryeni, Jamieson, 1970: 125-129, figs 5b,c; 8c,d; 9g.h; 10j (nine clitellate specimens). Woodwardiella youngi Boardman, 1932: 128-130, fig 2 (two specimens). SPECIES INCERTAE SEDIS “Heteroporodrilus fletcheri (Beddard, 1887), provenance in Qld. unknown. Cryptodrilus fletcheri Beddard, 1887: 544-548, il- lustr, (two specimens, one mature - types lost), Platellus fleicheri; (part excluding H. oxleyensis) Michaelsen, 1900: 79, Heteroporodrilus ? fletcheri, Jamieson, 1970: 114- 115, Note: Based on inadequacies of the original account, Jamieson (1970) considered this species to be ‘nomen et species dubium’ but its affinity with H. oxleyensis was alluded to by Michaelsen, 1900. A large heteropore specimen, unfortunately too macerated for adequate characterisation, was col- lected by C.H. Thompson in 1984 from Mt Buderim near Maroochydore, Old. It had the ex- ternal appearance of H. shephardi and three pairs of spermathecae. Two unpublished new species from south-east Old. have been identified by Dyne (1984) and a furthertwo species from the Adelaide region are currently in preparation by the author. KEY TO HETEROPORODRILUS 1. Large size (250mm); five pairs of calciferous glands; three pairs of spermathecne.. . H. trywar Four pairs of calciferous glands ....,..., 2 Three pairs of calciferous glands ....... 10 2. Three pairs of spermathecae 2.2... 2.24 3 Two pairs of spermathecue , ,.,,.,. g 3. Nephropores alternate between c line and d line in 4/5-8/9 (from Queensland) . ...... 4 Nephropores alternate between d line and mid-bc in 4/5-8/9 (Victoria and South Australia), . , 7 4. Prostates confined to xviii, penial setae absent; genital markings usually in xx Prostates xvii-xix or xvi-Xxxii, penial setae present; no genital marking8$in XX... ... 5. Prosiates with straight duct; spermathecal diverticula variable, single or paired . . . H. incommodus Prostates ducts Jong and sinuous; spermathecae each with a single, simple diverticulum . . . HM. raveni 6. Dorsum canaliculate; spermathecae cach with two diverticula, occasionally single, but often compound; seminal vesicles in ix and xii H. canaliculans Body circular; spermathecae each with a single, simple (even rudimentary) diverticulum; seminal vesicles in xi and xii H. mediterreus REVIEW OF HETEROPORODRILUS 23 7, Genital markings as cye-like pits in 17/18 and 18/19; penial setae present in xviii H. shephardi armatus Genital markings segmental i in some or all of xvii, xviii and xix; penial setae absent yR hebauc H. shephardi shephardi 8, Spermathecal diverticula bifid or trifid or various- ly mulri-lobed or numerous, genial markings in KURS sy sn 2 We oc Gre wb snnt Aon Spermathecae with simple ( paired) diverticula H. bongeen sp. nov. 9. Size 210mm; prostomium epilobous furrowed; spermathecae with large and concertinaed ampul- lae and compound diverticula ssela obu á ar belydr de H, thompsoni sp. nov. Size <105mm; prostomium tanylobous; sper- mathecae, with simple or bifid diverticula, open- ing to midsegment .... .... H. notatus 10. Spermathecae, three or more pairs . ... . 1) Spermathecae, two pairs... ........ 14 11, Size generally 100mm; genital markings in xvii- BX Tae ae Se oe te ee Bee Gt . 12 Size generally <100mm; genital markings weak or absent (immature?) ........... 13 12. Spermathecae 5 (or 4) pairs; genital markings widely paired in xvii-xx, presetal in xvii ~+- H. jamiesoni sp.nov. Spermathecae 4 (or 3) pairs; postsetal genital markings elongate or closely paired in xvii-xx ie oo ee H. oxleyensis 13, Spermathecae 3 pairs each with two diverticula dn Td yno i, a EE nl H. sloanei Spermathecae 3 pairs {occasionally one of ihe six absent) each with a single diverticulum, prostates absent or present. . . . H. dineciuy 14. Spermathecal pores segmental, presetal in viii endi... ep etoile y H, clarkei Spermathecal pores intersegmental in 7/8 and O I —— 15 15. Size 300; prostomium epilobous; preclitellar genital marking a series of papillae within tumid ventral pads in xii, xvii-xxi. .. .. H. minyoni Size 100; prostomium epilobous; preclitellar genital markings in vii, viii and ix; spermatheal diverticula paired H. doubei sp. nov. Size <80; prostomium tanylobous; preclitellar genital markings widely paired or lacking . 16 16, Spermathecal diverticula paired . j H. ‘lamingionensis Spermathecal diverticula single . bw. ein th ta. H. cooraniensis Associations between species can be inferred from their neamess in the above key. Accounts of species not covered in the following descriptions may be sought with reference to the preceding species list. DESCRIPTIONS OF SPECTES Heteroporodrilus bongeen sp, nov. (Fig.1) MATERIAL EXAMINED HoLoryPE; OMG210136, opposite the local school house at Bongeen (27 /34'S; 151°27°E) on the Con- damine River plain west of Toowoomba, Old, R.J. Blakemore, 09 Apr 92, drawn and dissected, posterior amputee. PARATYPE: same collection data as holorype. OMG210137, dissected, posterior amputee. OTHER MATERIAL: same collection data as holotype, QMG210138, two anterior amputees (losing first 5-7 segments); also collected were four damaged sub adults, HABITAT Under bare soil in dark brown cracking clays, Waco clay gilgai complex (Beckmann & Thompson, 1960) on roadside next to cultivated fields. EXTERNAL CHARACTERS Length 250+mm, body circular in section without dorsal groove although one specimen had the posterior 6 segments furrowed, Width 5,5- 7.5mm, widest point about viii. Mass 3-45 per adult. Segments 193-200, first ten segments smooth followed by moderate secondary annulation in succeeding segments. Colour dark slate grey pigmented anterior (with paler ventral lumescences on x, xi and possibly xii), dorsum and caudal segments with faint red/yellow iridescence: mid-body moderate pig- mentation (soil visible in gut). Clitellum various- ly darker or lighter brown colour. Prostomium open epilobous or closed by fine groove but ap- pearing tanylobous due to longitudinal dorsal groove extensions which may almost reach as far as 1/2. First dorsal pore 10/117 (difficult to deter- mine due to deep furrows), Setae small, & per segment from ii, a and & obscure (modificd’?) on xviii, lateral setal couples ed widely spaced (mean of two ratios,1.7:1.0;:1.4:2:4:0,28), Nephropores in D line 1/2-3/4, 6/7, 8/9, 10/11 or 11/12 then alternating: in B line 5/6, 11/12 or 12/13 then allernating, possibly in C line in 4/5- 7/8. There is slight variation between specimens, but for most of the body length there is clear alternation between B and D lines. Clitellum '/sxiii-xvii annular, setae and nephropores retained. Male pores minute in xvii on small raised porophores in B line. Female pores paired in xiv in line of anterior annulus within a common 24 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 1. Heteroporodrilus bongeen sp. nov. A, ventral view of H specimen; B, prostale (with two adjacent nephridia); C, spermathecae in situ; D, reduced lateral view; E, prostomium. darker patch anterio-median to a setae. Sper- mathecal pores 7/8 and 8/9 in B line concealed in deep intersegmental furrows. Genital markings (x-xii ventrally tumid?), in xvii almost central between a setae, three small discs in raised tumid area; in xviii, especially in anterior annulus, four or five pairs of discs (one pair ventral in setal arc) in furrowed, generally tumid area that fills the ventral aspect; in xix similar markings, but discs tend to be smaller and more numerous. INTERNAL CHARACTERS Septa 4/5-11/12 getting progres- sively thicker, from 12/13 becom- ing weaker (10/11-12/13 thickest); pharyngeal ligatures (tendons) ex- tend beyond viii. Dorsal blood ves- sel single, continues on pharynx. Commissurals in vi-ix, larger hearts in x-xiii. Gizzard compact, spherical and muscular in v, Cal- ciferous glands four pairs, compact and ventral on oesophagus in x, xi, xii, and xiii; those in x smaller, almost rudimentary, Oesophagus narrows in xiy (valve?) then widens in xv as intestine proper; typhlosole not found. Holonephridia, a pair in each seg- ment, from at least v, with large deflated and flattened, spherical terminal bladders. Nephridia closely associated with anterior septa (pre-septal funnels), Male or- gans holandric: iridescent funnels of testes in x and xi; racemose seminal vesicles in ix and xii. Ovaries in xiii, ventrally paired sheets of mesentery may be ovaries but individual oocytes not seen, the egg funnels were seen on the posterior septum. Five or six smal- lish, white spheres were noted in one specimen loosely attached to the septum, possibly parasitic Monocystis sp? Prostates a pair of large tongue-like racemose glands extending from xviii-xix and over- lying a small, flaccid duct that soon enters the body wall, Sper- mathecae two pairs in viii and ix: waisted ampullae clearly demar- cated from shorter ducts, each bearing a pair of opposed digitiform, iridescent diverticula. One specimen had a single diver- ticulum on only one of its spermathecae. Gut contents fine grey clay soil and a few round grits coated in mucus (i.e. geophagous). TAXONOMIC REMARKS H. bongeen has 4 pairs of calciferous glands and 2 pairs of (bidiverticulate) spermathecae. It is perhaps closest to H. canaliculatus (Fletcher, 1889) recorded from the Lachlan River at Forbes, NSW. It differs from H. canaliculatus in several REVIEW OF HETEROPORODRILUS 25 features but especially in lacking dorsal canaliculation, in having one fewer pairs of sper- mathecae and in the numbers and position of the genital markings. On these variations, together with its geographic location, it is separated from other species and jt is named after its type loca- tron. ECOLOGICAL/AGRONOMIC OBSERVATIONS In the same samples, a native Diporachaeta sp. was also identified. Neither of these, por any other worms, were found nearby, including under an adjacent sorghum (Sorghum bicolor) crop, despite an hour of searching. Perhaps cultivation, which has severe effects on burrow systems and soil physical propertics, and acrial spraying of insecticides, are adverse to its survival, The worms were collected from opposite the local school and thereby in an area presumably less affected by application of biocides. A local farmer from Jondaryan (K. McIntyre, pers. comm.) reported that when the natural grasslands were first cultivated, masses of earthworms were exposed, Possibly the present population is residual from what must have been a very abun- dant fauna, The type locality was known to local fishermen in the 1950's as a good source of bait (C.H. Thompson, pers. comm,). The behaviour of these long worms was to escape rapidly along their burrows but when caught they were only moderately active, al- though they exuded copious watery fluid which dripped from their bodies. The specimens were dug from burrows up to lem wide to a depth greater than 30cm. Waco clay is a strongly struc- tured soil that has a thin, fine granular ‘self- mulching’ surface, but it is possible that the surface casting noted for this worm contributed to the granular aggregates observed. It is likely that their burrows extend down to the deep cal- careous brown clay subsoils found at depths of up to Im as during dry periods these deep subsoils would provide a moist retreat for earthworms. The agronomic potential of collected specimens was investigated in a glasshouse ex- periment, probably the first assessment of this type for this genus. From twelve specimens, maintained in a sealed container filled with the soil for six months, four matures specimens (which had regressed to a “sub-adult” stage with suppressed clitella) were released into new soil in imigated 18kg cores of clay. None survived after four months (October 1992-February 1993) when soi] temperature ranged 17-35*C. Despite this, their burrows and granular casts indicated some initial activity and the final yield of grain sor- ghum (Sorghum bicolor) was increased, com- pared to uninoculated contrals, by 40%. Heteroporodrilus dioecius (Stephenson, 1933) (Fig. 2) MATERIAL EXAMINED 2 mature cig OMG210139, Brookfield, Old, (2T'30'S,152'55'E), C.H. Thompson, 17 May 92. CSIRO Samford farm, Qld, (2722'S,152'53'E), 5 ma- ture specimens, R.J. Blakemore, 9 Apr 92 and 22 Jun 92, in author's collection, HABITAT Under litter on bank of nver; on surface after rains. EXTERNAL CHARACTERS Length 58-70mm. Width 2.5-3mm. Segments 90-103, body cylindrica) but segments tending to become trapezoid with only slight secondary an- nulation. Colour of anterior dorsum pigment light to dark brown, clitellum yellow-orange with fain} indescence, ventrum pale, Prostomium closed epilobous tapering to notch that extends dorsally to 1/2 (not as far in all specimens). First dorsal pore: 5/6 faint becoming obvious by 8/9 and either obvious and continuous over clitellum or hot. Setae 8 per segment from ii: lateral setal couples widely spaced; setae a of vi to xii seen tn be damaged or dehisced in some specimens and, on xviii a setae may be modified, (Samford, 1.$:1:1.5:2.2:3.3:0.30} (Brookfield, 2:1:2:2:5:0.3). Nephropores in D line 1/2-3/4, 9/10 and 11/12 then alternating, C line in 4/5-8/9, in B line 10/11, 12/13 then alternating. There is clear alternation between B and D lines for most of body length (in one Brookfield specimen the alternalion was asymmetrical within segments). Clitellum raised and cingular '/3xiii,xiv-xvi, /3xvii (may be inter- Tupted ventrally in xvii); furrows, setae and nephropores retained. Male pores not found. Female pores puired in xiv in common darker field anterio-median to 2 setae (more closely paired in Brookfield specimens). Spermathecal pores 6/7, 7/8 and 8/9 small and concealed in furrows in B line. Genital markings none, but in Iwo Samford specimens, tmid pads seen in aa in x and/or xi and possibly small unpaired genital markings on xviii. INTERNAL CHARACTERS Septa 4/5-9/10 moderately thick, 8/9/10 the thickest, 10/11-14/15 weaker, Dorsal blood ves- 26 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Heteroporodrilus dioecius, Samford. A, ventral view; B, reduced lateral view, C, spermathecae; D, prostomium; E, prostomium of second specimen, G, ventral view; F, spermathecae. Prostates were aborted in all specimens. sel single continues on pharynx. Hearts in x to xiii, COMmissurals in vi or vii-ix. Gizzard small and spherical, moderately muscular in v. Cal- ciferous glands 3 pairs, ventral on oesophagus in xi, xii and xiii. Intestine widening in xy, no typh- losole. Holonephridia with large bladders, seen from at least vi, alternating in position as for nephropores. Iridescent funnels of testes free, or in sacs, in x and xi, seminal vesicles in ix and xi/xii. Paired tufts of egg strings seen in xiii or in xii (this anomalous positioning was seen in two specimens from Samford). Prostates absent. Three pairs of spermathecae in vii-ix, spherical ampullae taper to ducts each with a medium length diverticulum (iridescent in several cases). Gut contents fine colloidal material, dead roots and organic debris. REMARKS This species was erected on aprostatic specimens from Toowoomba (Stephenson, 1933), but Jamieson (1970) col- lected specimens at Petrie near Brisbane that had racemose pros- tates and male pores. Possibly similarly male-fertile specimens were present in the populations from which aprostatic specimens were collected in the current study, as the diverticula were in- seminated. These specimens from Sam- ford and Brookfield clearly lie within the descriptions of H. dioecius; a variation from Jamieson (1970) (apart from lack of prostates) is that the pros- tomium is here interpreted as beìng furrowed, closed epilobous rather than tanylobous. Seminal vesicles were recorded in several of these specimens in ix and xi/xii which is unusual and possibly the material filling xi was mucal coagulum. Jt is unusual too to observe ovaries in xii (i.e. progynous). Stephenson (1933) found in one specimen that one spermatheca, from the three pairs, was absent. Prostatic morphs of this species differ from H. raveni principally in having one fewer pairs of cal- ciferous glands which are sessile rather than on short-stalks. Heteroporodrilus doubei sp. nov. (Fiz. 3) MATERIAL EXAMINED HOLOTYPE: QMG211897, 10km NW of Lismore, NSW, 28°48'S, 153° 17E, Dr. B. Doube, 26 Feb 1992, mature, drawn and dissected. PARATYPES: QMG211898, mature complete; OMG211899, mature posterior amputee. Same collec- tron data as holotype. REVIEW OF HETEROPORODRILUS 27 HABITAT In deep unstructured black clay soil-prairie podsol in deep vertical burrows in association with a Digaster sp. nov. EXTERNAL CHARACTERS Length 95-120mm. Width 3.5-4.0mm. Seg- ments 91 (holotype) body circular. Unpigmented, iransparent in alcohol, Prostomium closed epilobous. First dorsal pore 5/6, not clear on clitellum, Setae 8 per segment from ii, ab absent from xviii, whole length of setae can be seen through body wall after clitellum (2.13:1.0:1.63:1.75:6.0:0.36). Nephropores in D in 1/2-3/4? and 6/7 on left hand side, in C 5/6-8/9 in B in 9/10 on right hand side and 10/11 on left hand side then alternating between D and B for remainder of body. Clitellum J/2xiii-xvii pale coloured but slightly tumid. Male pores small in site of missing b setae on slightly raised mound. Female pore on xiv on left hand side only (holotype) or small and paired (paratype 1). Sper- mathecal pores concealed in furrows 7/8/9 in B. Genital markings single, ventrally offset disc on vii; two closely paired in anterior of viii with more lateral sets in longitudinal series in line with spermathecal pores; similar discs to vili on ix except the median discs are in the setal arc; pos- sibly the raised mounds on xviii are genital mark- ings. Markings not as distinct in paratypes. INTERNAL CHARACTERS Septa 5/6 weak and finely attached to sides and base of gizzard; 6/7-10/11 moderately thickened; 11/12 on thin. Dorsal blood vessel single onto pharyngeal mass in iv. Commissurals in vi-ix; hearts in x-xiii, the last three pairs larger, Ventral vessel bifurcated under gizzard. Fine, supra- oesophageal vessel seen in xii and xiii with lateral branches. Gizzard large, muscular in y but dis- placed almost to 6/7. Calciferous glands small, white, ventrally sessile pairs of glands in xi-xiir. Intestine widing suddenly in xv (acaecate, aty ph- losolate). Holonephridia from anterior with large, flimsy vesicles obvious in the clitellar region, alternating in position, Testis/sperm funnels iridescent, seen in x and xi but fragmented during dissection; large pairs of racemose seminal vesicles posteriorly in ix and anteriorly in xii, Ovaries not located. Prostates tubulo-racemose, confined to xviii but folded over flaccid duct. Spermathecae two pairs in viii and ix, bulbous ampullae narrow to shorter ducts, each bearing a pair of opposed clavate, iridescent diverticula. Gut contents fine soil and Acarina (litter mites) FIG. 3. Heieroporodrilus doubei sp. nov, A, ventral view: B, tubuloracemose prostate; C, spermathecae; D, prostomium. suggesting geophagous/deirifiyores diet. Mucus invests several antenor segments. 28 MEMOIRS OF THE QUEENSLAND MUSEUM REMARKS Heteroporodrilus doubei has 3 pairs of cal- ciferous glands and two pairs of bidiverticulate spermathecae, and is named after the collector. Similarities in allocation of calciferous glands and spermathecae are with Heteroporodrilus minyoni which, however, is much larger (400mm), and also with the bi-diverticulate H. lamingtonensis in which the position of the geni- tal markings are in x and xi and xvi-xxiratherthan in vii-ix. Heteroporodrilus doubei vindicates the separa- tion of Heteroporodrilus s. mihi from Plutellus s. stricto, outlined in the emendation of the genus above, as it has sessile calciferous glands and tubuloracemose prostates that would have bridged the previous generic definitions. Heteroporodrilus jamiesoni sp. nov. (Fig. 4) MATERIAL EXAMINED HOLOTYPE: OMG211900, Mt Glorious, Old, 27720'S, 152746'E, B. Jamieson collection, labelled “Or5 OSp 27", 12 Aug J971, previously dissected mature specimen. HABITAT ?rainforest EXTERNAL CHARACTERS Length 110mm (posterior amputee). Width 6mm. Segments 154+. Body spherical without dorsal canaliculation, peristomium wrinkled, preclitellar ventrum somewhat corrugated, secondary annulation slight, Unpigmented buff in alcohol, clitellum darker. Prostomium closed epilobous with distinct central cleft. First dorsal pore 6/7? wide, continuous on clitellum. Setae 8 per segment obvious from iii, a and b setae absent from xviii; lateral setal couples widely spaced (2.0:1.0:1.8:3.0:4.0:0.23). Nephropores at anterior margin of segments: 1i/D, iii/D, iv/D or C, VIC, vi/C, vii/C, viii/C or D, ix/C or D, x/D, xi/B then symmetrically alternating for remainder of body. Clitellum annular /axiii- /2xvii, interrupted ventrally in xvii. Furrows, setae and nephropores retained. Male pores on elongate porophores on xviii in B line. Female pores paired on xiv anterio-median to a setae, Spermathecal pores five pairs 5/6-8/9 in B line. Genital markings in xvii-xx widely spaced by the width of the a setae, wholly presetal, laterally flattened pads which just impinge on the ventral setal couples. FIG. 4. Heteroporodrilus jamiesoni sp. nov, A, ventral view; B, prostate; C, spermathecae in situ; D, dorsal view of calciferous glands with the dorsal vessel removed. The peristomium was damaged dorsally and was not drawn, INTERNAL CHARACTERS Septa 5/6 weak, 6/7-12/13 becoming progres- sively stronger, then thinning; 5/6 and 6/7 dis- placed by gizzard, 10/11 and 11/12 converging and thickened. Dorsal blood vessel single con- REVIEW OF HETEROPORODRILUS 29 tinuous on pharynx. Commissurals noted in vi-ix, hearts in x-xiii. A supra-oesophageal vessel in x, strengthens from xi-xiij and, on each side, sup- lies the centre of each calciferous gland in these ast three segments with a connective that dis- sipates into capillaries. (Ventral vessels not in- spected). Gizzard muscular in v displacing septum 5/6 and 6/7, with wide nm anteriorly then lapering to appear cone shaped. On the oesophagus or posterior septum of vi-ix, small opalescent glands or sacs were also observed. Calciferous glands three pairs of spherical. al- most reniform calciferous glands ventro-lateral to oesophagus in xi, xii, and xii with numerous lamellae seen externally. To each gland a short stalk extends laterally from the oesophagus. and is directed downwards to the dorsal centre of each gland i.e. on short-stalks. Intestine widens in xv to reach its full width in xvi; typhlosolenol found. Holonephric throughout with large convoluted tubes and flauened, round terminal bladders in the position of nephropores. Male organs holandric; especially x, but in xi too, fairly thick- ened pericardic (testis) sacs extend between the septa and encompass the blood vessels, oesophagus and the large indescent funnels of the testes. Seminal vesicles are paired and racemose in ix posteriorly and xii anteriorly on septa, Ovaries smal] compact glands found in the anterior ventrum of xiii but individual ova not visible. Prostates compact, tongue-like. racemose glands confined to xviii each with a short, flaceid duct. Spermathecae five pairs in v-ix (the anterior pair obscured by gizzard and septa). Elongate, conical ampullae attach to tapering ducts each with a single simple digitiform diverticulum. The right side posterior diverticulum was bifid with a smaller, blunt branch, Iridescence was noted for each diverticulum. Gut contents almost entirely consisting of woody organic matter well mixed with some soil. REMARKS Heteroporodrilus jamiesoni has three pairs of calciferous glands and five (or four) pairs of spermathecae and corresponds with Jamieson's (1970) Mt Glorious specimen he placed, despite several differences in the accounts, in H. ashwaor- thi (Stephenson, 1933) (here considered a junior synonym of H. oxleyensis). The present specimen is especially similar with regards to the illustra- tions (Jamieson, 1970: figs 2; 3a; 8a,b; 9a; 10a). Differences are that only four, rather than five spermathecae were found, testis were free rather than in testis sacs as here and the ovaries and funnels were (erroneously) recorded in xiv. Dis- tributions of genital markings concur except for the absence in the specimen before me, of an obvious pair of presetal elliptical pads in ix and x. This species can be separated from H. oxleyensis on the basis of genital markings, but not on testis sacs, which have been recorded as or absent for both species, The spermathecae of H. oxleyensis and H. jamiesoni are similar in appearance and arc recorded as 4 or 3 pairs and 5 or 4 pairs, respectively. Separating species principally on genital markings, which may vary with season or physiological or ontological state, is perhaps tenuous. But, in cases where population ranges overlap, such features are perhaps emphasised and may have a greater impor- tance for speciation. Such small apparent variations in genital markings could be crucial deciders for mate recognition. Of interest is that the short stalks on the cal- ciferous glands in this specimen are similar Io those for H. oxleyensis (Mt Collon specimen) as well as those illustrated for Hereroporodrilus minyoni comb. nov. (Dyne, 1980: fig. 2b). Heteroporodrilus oxlevensis (Fletcher, 1889) (Figs 5, 6) MATERIAL EXAMINED OMG210140, 2 mature, dissected specimens, Lower Savages Rd.. Brookfield (27°30°S,152°55°E), early mornings, C.H. Thompson, 17 May 1992; I mature specimen, same locality data as previous, 31 Dec 1992, author's collection; OMG210141, 1 mature specimen (with the posterior amputated but retained) and three sub-adult and two immature specimens, Queensland University farm, Mt Cotton (27°53'5, 153" IFE), R.J, Blakemore, S. Jeffries, 19 Jan 1993; sub-adult specimens, same collection data as previous, A. Wilkie, Aug & Oct, 1992, author's collection; one mature specimen, possibly a postenor amputee, Samford CSIRO farm (27°22'S, 152°53’E), R.J. Blakemore, V.R. Catchpoole, 5 Feb 1992, author's collec- tion. Descriptions following are in the same order as Lhe three locations. HABITAT On road near remnant rainforest: under “blue gum’ (Eucalypres rereneornis) and Leucaena leucociphala plantation, particularly in red ped- solic soil; under grass in a mon-calcic brown soil, EXTERNAL CHARACTERS Length of mature specimens: 100-120mm; 140mm; >92mm (less mature specimens: 33- 110mm). Width of larger specimens: 5.5-6mm; 4mm; $mm. Segments of mature specimens: 30 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 5. Heteroporodrilus oxleyensis, Mt Cotton, ma- ture specimen, A, ventral view; B, prostate; C, sper- mathecae in situ. D-F, dorsal views. D, prostomium; E, calciferous glands; F, posterior. 161, 185, 190; 167; >119 (others: 113-159). Secondary annulation minimal, Slight dorsal fur- rows in caudal segments. Colour of dorsum, espe- cially in anterior and caudal segments, pigmented light yellowish-brown with slight iridescence (Samford specimen dark grey), ventrum pale; clitellum buff. Prostomium closed epilobous with distinct dorsal cleft extending from tip of pros- tomium through to 1/2 or 2/3 (seen in all larger specimens). Immature specimens have furrow to 1/2 only but short grooves also flare from pros- tomial junction. Pharynx may evert on preserva- tion. Peristomium usually rugose. First dorsal pore 5/6 faint, 6/7 wide; 6/7 rudimentary, from 7/8 distinct; obvious from 7/8. Setae 8 per seg- ment from ii, a and b (always?) absent from xviii; lateral setal couples widely spaced, (2:1:2:2:4.3:0.27); | (2.0:1.0:2,0:2.3:3.7:0.23); (2:1:2:2:3:0.3). Nephropores in D line 1/2-3/4, 5/6 or 6/7, then alternating; in C line, when not in D in 4/5-7/8 and alternating in B line from 9/10. Clear alternation between B and D lines for remainder of body, In some specimens, FIG. 6. Heteroporodrilus oxleyensis, Samford specimen. A, ventral view; B, prostate; C, spermathecae; D, pros- tomium; E, ventral view of calciferous glands, REVIEW OF HETEROPORODRILUS al nephropore alternations were asy! ical in B and D per side. Clitellum Mxiii-/axvii,xvii in- terrupted ventrally in xvii; setae, nephropores, furrows and dorsal pores retained, Male pores on xviii on slightly raised porophores, ap- proximately in B line (on immature and sub- adult specimens seen within small lateral creases). Female pores on xiv a pair anterio- median to a setae in common darker lateral patch. Spermatheca] pores four pairs in 5/6-8/9 in furrows in B line. Genital markings of Brookfield specimens: in some of vi-xi small closely paired, median pads, tn xvii closely paired or clongate postsetal pads, xviii-xx (and xxi in one unusual specimen) tumid, elongate, mostly presetal pads; Mt Cotton: in vii and viii single or paired faint discs anterior to ventral setal couples, in ix and x closely paired discs between @ setae, also tn ix a smaller pair of markings just behind the spermathecal pores, in xvii a pair of postsetal pads, closely paired in common ventral tumid area and mostly prese- tal in each of xviii-xx: elongate smooth pads sunken in otherwise raised glandular areas; Sam- ford: in ix-xi small, ventrally paired discs, in xvii postsetally and presetal in xviii-xx large, paired pads elongate from mid-ventrum to B lines. INTERNAL CHARACTERS Septa 4/5 membranous to anterior of gizzard, 5/6 thin and much displaced to accommodate gizzard, 6/7-11/12 only slight, but progressive thickening, 12/13-14/15 thinning and thereafter membranous. Septa 4/5-8/9 are posteriorly dis- placed to a lessening degree, 9/10 and 10/11 are adherent, Pharyngeal tendons extend through anterior septa. Dorsal blood vessel single con- tinuous onto pharynx. Commissurals from vi-x, hearts in xi-xiii. In Mt Cotton specimens: supraoesophageal vessel not found in xi but developed in xii and xiii with small connective to dorsal heart valves and lateral branches attached to stout pink stalks that pass to dorsal centres of the calciferous glands on each side, The hearts pass to ventral Vessel, dorsal to which is a separate vessel Ihat seems to supply the calciferous glands al their ventral axes. The dorsal vessel from xiv has two small pairs of vessels in each segment that pass to each side of the alimentary canal, Gizzard tubular to spherical and muscular in v. Calciferous glands three pairs in xi, xii, and xiii; spherical glands with white lamellae in section and visible externally, lay ventro-lateral to the desophagus on shart stalks (in one Brookfield specimen the glands are anomalous in xi then in xiv and xv on left side only). Intestine widening through xv; no typhlosole, Nephridia paired holonephridia with long convoluted tubes and deflated, subspherical to elongate, terminal blad- ders, from at least v, corresponding in position to the nephropores. lridescent testes and funnels ventrally in x and xi within pericardic testis sacs (strong in x, more membranous in xi); a small pair of racemose seminal vesicles posteriorly on septa in ix (and xi) and (Jobulate) anteriorly in xa, A compact pair of glandular ovaries (individual ova not discemible) in antenor and nebulous oviducis in posterior of xiii. Prostates paired racemose glands, confined to xviii each with a short, en- sheathed duct bent in single loop. Spermathecae four pairs in vi-ix; conical, fleckled ampullae connect to tapering ducts each with a singlediver- ticulum, Tridescent (inseminated) diverticula only noted in Samford specimen. Gut contents mostly organic matter including woody (dead root) material, plus some fine and some coarser soil, REMARKS Despite some variation between the specimens described above from the three locations, iL was decided to include this new material in H. ox- leyensis, The original brief description by Fletcher (1889) was based on one aclitellate specimen (plus two incomplete specimens). Sub- adult specimens from Mt Cotton differ insubstan~ tially from this original description apart from having about 15-20 fewer segments. Moreover, the mature specimens from Mt Cotton and the other two locations above agree sufficiently with each other and with Stephenson’s (1933) single specimen of Woodwardiella ashworrhi, here regarded as a junior synonym of H. oxleyensis. Justification for this synonymy is that Stephen- son (1933) omilled to refer to the earlier descnp- tion, yet his specimen from Brisbane agrees on every point in common with Fletcher's (1889) H, oxleyensis apart from those attributable to its being mature, The only other records of H. ox leyensis are the two specimens (from Kholo Creek und Marrickville) described and il- lustrated by Jamieson (1970), These also con- cur with Stephenson's (1933) specimen except thal they are slightly truncated, have dorsal pores in 5/6 or 6/7 (rather than 7/8), do not retain setae a of xviii, one has three pairs of spermathecae (rather than four) and they have paired postsetal genital markings in xvii (rather than an analogue, Stephenson, 1933: fig. 12). 32 MEMOIRS OF THE QUEENSLAND MUSEUM xv FIG. 7. Heteraporodrilus thompsoni. A, ventral view of paratype; B, prostate: Heteroporodrilus thompsoni sp. nov, (Fig. 7) MATERIAL EXAMINED HOLOTYPE: OMG210142, Lower Savages Rd. and near Gold Creek, Brookfield, Old (27°30°S,152°55'EB), C. H. Thompson, 08 Apr 1992. PARATYPES: 6 specimens, QMG210143- G210146, same collection data as holotype, 15 Apr 1992, 17 May 1992, 30 Dec 1992. HABITAT Under leaf litter of rain- forest and eucalypt woodland and on road surface, especial- ly after rain. EXTERNAL CHARACTERS Length 210-270mm (holotype: 210mm, but some damage to middle area), Mass 21.5g(holotype). Width (midclitellar) 11-13mm. Seg- ments 181-235, (holotype 235). Faint secondary annula- tion, especially after the clitellum. One specimen had marked dorsal furrow (i.e. canaliculate) in the latter half of its body. Unpigmented, grey in alcohol; clitellum darker pink-grey and faintly iridescent. Prostomium closed C, spermathecae in sim; D, dorsal view of prostomium; E, ventral view of epilobous to pro-epilobous calciferous glands with ventral vessel cut away; F, anterior view of calciferous glands from xiii with left-side sectioned; G, posterior. Confusion over the definition of H. oxleyen- sis that resulted mainly from its designation on sub-adult material (as was H. sloanei, also by Fletcher, 1889) has been reduced by this revision. The combination of three pairs of calciferous glands in xi-xiii (mislabelled in Jamieson, 1970: figs. 3d, 5g as in ix), four (or three) pairs of spermathecae and postsetal geni- tal markings in xvii are unigue to this species. The distribution in the Brisbane region of southeast Qld. is dissipated only by the one anomalous record from the suburb of Marrick- ville, Sydney, NSW (collector M.E. Grey, 1933). with definite furrow that bisects prostomium and ex- tends to 2/3 as a groove, Peris- tomium rugose. First dorsal pore not clearly detectable in deep furrows but seen in 10/11 and 13/14 at least, and again from 17/18. Setae 8 per segment, from ii, dark tipped, ab absent from xviii (mean of four specimens; 1.7:1.0:1.5:2.5:3.8:0.26). Nephropores at anterior margin of segment almost in furrow, obvious on clitellum. The most common arran- gements: ii/D, iii/D, iv/C or D, V/C, vi/C, vii/C, viu/C or D, ix/D or C, x/B, xi/D, xii/B, xiii/D, xiv/B, xv/D, xvi/B, xviii/D, xviii/B, xix/D, xx/B, xxi/D, etc, Several specimens had ir- regular pore distributions to the above scheme, one had the opposite arrangement from x, REVIEW OF HETEROPORODRILUS 33 another two had asymetrical alternations within a segment (i.e. a pore in B and one in D}. Clitellum annular and tumid xiv-xvji, setae and nephropores retained often in deep dimples. Male pores in xviii in site of b setae on slightly raised porophore. Female pores on xiv variously closely paired anterio-median to aa, sometimes concealed in a lateral groove, alternatively a single pore on a slightly tumid pad anterio- median to aa setae. Spermaihecal pores two pairs in 7/8 and 8/9 almost hidden in furrows in B line but visible under traction. Genital mark- ings in x in all specimens: a distinct pair of circular, tumid pads occupying the longitudinal width of the segment and encompassing the ventral setal pairs, In xviii, the areas of the missing ventral setal couples are generally tumid with a mid-ventral hollow and in xix is a similar glandular area which extends as far as the limit of the ventral setae. INTERNAL CHARACTERS Septa 4/5-7/8 moderately thickened, 8/9- 12/13 becoming progressively much thicker, then, after 13/14, thinning. Dorsal blood vessel single, Continuous on pharynx, Commissurals in vii-ix, large hearts in x-xiii. From ix or x to xiv or xv Jarge bladders dominate the dorsal yessels, in xv the enlarged dorsal vessel has two pairs of ventrally directed vessels which sur- round the oesophageal valve, Bifurcate ventral blood vessels form a pair of hairpin loops under gizzard before moving anteriorly. Gizzard large, muscular and barrel shaped in v with slight anterior rim, compressing septa 5/6 onto 6/7 and reaching as far back as vii, Calciferous glands four pairs in x-xiul, the anterior pair the smallest, as spherical, white, ventral pouches sessile on oesophagus cach with numerous fine lamellae internally. A pair of sub-oesophageal blood ves- sels run between the lobes of each pair of glands. Intestine origin abrupt in xv or xvi, no typh- losole. Holonephric, with large, subsphencal to elongate, flattened bladders corresponding in position with external nephropores. Male or- gans holandric: large paired, iridescent fun- nels of testes free (or appearing in testis sacs in at least lwo specimens) in x and xi. Seminal vesicles medium to small size, paired and racemose in ix and xii (the latter Jobulated on anterior septum). In ix white coagulum, the same texture as in seminal vesicles, frequently scen- Ovaries as a small pair of egg-string tufts anteriorly im xiii and paired diaphanous oviducts on posterior ventrum, Prostates con- fined Io xviii, a pair of compact, sguarish, racemose glands with short, bent duct joined en- tally by vasa deferentia. In section the prostates appear solid without obvious central lumina. Spermathecae two pairs in viii and ix: large bulbous but slightly deflated or deflected subspherical ampullae with fine concertina- like corrugations, attach to shortish ducts bearing several (two to six), small digitiform or bi-. tri-, or multi-lobed diverticula encir- cling the duct near the ectal end, Stalks (and occasionally termina) of several diverticula have lustre indicating insemination. Gut con- tents mostly reddish silt with some grits and sometimes organic debris including woody remains (geophagous/detritivorous). REMARKS The combinations of four pairs of calciferous glands and two pairs of polydiverticulate sper- mathecae identify H, thompsoni which is named after the collector. This species has close affinities with the similarly large H. tryoni, also collected from this location. It dif- fers from H. tryoni in having one fewer pairs of calciferous glands, one fewer pairs of sper- mathecae, different setal ratios and in having a distinct pair of genital markings in x, H. tryoni usually has a pair or trio of tortuous diverticula on each of it's three pairs of spermathecac, often with sessile seminal chambers. It is noteworthy that Boardman's (1932) description for H. youngi (syn. H. tryoni) has compound diverticula similar to those described here for H. thompsoni, but here again there are three pairs of spermathccae (even in his paratype where he reports only dissecting the first two sper- mathecae from segment vii and viii). Heteroporodrilus tryoni (Fletcher, 1889) Fig. 8) MATERIAL EXAMINED OMG210147, Adavale St and Lower Savages Rd, Brookfield, Old. (27 3075,152'55'E), C.H. Thompson, 8 Apr 1992, one damaged mature specimen dissected and drawn; QMG210148, 17 May 1992, one intact specimen; OMG210149, 26 May 1992, two damaged specimens dissected. HABITAT Under litter, on surface after rains and at about 30 cm depth in a deep surfaced euchrozem soil with clay-loam surface texture (field pH 6.0). 34 MEMOIRS OF THE OUEENSLAND MUSEUM EXTERNAL CHARACTERS Length 270-300mm. Mass 35.85g (damaged but complete specimen). Width (midclitellar) 14-17mm. Segments: 235-236. Peristomium wrinkled, second segment less so, Marked dor- sal furrow in latter part of body. Pigmented dorsum dark grey in alcohol; clitellum slightly darker with faint green tinge. Prostomium closed epilobous to pro-epilobous with definite furrow that bisects prostomium and extends to 2/3 as a groove, First dorsal pore in 6/7 or 7/8. Setae small, 8 per segment from ii, ventral setal couples widely spaced; ventral setae absent on xviii. (mean; 2.0:1.0:1.8:3.5:4.3:0,24). Nephropores at anterior margin of segment in furrow, irregular alterations noted for two specimens: ii/DD, i;VDD, iwDD, V/CC, vi/DB, vii/CD, viii/DC, ix/CD, x/DC, xi/CD, xii/DB, xHHBD, xiv/DB, xv/BD, xvi/DB, xvii/BD, XViii/DB, xix/BD, xx/DB, xxi/BD this alternation continued for the remainder of the bodies Nephropores do not open in B lines in vii, viii nor ix as these positions are occupied by the spermathecal pores. Clitellum annular and tumid: xiii, xiv-xvii but interrupted ventrally on xvii by genital markings. Setae and nephropores retained. Male pores paired in xviii on small circular porophores at centre of generally tumid patch centred at site of aborted b setae, Female pores on xiv in lateral groove stretching between a setae, but just anterior to setal arc. Spermathecal pores in 6/7, 7/8 and 8/9 almost hidden in furrows in B line but visible under traction as minute pores. Accessory geni- tal markings in x and xi slightly raised pair of tumid pads filling the antenor aspects and just incorporating the ventral setal pairs; in xvii a pair of sucker-like markings with the ventral setae in the lower edge; in xix a similar pair of pads although possibly slightly wider. INTERNAL CHARACTERS Septa 5/6 and 6/7 weak but adpressed, 7/8- 12/13 thick, 13/14 moderately thick, thereafter membranous. Dorsal blood vessel: single and considerable. Commissurals in vii-ix, large hearts in x-xiii. Gizzard moderately muscular and cone shaped in y with wider anterior rim. Calciferous glands five pairs in ix-xiii, the anterior pair the smallest, as spherical, white, ventral pouches sessile on oesophagus each with numerous fine lamellae internally. In- testine origin in xv, no typhlosole. Holonephridia with largish, subspherical col- lapsed bladders in position of external v . A ND ~ aor D C ES 5», M EN FIG. 8. Heteroporodrilus tryoni. A, ventral view of an anteriorly damaged specimen; B, prostate gland; C, spermathecae in situ; D, enlargement of a sper- mathecae, nephropores, Iridescent funnels of testes free in x and xi close to ventral nerve cord. Seminal vesicles small and racemose in ix and xii. Ovaries as tufted strings in xiii with paired egg funnels on posterior ventrum. Prostates confined to xviii, a pair of tongue-like racemose glands with thin, straight, non-muscular duct, Spermathecae three pairs in vii, viii and ix: constricted, bulbous am- pullae narrow abruptly to ducts bearing ventrally, threemedium-sized, tortuousdiverticula, The central diverticulum in each case was the most convoluted. Iridescence was seen in all diver- ticula. The same grey coagulum as was found in the body cavity reappeared in the gut, The ingesta was mainly dead plant remains including large woody husks and small leaf skeletons. REVIEW OF HETEROPORODRILUS 35 REMARKS These large specimens are identified as H. Iryoni having five pairs of calciferous glands and three pairs of poly-diverticulate spermathecae. The present description consolidates that of Jamieson (1970) as the specimens here descnbed appear to have intermediate characters within the range of the earlier species description. The form is intermediate to the two heteromorphic specimens illustrated in the previous account (Jamieson, 1970: figs. 5b, 5c). H. tryoni appears closely related to, but discrete from. the sympatric H. thompsoni. Plutellus Perrier, 1873 emend. TYPE SPECIES Plutellus heteroporus Perrier, (presumably) Australian origin in NSW, 1873 nf REMARKS The generic description largely reverts to that of Jamieson (1971a), combined with features af the single Lord Howe Island species, P. hatchingsae Jamieson, 1977 (as shown in the brackets in the following diagnosis), DIAGNOSIS Small to moderately large terrestrial worms. Pros- tomium tanylobous (or epilobous). Combined pores of a pair of tubular (or ‘thickly tubular’) prostates and male pores in xviii. Spermathecal pores 4 or 5 pairs, the last in 8/9 in B lines (A lines), 8 setae per segment. Holonephric with large nephridial blad- ders; anterior nephropores in C or D lines and, from v-vii posteriorly, alternating from D to B lines. Gizzard, strong in v. Last heart xii (xiii). Four pairs of long-stalked ie. better than twice as long as broad, reniform calciferous glands in x-xiii. Holandric. Gymnorchous. Spermathecac monodiverticulate, DISTRIBUTION Restricted to the central coastal region of New South Wales; Lord Howe Island. SPECIES OF PLUTELLUS 1. Plutellus heteroporus Perner, 1873. Plutellus heieroporus Perier, 1873; 245-268, figs 1-3; Beddard, 1895: 487; Michaelsen, 1900: 174; Jamieson, 1971a: 1300-1310, fig. I (portions of type senes). 2, P. hutchingsae Jamieson, 1977:276-277, figs. 1; 2b; 6h; 7e-1; from Lord Howe Island, 3. P. manifestus (Fletcher, 1889) from Audley National Park and Bulli, NSW. Cryptodrilusmanifestus Fletcher, 1889: 1538-1539, (seven specimens); Buchanan, 1910: 215-216, figs 4,7. Plutellus monifestus; Michaelsen, 1900: 173; Jamieson, 1970: 129-132, figs 6a, b, 8e, £ 9h; {eleven clitellate and some aclitellate specimens), SPECIES INQUIRENDUM Plutellus rubens (Fletcher, 1887), from Mt Wil- son, NSW, Cryptodrilus rubens Fletcher, 1887a: 381-383, (a few small spirit specimens) Megascolides rubens; Beddard, 1895, Plutellus rubens; Michaelsen, 1900. Note: Fletcher conceded that spermathecal and nephridial pores were not visible in this holonephric species with “long narrow” prostates, although four pairs of calciferous glands were recorded in x-xiii and the last heart in xii. He went on to remark that the spermathecae and gizzard may have been over- looked for this small (SSmm) species, This imper- fect characterization has not subsequently been corroborated, KEY TO PLUTELLUS 1; 5 pairs of spermathecal pores, al anterior margins a” nia MNE reper poche b's 2 4 pairs of spermathecal pores, at anterior margins of vi- ix, almost contiguous midventrally P, manifestas E E 2. Spermathecal pores in B lines. Last hearts in xii Mb a Ee Ne P. heieroporus Spermathecul pores tn A lines. Last hearts in xiii ———' Or , P, hutchingsae —————'es'rumN:xr Plutellus heteroporus Perrier, 1873 (Fig. 9) MATERIAL EXAMINED Port Macquarie, NSW (31*27'S, 152°55'E), B.G.M. Jamieson collection: 2 specimens (mixed with species of several other native genera): one aclitellate, onc posterior amputee al 16/17; both drawn and described here, 8 miles from Beechwood on Bellangry and Wil- son River St. Forest Rd. Riverine forest sloping into gully, black soil under logs and in carth, W, Nash, R. Raven, 17 Jul 1975; 5 specimens tagged PH] - PHS (PH1 dissected, PH2 drawn, other specimens in- spected), near Cairncross State Forest, 6.4km S of Telegraph Point turnoff by Pacific Hwy. Wet black soil in paperbark and tea tree swamp, W. Nash, R. Raven, 16 Jul 1975. 36 MEMOIRS OF THE OUEENSLAND MUSEUM FIG. 9. Pluzellus heteroporus. A, ventral view of aclitellate specimen; B, tubular prostate; C, spermathecae; D, prostomium; E, dorsal view of calciferous glands in situ; F, anterior view of calciferous glands of xiii showing long ducts beneath lateral hearts; G, ventral view of specimen PH2; H, ventral view of posterior amputee specimen; I, spermathecae in situ (prostates lost). EXTERNAL CHARACTERS Length 77-175mm. Width 4-Smm. Segments 185-209. Body circular in section with secondary annulation dorsal from x and in post-clitellar seg- ments. Pigmentless buff in alcohol, not irides- cent, clitella manilla. Setal lines appear darker in some specimens. Prostomium tanylobous, First dorsal pore 7/8. Setae, especially ventral couples, dark, 8 per segment from ii, a and b absent from xviii, (mean of three setal ratios, 1.7:1.0:2.3:2.1:2.9:0.2). Nephropores paired in D line 1/2, 7/8, 9/10, 11/12, 13/14, 15/16, 17/18, etc.; in C line 2/3, 3/4, 4/5, 5/6 (one specimen in 6/7 too, another specimen in 8/9 also); in B line 6/72, 8/9, 10/11, 12/13, 14/15, 16/17, 18/19, etc. One specimen had irregular altemations, but mostly alternation between D and B in successive segments. Clitellum '/sxiii(dorsally), xiv-xvii an- nular, setae and nephropores retained, interseg- mental furrows faint, dorsal pores occluded. Male pores minute in xviii on small raised porophores in B line, each within a smooth, flattened, sub- spherical dish that fills the segment longitudinal- ly. Female pores in xiv a mìnute pair widely spaced but anterio-median to a setae, Spermathe- cal pores five pairs in B lines from 4/5 - 8/9. Accessory genital markings in one specimen, a pair of large but lowset presetal pads with darker centres between ab in x and xi extending from the setal arc to the anterior furrow. In other specimens, these marks were not clearly defined but a similar pair of pads were present in one in xi, on the left side only in one and in xii in another. Postsetally in xvii, a pair of ellipsoid smooth dishes between ab with translucent centres and shelving towards xviii; in xix a similar pair of marks presetally. Ellipsoid dishes, presetal, closely paired (occasionally single) and ventral to aa also present in each specimen in some of xx-xxiv (commonly in xx-xxii). REVIEW OF HETEROPORODRILUS 7 INTERNAL CHARACTERS Septa 4/5 membranous, 5/6-12/13 moderately thickened (especially the first three impinging rearwards), 13/14 and 14/15 thinner, then membranous, Dorsal blood vessel single con- tinues on pharynx. Commissurals vi-ix, larger hearts in x-xii , Gizzard thin walled but muscular in v, tubular or tapering. Calciferous glands four pairs laterally in x, xi, xii and xiii; rosette or reniform with radiating lamellae attaching to the oesophagus dorso-laterally via long curved ducts, one from each gland, Ducts well supplied with blood capillaries and originate near the centre of each gland. The glands are solid and consist of numerous internal lamellae. Oesophagus narrows in xiv (valve?) then widens in xv as intestine proper; in xv and xvi it is especially dilated; typhlosole and caeca not seen. Holonephric from ii. Large elongate bladders are associated with the nephropores i.e. they have altemate extensions in successive segments for most of the body, Indes- cent funnels of testes free in x and xi; racemose seminal vesicles occupy ix and xii, Ovaries paired ventrally in xiii, numerous egg strings are loosely attached to give fan-like appearance. Prostates a pair of elongate and tortuous tubular prostates (internal lumina not visible) overlaying short muscular ducts (dilated near body wall) in xviii. The prostates extend from xviii to xix (one abnor- mal specimen had a continuation of the gland directed forwards as far as xvi). Spermathecae five pairs im v-ix: elongate ampullae taper 1o narrowing ducts each bearing a single clavate diverticulum on stalks almost as long as ampulla (in mature specimens the diverticular bulbs were iridescent), Gut contents colloidal dark soil with no obvious Organic debris. REMARKS The above specimens correspond with previous descriptions of the type-series of the genus Plutel- lus, in particular that of Jamieson (197 1a). Addi- tional features recorded here are the first nepndiopores in D line in 1/2, new body dimen- sions, segmental counts, setal ratios and acces- sory genital markings. Large ventral disks in x and xi developed in one specimen resemble those seen in P, manifestus (see Jamieson, 1970: fig 6a), New illustrations are provided, Plutellus heteroporus has very close affinities with P. manifestus from NSW and P. hutchingsae from Lord Howe Island, not least in having tubular prostates. Characteristics of the genus are the alternation of nephropores and, separating it from both the allied Australian genera Heteroporodrilus and the monotypic Lord Howe Island Paraplutellus Jamieson, 1972 (see Jamieson, 1977) the possession of four pairs of clearly stalked calciferous glands. Jamieson (197 1a), related how two lots of specimens were mixed in the same bottle in the Paris museum where Perrier first descnbed type material in 1873. At about the same time, Digaster lumbricoides Perrier, 1872 was erected from specimens collected at Port Macquarie in 1846 (by M. Le Souef?). It is significant that this locality is the same as for the P. heieroporus specimens described above. The provenance of this new material then, supports the probable type-locality as being in Australia, and in par- ticular Port Macquarie NSW, rather than Pen- nsylvania, (USA) as was presumed by Perrier. DISCUSSION Plutellus clearly retains the more ‘primitive’ tubular form of prostates, Heieroporedrilus is thus considered an apomorphic sibling-group of Plutelius that has reached a climax in rainforest and nverinc environments in the Old. subtropics. This view contrasts with that of Dyne (1981: 105) who considered sessile calciferous glands as a pleisiomorphic condition and the stalk-glanded Plutellus us the more derived genus. Evidence that Heteroporodrilus has diversified in south-east Qld. is provided here by surveys that have located four of the six previously described species for this region, and added several more to the list. One small area at Brookfield in Brisbane has yielded four species in association: H, tryoni, H. thompsoni, H. dioecius, H. oxleyensis, often in the same sample. If the origin of the genus is taken 10 be south-east Qld., then the focus of endemicity (as indicated by the abundance and diversity of species and specimens) may well be in the catchments and rainforest foothills of the D' Aguilar Range of which Mt Glorious is n. high point, Nevertheless, the diversity of Heteroporodrilus species around Brisbane may be an aberration of sampling effort especially as much of the inland areas of the eastern sub- region have yet to be systematically sampled and are expected to provide further species, A fuller picture of the biogeography, as with the systematics of these and other species, may await genetic analyses. That the types of H. dioecius were trom Toowoomba on the Great Dividing Range demonstrate that this escarpment is no barrier to heteroporodrilid migration. Another species, 37 MEMOIRS OF THE OUEENSLAND MUSEUM H, bongeen, was recorded from the Condamine River plain west of Toowoomba in the Darling Downs region. This river, which at one time may have united with the Burnctt River, even- tually connects with the Murray-Darling river systems of New South Wales, Victoria and South Australia where distributions of other Heteroporadrilus spp. have been recorded. The riverine transport! mechanism for this genus is thus supported. The propensity for nocturmnal or crepuscular wanderings (particularly after flooding?), noted for several] species above, may further enhance their ability to disperse. Several members of this genus are associated with ripanan habitats where their burrows can easily access the water-table, they may thus survive apparently dry conditions, Osmotic regulation and secretion via large, exonephric terminal bladders, are perhaps enhanced by nephrophore alternation. Possibly this adapta- tion serves partly to maintain a more even body surface moisture, for respiration, lubrication and defence, especially in worms with larger surface areas, than would a regular series of nephropores. Both Piutellus and Heteroporodrilus contain medium to large sized members. However, alternation of nephropores is found in other gencra too eg. Maoridrilus Michaelsen, 1899, from New Zealand and in at least one member of the Australian native Diploirema (D. heteropora Dyne, 1979). ACKNOWLEDGEMENTS Professor B.G.M. Jamieson is thanked for suggestion of the topic and for critical appraisal of the manuscript. Mr C.H. Thompson, CSIRO soil scientist. Other collectors are acknow- ledged in the text. This paper was completed thanks to the provision of facilities by Dr Mau Colloff at CSIRO Division of Entomology, Canberra- LITERATURE CITED BAGE, F. 1910. Contribution to our knowledge of Australian earthworms. The nephridia. Proceed- ings of the Royal Society, Victoria 22: 224-243, BECKMANN G, G. & THOMPSON, C. H. 1960, Soils and land use in the Kurawa area, Queensland, CSIRO Australia, Division of Soils, St, Lucia, Series No. 37. BEDDARD, F. E. 1887. Contributions to the anatomy of canthworms - No. IV. Proceedings of the Zoological Society of London 1887; 544-548, 1895, ‘A monograph of the Order Oligochaeta’, (Oxford: Clarendon Press). BOARDMAN, W. 1932,Some earthworms from Queensland. Memoirs of the Queensland Museum 10: 125-130. BUCHANAN, G. 1910, Contributions to our knowledge of Australian earthworms. The blood vessels - Part II, Proceedings of the Royal Society, Victoria 22: 59-84, DYNE, G. R. 1979. A new species of bioluminescent earthworm from North Queensland belonging to the subgenus Diplotrema (Oligochaeta:Megas- colecidae). Memoirs of the Queensland Museum 19(3): 373-379. 1981. Three new species of the earthworm Genus Plutellus s. strict. (Megascolecidae: Oligochacta) from New South Wales and Queensland. Proceedings of the Linnean Society of New South Wales 105(2): 95-106, 1984. Systematics of Australian Megascolecoid Earthworms. (Unpublished PhD, Thesis: Queensland University). FLETCHER, J. J. 1887, Notes on Australian Earthworms, Part II. Proceedings of the Linnean Society of New South Wales (2)2: 375-402. 1387b, Notes on Australian Earthworms. Part TV, Proceedings of the Linnean Society of New South Wales (2)2: 601-620. 1889. Notes on Australian Earthworms. Part V. Proceedings of the Linnean Society of New South Wales (2)3: 1521-1558. 1890, Notes on Australian Earthworms, Part VI. Proceedings of the Linnean Society of New South Wales (2)4: 987-1019, GATES, G.E. 1959, On a taxonomic puzzle and the classification of the earthworms, Bulletin of the Museum of Comparative Zoology, Harvard 123(6): 229-261, 1961, On some Burmese and Indian earthworms of the family Acanthodrilidae, Annals and Magazine of Natural History (13)4: 417-429, JAMIESON, B.G.M. 1970. A revision of the Australian genus Woodwardiella with descriptions of two new genera (Megascolecidae: Oligochaeta), Jour- nal of Zoology, London 162: 99-144, 1971a. Description of the type-species of the earthworm genera Plutellus and Digaster (Megascolecidae: Oligochaeta). Bulletin du Museum National d'Histoire Naturelle, Paris 42(6): 1300-1310, 1971b. A review of the Mcgascolecoid carthwnrm genera of Australia. Part lll - The subfamily Megascolecinac. Memoirs of the Queensland Museum 16(1): 69-102. 1974, Earthworms (Megascolecidae: Oligochacta) from South Australia. Transactions of the Royal Society, South Australia 98(2): 79-112. 1977. The indigenous earthworms (Megas- colecidae: Oligochaeta) of Lord Howe Island. Records of the Australian Museum 30: 272-308. REVIEW OF HETEROPORODRILUS 39 JAMIESON, B.G.M. & NASH, W. 1976. The first record of Plutellus (Megascolecidae: Oligochaeta) in Queensland P. incommodus and P. raveni spp. nov. Proceedings of the Royal Society, Queensland 87: 45-52. JENSZ R.L. & SMITH, B.J. 1969. Catalogue of Baldwin Spencer types in the National Museum of Victoria, Australia. Memoirs of the National Museum of Victoria 29: 85-110. MICHAELSEN, W. 1900. 'Das Tierreich', Vermes, Lief. 10, Oligochaeta. (Friedlander: Berlin). 1907. ‘Oligochaeta in Die Fauna Siidwest- Australiens'. 1(2): 117-232. (Jena: Gustav Fis- cher). 1916. Results of Dr E. Mjobergs Swedish Expedi- tions to Australia 1910-1913. Kungliga Svenska Vetenskapsakademie ens Handlingar 52(13): 3- 74 PERRIER, E. 1873. Etude sur un genre noveau de lombriciens (Genre Plutellus E.P.). Archives de Zoologie Experimentale et Generale 2: 245-268. SPENCER, W. B., 1900. Further descriptions of Australian earthworms. Proceedings of the Royal Society, Victoria 13(1): 29-67. STEPHENSON, J. 1933. Oligochaeta from Australia, North Carolina, and other parts of the world. Proceedings of the Zoological Society, London 1932: 899-941. WALLACE, C.C. 1972. An examination of the clas- sification of some Australian Megascolecid earthworms by numerical methods. Memoirs of the Queensland Museum 16(2): 191-209. 40 MEMOIRS OF THE QUEENSLAND MUSEUM PIGMENTATION BANDS ON AN EARLY DEVONIAN NAUTILOID. Memoirs of the Queensland Museum 37(1): 40. 1994:- A specimen of an orthoceratid nautiloid was collected in 1990 from the Martins Well Limestone, Broken River Province, north Queensland (QM Locality 549). The Martins Well Limestone member of the Shield Creek Forma- tion contains a diverse fauna indicative of an Early Devonian (Lochkovian- Pragian) age (Jell et al., 1993). The specimen (OMF32208) is a 92mm long fragment of an orthoconic shell (Fig. 1) and is exposed only on the lateral parts of the shell with the dorsum and venter apparently obscured. The shell had been encrusted by an actinostromatid stromatoporoid skeleton, and the preserved shell surface represents this encrusted surface. Upon the outer shell surface are preserved thin, light grey sinuous bands 1.8-2.0mm thick, separated by thicker dark grey bands 3.3-4.4mm thick. The pattern is restricted to the exterior shell surface, and the pigmentation does not appear in longitudinal section as sutures or septal remnants. Ribbing is discounted given that a longitudinal section revealed a thin smooth, un-ribbed shell. The pattern is thus considered as a relict surface pigmentation. Preservation of pigmentation pallerns on Palaeozoic cephalopods is rare (Foerste, 1930; Teichert, 1964). Preserva- tion, in this case appears to have been facilitated by the early encrustation of the stromatoporoid, providing protection against crushing, and a significant buffer to destructive diagenetic processes. Literature Cited Foerste, A. F, 1930, The color patterns of fossil cephalopods and brachiopods, with notes on gasteropods and pelecypods. Contributions from the Museum of Paleon- tology, University of Michigan 3 (6): 109-150. Jell, J. S., Simpson, A., Mawson, R. & Talent, J. 1993. Biostratigraphy. Pp. 239-245. In Withnall, 1. W. & Lang, S. C. (eds), ‘Geology of the Broken River Province, north Queensland’. Queensland Geology 4. (Department of Minerals and Energy: Brisbane). Teichert, C. 1964, Morphology of hard parts, Pp, K13-53. In Moore, R. C, (ed.), ‘Treatise on Invertebrate Palaeontology’. Part K. Mollusca 3. (Geological Society of America and University of Kansas Press: Lawrence, Kansas) Alex G. Cook, Queensland Museum, PO Box. 3300, South Brisbane, Queensland 4101, Australia; 11 November 1994. FIG. 1. QMF32208, 7orthoceratid nautiloid showing pigmentation patterns, x 1. THE MARINE ISOPOD NEOCIROLANA HALE, 1925 (CRUSTACEA: CIROLANIDAE) FROM TROPICAL AUSTRALIAN WATERS NIEL L, BRUCE Bruce, N.L. 1994 12 01: The marine isopod Neocirolana Hale, 1925 (Crustacea: Cirolanidae) feet tro rh waters. Memoirs of the Queensland Museum 37(1): 41-51. Brisbane. I 79-8835. Two new species of Neocirolana are described from sub-littoral tropical waters of northern Australia: Neocirolana salebra sp. nov., from the Arafura Sea and Neocirolana tricistata sp. nov. from the vicinity of Torres Strait; new localities are also recorded for Neaciralana hermitensis (Boone, 1918) and N. excisa (Richardson, 1910), Observations indicate that Neocirolana hermilensis is a brood predator of hermit crabs. Comments are given on the current status of the genus, together with a list of all species and a key to the Indo-Pacific species. [_] Jsopoda, Cirolanidae, new species, northern Australia, taxonomy. Niel L, Bruce, Zoologisk Museum, University of Copenhagen, Universitetsparken 15, DK 2100, København Ø, Denmark; 30 August 1994. Neocirolana Hale, 1925 was established as a mo ic genus, later being revised by Bruce (1981, 1986). Since then Javed & Yasmeen (1990) placed a further two species in the genus, transferring Neocirolana obtrincata (Richardson, 1901) from Cirolana Leach, 1818, and describing the species Neocirolana arabica Javed & Yasmeen, 1990; Neocirolana tayronae Müller, 1993 was described from Colombia. In Australia the genus is represented by seven species, including the two new species described bere, six of which occur only within the tropics. METHODS Methods follow those outlined in Bruce (1986, 1993), ABBREVIATIONS BRR- Bureau of Rural Resources; CP- cir- cumplumose; CP- circumplumose setae; NT- North- em Terrilory; Northern Teritory Museum, Darwin, N.T.; PMS- plumose marginal setae; Qld- Queensland, Australia; QM- Queensland Museum, Brisbane; USNM- National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C.; WA- Western Australia. TAXONOMY Family CIROLANIDAE Dana, 1852 Neocirolana Hale, 1925 Neocirolana Hale, 1925: 153; Holdich ct al., 1981: 583; Brice, 1981: 955, fig. Gg, h; 1986: 200; Javed & Yasmeen, 1990: 71. TYPE SPECIES Neocirolana obesa Hale, 1925, by monotypy. DIAGNOSIS Head narrow, 50 - 60% width of pereonite 1. Pleon with 5 unfused segments; posterolateral margins of pleonite 4 encompassing and ex- tending posteriorly beyond posterior of leonite 5. Frontal lamina ventral surface flat. andible incisor narrow, usually less than 50% of maximal mandible width; spine row usually with 3 - 7 spines, molar usually reduced in size (in comparison to Cirolana), weakly toothed, Maxillule entire, or with reduced spination. Maxilla entire or often with reduced setation and reduction in size of lobes. Pereopods robust, ambulatory, not markedly flattened or expanded; pereopods 1-3 with anterodistal angles of ischium and merus not produced or lobate. Pleopods with PMS on all rami except endopod of pleopod 5; peduncles without lateral lobes; appendix masculina basally in- serted. Additional characters. Body 2-3 times as long as wide; pereonite 1 distinctly longer than pereonite 2; posterolateral margins of pleonite 3 often extending to or beyond posterior of pleonite 5. Antennnle peduncle with 3 major articles, article 4 minute; articles I and 2 subequal in length, partly or wholly fused. Mandible without lacinia mobilis. Maxilliped entire, or with various setal reductions, sometimes with endite reduced. Antennal peduncle 5 articled, Mandible palp 3 articled, extending beyond in- cisor. Pereopod dactylus with distinct secondary unguis. Penes short papillose or vasa deferentia 42 MEMOIRS OF THE QUEENSLAND MUSEUM opening flush with surface of stemite 7. Uropod peduncle inner margin produced along endopod medial margin; rami provided with PMS inter- spersed with spines. REMARKS Species of Neocirolana all have al least some reduction of the mouthpart appendages; most species (7 of 9) have a narrow mandibular incisor, often (7 of 9) accompanied by a reduced spine row of less than 5 spines and a molar process that is smaller than that of Cirolana; the narrow man- dible incisor appears to be associated with a nar- rower head width. Modifications also occur in the other mouthpart appendages, but these are mostly restricted to single species, and where similar may not be homologous. Meocirolana excisa, N. maculata and N, obesa all share a similar mouth- part, pereopod and uropod morphology, and ap- pear to form a group of related species, as indicated by Bruce (1986). Neocirolana arabica differs from Cirolana only in having few (3) spines on the mandibular spine row, and could arguably be better placed in Cirolana. Similarly, Neocirolana salebra, while very similar to Cirolana with regard to most genene characters, differs substantially in the spination of the muxi- llule, In contrast, all the mouthparts of N. obirun- cara are reduced or otherwise modified. Two species, N. bicrisia and N. tricristata, have longitudinal ridges on the pleoiclson, and also have similar mouthparts, but the pereopodal mor- phology of the two species is distinct. The conclusion drawn by Javed & Yasmeen (1990) that Neocirolana is a paraphyletic taxon for which the principal uniting character is a homoplasy is further supported by the two new species described here. The species currently placed in the genus are better retained in their present combinations, as other existing genera are less appropriate. SPECIES OF NEOCIROLANA Neacirolana arabica Javed & Yasmeen, 1990. Karachi coast of Pakistan, northern Indian Ocean. Types held at the QM. The species is otherwise very similar to those species of Cirolana which have a nodulose pleon and pleotelson, and uropods with dense seral development in the mees such as Cirolana pleonastica Stebbing, Neocirolana bicrista Holdich et al., 1981. Known only from shallow subtidal depths, Halifax Bay, Townsville, Queensland. Types held at the OM, Neocirolana excisa (Richardson, 1910). Type locality Cabalian Point, Jolo, Phillipines; com- mon in tropical Australian waters. Syntypes held at the USNM. Neocirolana hermitensis (Boone, 1918). An associate of hermit crabs, probably a brood predator (see species “Remarks” given here for that species). Common in tropical Australian waters. Location of the type matenal is unknown. Neocirolana maculata Bruce, 1986. Known only from the onginal records, Wistari Reef, Capricom Group, southern Great Barrier Reef and Marion Reef, Australian Coral Sea. Types held at the QM. Neocirolana obesa Hale, 1925. Type species. Types held at the Australian Museum, Sydney. Recorded distribution in southeastern Australia from Moreton Bay, southeastern Queensland to the Sydney region of New South Wales Neocirolana obtruncata (Richardson, 1901). Jamaica and Caribbean coast of Mexico (Martin & Felder, 1984). Types held at the USNM. Neocirolana salebra sp. nov. Described herein, Neocirolana tayronae Miiller, 1993. Caribbean coast of Colombia, in shallow water. Holotype held at the Zoologisches Museum, Berlin. Neocirolana tricristata sp. nov. Described herein. KEY TO THE INDO-WEST PACIFIC NEOCIROLANA 1. Dorsal surfaces nodulose Dorsal surfaces smooth, without nodules . . 2. Pereon, pleon and pleotelson with dsitinct nodules . 3 Pereon and pleon without distinct nodules, pleotelson with distinct longitudinal carinae . 4 3.Pereonites 7 only with nodules, pleon and pleotel- son with fine nodules and pleonites 3-5 with larger paired nodules; male uropod rami lateral margin with dense mass of setae . . N. arabica Pereonites 5-7, pleon and pleotelson heavily nodulose; male uropod rami without setal mass Si i a Mn ee N. salebra sp, nov. 4,Pleotelson with 2 widely spaced carinae and weak median carina; pereopod dactylus slender with minute secondary unguis N. bicrista Pleotelson with 3 closely spaced carinae; percopod dactylus robust with prominent secon- dary unguis N. tricristera sp. nov. 5.Uropod lateral margin without excision, apices not bifid; maxilliped endite reduced, without CPS or coupling hooks N. hermitensis r un r NEW MARINE ISOPODS 43 Uropod lateral margin with excision, apices bifid; max- illiped endite with CPS and 2 coupling hooks , .., 6 6.Uropod end deeply excised; maxilla middle and lateral lobes expanded ....... N. excisa Uropod endopod shallowly excised; media mn dle and lateral lobes not expanded , 7.Cephalon anterior margin without distinct rostral point; antennular flagellum hall as long as pedicle. «-c'ebe 6. sa ot awe Wy'n N. obesa Cephalon anterior margin with rostral point; anten- nular flagellum longer than peduncle . N. maculata Neocirolana salebra sp. nov. (Figs |, 2) MATERIAL EXAMINED HOLOTYPE: â (9.7 mm). Northern Territory, Anifura Sea, 11713.5'S, 129*42'E, 12 Dec 1989, dredged 23.8m, BRR,QMW 18711. DESCRIPTION Male holotype. Body about 2.7 times as long as greatest width; maximum width at pereonites 4-6, margins sub-parallel. Cephalon without rostral point in dorsal view; with submarginal anterior suture, with obscure ridge along medial margin of eyes; eyes large, occupying 50% width of cephalon in dorsal view; ocelli distinct. Pereonites 1-7 each with transverse suture, pereoniles 4-7 becoming increasingly nodulose; coxae 2-7 not prominent in dorsal view, all with prominent entire oblique carina; posterolateral angles of coxae 5-7 acute. Pleonites all visible; pleonite 1 smooth; pleonites 2-5 nodulose, with median nodules most prominent on pleonites 3-5. Pleotelson nodulose, median nodules most prominent. Antennule peduncle 2.8 times as long as width at article 2, articles 1 and 2 wholly fused, article 3 slightly shorter than combined lengths of ar- ticles 1 and 2; flagellum short, extending to anterior of pereonite 1, with 9 articles bearing abundant aesthetascs. Antenna peduncle articles | and 3 short, articles 4 and 5 subequal in length; flagellum with 14 articles, extending to posterior of pereonite 2. Frontal lamina pentagonal, about twice as Ìong as wide, with 2 submedian grooves. Mandible incisor with cusps indistinct, nearly as wide as maximal width; spine row with 13 spines, molar process with setules, anterior margin with 11 Widely spaced spines: palp articles 2 and 3 with 13 and 14 serrate spines respectively. Maxillule lateral lobe with 11 slender spines on gnathal surface, some of which are nodulose; medial lobe with 3 very slender spines. Maxilla lateral and middle lobe each with 3 setae; medial lobe with 2 CP spines and 9 simple setae. Maxilliped palp medial margins of articles 2-5 provided with short serrate spines, lateral margins with few setae; endite with 2 coupling hooks, 3 short simple setae and 4 long CPS. Pereopods with few setae. Pereopod 1 merus posterior margin laterally with 6 blunt tubercular spinesans 1 short acute spine, medially with 5 acute spines; posterior margin of carpus with ] acute spine; propodus with 3 acute spines, distal spine opposing dactylus base 20% length of dac- tylus, Pereopod 2 posterodistal margin of ischium with 1 tuUeFodiwr and I large blunt spine; posterior margin of merus with 3 shorttubercular, 2 longer stout blunt spines, and 3 acute spines; carpus with 2 acute and J blunt spines at posterodistal angle; propodus palm with 3 acute spines, Pereopod 7 with anterodistal angles of ischium, merus and carpus each with cluster of large spines, some of which are serrate; posterior margins of ischium, merus and carpus with short and stout spines and cluster of spines ar distal angle. Pleopod 1 with both rami distally rounded, endopod with ~13 PMS, about 0.3 times as wide as exopod, exopod with ~40 PMS and spine al proximolateral margin. Pleopod 2 exopod and endopod with ~50 and 15 PMS respectively, ap- pendix masculina straight with apex abruptly nar- rowed, extending very slightly beyond distal margin of ramus. Pleopods 3-5 exopods with the distal murgin distinctly indented,with transverse suture and with about 50 PMS; endopods each with -12, 10 and 0 PMS respectively. Uropod exopod about 0.8 times as long as endopod, straight, about 2.5 times as Jong as proximal widîh, with 2 prominent lateral submarginal nodules, lateral margin sinuate with 5 acule spines set among prominent PMS, medial margin with 4 spines among PMS, apex subeguall y bifid; endopod lateral margin indented at distal two- thirds of length, with prominent subapical nodule, with PMS and 1 spine adjacent to apex, medial margin smoothly convex, with 7 stout peg-like spines set among PMS; apex sub-bifid with lateral process prominent. REMARKS The nodulose pereon and pleon immediately distinguishes Neocirolana salebra from all other species in the genus. N. arabica has a nodulose pleon and pleotelson, but the nodules are fine except for the pairs of prominent submedial nodules on pleonites 3 and 4. There are numerous da MEMOIRS OF THE OUEENSLAND MUSEUM Uu, “gd cy “ ea) J rye wd ~, yt (Ee t 33 FIG. 1. Neocirolana salebra sp. nov. A, dorsal view; B, lateral view; C, frons; D, antennule, and detail of terminal flagellar articles; E, antenna, and detail of flagellar articles; F, maxilla; G, mandible; H, mandible, detail of incisor, molar and spine row; I, maxillule; J, maxilliped, j spine from medial margin of palp article 5. Scale = 2.0mm, 45 NEW MARINE ISOPODS pleopods FIG. 2. Neocirolana salebra sp. nov. A, pereopod 1; B, pereopod 7; C, pereopod 2; D, uropod; E-H, 1-3, 5 respectively. 46 MEMOIRS OF THE OUEENSLAND MUSEUM other differences between these two species, in- cluding the ornamentation of the frontal lamina, setation of the uropods and morphology of the mouthparts, those of N, arabica scarcely differ- ing from Cirolana, while the mandible, maxillule and maxilliped of M. salebra differ significantly from that of Cirolana. ETYMOLOGY From the Latin xzlebra— mgged, uneven, alluding to the nodulose dorsal surfaces. Neocirolana tricristata sp, nov. (Figs 3-5) MATERIAL EXAMINED HOLOTYPE: ô (3.4 mm), Gulf of Carpentaria, Queensland, 10°58.5°S, 141712'E, 3 Dee 1990, dredged 31,0m, S. Cook on FRY Southern Surveyor, QMW 18717, DESCRIPTION Male holotype. Body about 2.2 times as long as greatest width; maximum width al pereonites 3-6, margins sub-parallel, Cephalon without rostral point in dorsal view; anterior margin thickened, with submarginal anterior suture, with obscure longitudinal ridge along medial margin of eyes; eyes occupying 44% width of cephalon in dorsal view; ocelli distinct. Pereoniles 1-7 each with transverse suture, becoming laterally more prominent on pereonites 5-7; coxae 2-5 not prominent in dorsal view, ‘all coxae with prominent entire oblique canna; posterolateral angles of coxae 5-7 acute. Pleonites all visible, pleonites 2-5 with finely nodulose posterior mar- gin. Pleotelson with 'submarginal nodule at anterolateral angle; lateral margins obscurely thickened, dorsal surface with 3 distinct lon- gitudinal ridges consisting of median ridge flanked by further ridge on either side; posterior margin evenly rounded, with 4 spines set among PMS. Antennule peduncle robust, 2.5 times as long as width at article 2, articles 1 and 2 partially fused, article 3 about half as long as the combined lengths of articles 1 and 2; flagellum short, ex- tending to anterior of pereonite 2, with 9 articles bearing conspicuous spray of nesthetascs. Anten- na peduncle dorsoventrally compressed, article 1-3 short, articles 4 longest, 1.4 times longer than article 5; flagellum with about 9 articles, extend- ing ta postenor of pereonite 2. Frontal lamina pentagonal, about twice as long as wide, lateral margins weakly concave, with 2 submedian grooves, Mandible incisor namnw, about 50% maximal width, with distinet cusps; spine row with several setae and 3 spines; molar process short, with few setules, anterior margin with about 7 widely spaced spines: palp articles 2 and 3 with about 10 and 7 serrate spines respec- tively, Maxillule lateral lobe with 12 slender spines on gnathal surface, several of which ser- rate; medial lobe with 2 CP spines, Maxilla lateral and middle lobe with 3 and 4 setae respectively; medial lobe with 5 CP spines and 3 simple setae, Maxilliped palp articles lateral margins with few setae; endite with 1 coupling hooks, 3 short CPS, Pereopods robust, with few setae. Pereopod ! merus posterior margin with 2 blunt tubercular spines and | acute spine; posterior margin of carpus with 1 acute spine; propodus with | acute spine, and distal spine opposing dactylus base 38% length of dactylus. Pereopod 2 posterodistal margin of ischium with | tubercular pine, postenor margin of merus with 3 tubercular and 2 Jong acute spines; carpus with 2 acute spines at poslerodistal angle; propodus palm with | acute spine. Pereopod 7 with anterodistal angles of tschium, merus and carpus each with cluster of large spines, some of which are serrate; posterior margins with short simple acute spines at distal angles, ischium with 3 short acute spines along posterior margin, merus, carpus and propodus with 1 each, Pleopod 1 with both rami distally rounded, endopod about 3 times as long as wide, about 0.5 as Wide as exopod; exopod with ~20 PMS and spine al proximolateral margin, endopod with ~10 PMS, Pleopod 2 exopod and endopod with ~26 and 9 PMS respectively, appendix masculina distally curving slightly towards lateral margin, apex extending beyond distal margin of ramus by half of its length. Pleopods 3-5 with similar, ex- opods each with the distal margin rounded, each with transvers¢ suture and with about 25 PMS; endopods each with - 10, 6 and 0 PMS respective- ly. Uropod exopod about 0.9 times as long as endopod, straight, about 3 times as long as proximal width, lateral margin straight, irregular- ly notched with 4 short spines, each set in notch, and 7 PMS, medial margin with 2 spines among PMS, apex subequally bifid with cluster of setae and small spine within apical notch; endopod lateral margin indented with 4 PMS and 1 spine adjacent to apex, medial margin serrate, smoothly convex, with 3 stout peg-like spines set among PMS; apex subequally bifid with cluster of setae and two small spines within apical notch, NEW MARINE ISOPODS 47 FIG. 3, Neocirolana tricristata sp. nov. A, dorsal view; B lateral view; C, frons; D, antennule; E, antenna; F, antennule, showing spread of aesthetascs; G, maxillule, lateral lobe; H, maxilliped; I, maxilla; J, mandible; K, maxillule, medial lobe in oblique view. Scale = 1.0mm. 48 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4, Neocirolana tricristata sp. nov. A, pereopod 1; B, pereopod 1, dactylus, medial view; C, pereopod 2; D, pereopod 6; E, pereopod 7. NEW MARINE ISOPODS 49 FIG. 5. Neocirolana tricristata sp. nov. A-D, pleopods 1-3, 5 respectively; E, uropod. REMARKS This species is readily separated from others of the genus by the three prominent pleotel- sonic ridges and the antennular morphology. Neocirolana bicrista has two ridges on the pleotelson and a third weakly defined median ridge. N. tricristata has a maxilla and maxi- lliped that are similar to that of Cirolana, the maxilliped having the mediodistal angles of palp articles 2 - 4 somewhat produced and lobate, and provided with abundant slender setae. In contrast, in N. bicrista the maxilla has no large circumplumose setae on the medial lobe and has prominent recurved setae on the middle and lateral lobes; the maxilliped has cuticular setules along both margins of the palp, but few setae, and the mediodistal angles are scarcely produced and have only 1 or 2 short setae. The pereopods of N. tricristata differ from that of N. bicrista being more robust and in having a prominent spine opposing the dactylus base and a prominent secondary un- 50 MEMOIRS OF THE OUEENSLAND MUSEUM guis to the dactylus, both of these being minute in N. bicrista. The antennule flagellum has a dense mass of aesthetases, and is similar in form to a cal- lynephore (Lowry, 1986). The definition of a callynophore includes the total or partial fusion of the proximal articles (Lowry, 1986), and as the articles of N. tricristate are unmodified in thal way, nor are they particularly enlarged, the aesthetasc mass is not regarded as a cal- lynophore. ETYMOLOGY From the Latin crista = ridge, alluding to the three pleotelsonic ridges. Neocirolana excisa (Richardson, 1910) Cirolana excisa Richardson, 1910: 6, figs 4, 5; Nierstrasz, 1931: 156; Bruce, 1980; 127, 128. Neocirolana excisa, Bruce, 1981: 957; 1986: 207. figs 144, 145; Javed & Yasmeen, 1990: 72, table 1. MATERIAL EXAMINED ARAFURA SEA; Y, non-ovig, 10°32,6°S, 134916.6'E, 13 Oct 1992, 60m, R.W. Williams, NTMCr009778: â, 11*18.3'8, 129°24.0°E, 21 Nov 1989, 34, 7m, BRR, OMWI8710; 9, 11700.0'S, 132947.0'E, 30 Oct 1989, 18.3m, BRR, QMW 18712. NORTHERN TERRITORY: 4, Channel ls, Darwin, 20 Aug 1982, ex sponge 'A', lim, P. Horner, NTMCr005822; 3 9, 2 ovig, I non-ovig, Coral Bay, Port Essington, Coburg Peninsula, 1] Aug 1986, ex sponge, R. Williams, NTMCr004142; 2, non-ovig, north of Wessel Island, 16°32.5'S, 121°24.5'E, 17 Apr 1983, 40m, W. Houston, NTMCr002976. GULF OF CARPENTARIA: 2 4, far northern Gulf, Old, 10°04,1'S, 141724.4'E, 1 Apr 1989. dredged, 61.0m, BRR, QMW 19715. REMARKS Previously known from two records only, the species is not uncommon along Australia’s northern coasts from the intertidal to a depth of 61m; two records are from sponges. It has not been taken from coral reef habitats and may inhabit only inshore and offshore habitats with a high silt load. DISTRIBUTION Originally recorded (Richardson, 1910) from the Philippines, the species is now recorded from Australia's tropical northern waters from the Arafura Sea to Torres Strait at depths from 11 to61m. It has not been collected from tropi- cal eastern Queensland, despite collecting ef- forts in that region, and its range may not extend to that area, Neocirolana hermitensis (Boone, 1918) Cirolana hermitensis Boone, 1918: 592, pl. 31, fig. 2; Hale, 1925: 132. Neocirolana hermitensis, Bruce, 1981: 956; 1986: 202, figs 140, I41; Javed & Yasmeen, 1990: 72, table 1. MATERIAL EXAMINED TIMOR SEA: g, Oxley Island, 10759.U'S, 132948.8'E, 19 Oct 1982, low water spring level, ex Dardanus sp. A.J. Bruce, NTMCrD02282; 9, ovig, Hibernia Reef, 11759'S, 123722'E, 15 May 1992, 0- 0.5m, on reef flat in Turbo shell with Calcinus gaimar- dii, J, Short, QMW14232; 3 â, 2 ?, ovig, ‘Ann Millicent’ wreck, Cartier Reef, WA, 12°32.8'S, 123932.9'E, 4 May 1992, intertidal, in Bursa granularis shell, R. Williams & B. Russel, NTMCr009613; d, 9, non-ovig, North West Island, Ashmore Reef, 21 Sept 1987, intertidal, ex Dardanus, H. Larson, NTMCr005981. TORRES STRAIT: Juvenile, 10902'S, 142°31°E, 21 Apr 1974, 13m, Old Fisheries Service, OMWI17960; d, 9°56.2’S, 141955.5'E, 3 Apr 1989, 39m, dredged. BRR. QMW 18714; d, 10°22.4'S, 141730.37E, 4 May 1989. 65m, dredged, BRR, QMW 18716. REMARKS This species is regularly recorded from gastropod shells that are, or have recently been occupied by hermit crabs prior to collection, usually by species of Dardanus. Recently a specimen was collected by J.D. Shields at Bird Islet, Lizard Island in association with a female Dardanus scutellatus. The isopod left when the shell was cracked open to remove the crab, and immediately swam to the ovigerous host and was seen to be eating the eggs. The remains of eggs were also observed in its gut. It seems very prob- able, therefore, that Neocirolana hermitensis is an egg predator, a habit which has been recorded only for one species of deep-water lysianassoid amphipod (Bowman & Wasmer, 1984), but not previously for the Cirolanidae, nor any other marine isopod, DISTRIBUTION Most records are from offshore coral reef habitats around the Australian coast, with the exception of one record from inshore coral reefs at Pon Essington, NT (Bruce, 1986); from the Montebello Islands, WA, to Lizard Island, Qld, Australia. NEW MARINE ISOPODS 51 ACKNOWLEDGEMENTS I am indebted to Dr Jeffrey D, Shields (Virginia Institute of Marine Science, The College of Wil- liam & Mary, Virginia, U.S.A.) for providing the information on his observations of the dietary habits of Neocirolana hermitensis. I also thank Stephen Cook (OM) for delivery of specimens from the Gulf of Carpentaria, and P.J.F. Davie (OM) and A.J. Bruce (NTM) for the opportunity to examine material under their care. I thank Jørgen Olesen for his inking of the figures. LITERATURE CITED BOONE, P.L, 1918. Descriptions of ten new isopods. Proceedings of the United States Nationa! Museum 54: 591-603. BOWMAN, T.E. & WASMER, R.A. 1984. The deep- sea amphipod Paracyphocaris praedator (Gam- maridea: Lysianassidae) associated with the pelagic shnmp Oplophorus novaezeelandiae as an egg mimic. Proceedings of the Biological Society of Washington 97: 844-848, BRUCE, N.L. 1980. The Cirolanidae (Crustacea: Isopoda) of Australia. Heron Island and the Capricom Group. Bulletin of Marine Science 30: 108-130, BRUCE, N.L. 1981. Cirolanidae (Crustacea: Isopoda) of Australia: Diagnoses of Cirolana Leach, Metacirolana Nierstrasz, Neocirolana Hale, Anapsilana Paulian & Debouteville, and three new genera - Natatolana, Politolana and Car- tetolana. Australian Journal of Marine and Fresh- water Research 32: 945-966. BRUCE, N.L. 1986. Cirolanidae (Crustacea: Isopoda) of Australia. Records of the Australian Museum, Supplement 6; 1-239. BRUCE, N.L. 1993. Two new genera of marine isopod crustaceans (Cirolanidae) from Madang, Papua New Guinea. Memoirs of the Queensland Museum 31:1-15, HALE, H.M. 1925, Review of Australian isopods of the cymothoid group, Part I. Transactions of the Royal Society of South Australia 49: 128-185. HOLDICH, D.M., HARRISON, K, & BRUCE, N.L. 1981. Cirolanid isopod crustaceans from thc Townsville region of Queensland, Australia, with descriptions of six new species. Journal of Natural History 15; 555-605. JAVED, W. & YASMEEN, R. 1990, A new species of cirolanid isopod of the genus Neocirolana trom Pakistan with a review of the genus, Crustaceana 58: 67-73. LOWRY, K.L. 1986. The callynophore, a eucaridan/peracaridan sensory organ prevalent among the Amphipoda (Crustacea). Zoologica Scripta 15: 333-349. MARTIN, J.W. & FELDER, D.L. 1984. Rediscovery and redescription of Cirolana obtruncata Richardson, 1901 (Peracarida: Isopoda: Cirolanidae) from the east coast of Mexico. Proceedings of the Biological Society of Washington 97: 30-34, MULLER, H.G. 1993. Cirolanidae of the genera Calyp- tolana, Cirolana, Neocirolana, Colopisthus and Excirolana from the Santa Marta arca, Caribbean Sea of Colombia (Crustacea: Isopoda: Cymothoida). Zoologischer Anzeiger 230; 191- 225. NIERSTRASZ, H.F. 1931, Die Isopoden der Siboga- Expedition. 3. Isopoda Genuina. 2. Flabellifera. Siboga Expeditie Monographs 32c: 123-233. RICHARDSON, H. 1901. Keys to the isopods of 1he Atlantic coast of North America, with descriptions of new and little-known species. Proceedings of the United States National Museum 23: 493-579, RICHARDSON, H. 1910. Marine isopods collected in the Philippines by the U.S. Fisheries steamer Al- batross in 1907-1908. Department of Commerce and Labor, Bureau of Fisheries Document 736; 1-44, STEBBING, T.R.R., 1900. On Crustacea brought by Dr Willey from the South Seas. Pp.605-690. In Wil- ley, A. (ed.), ‘Zoological resulls based on matenal from New Britain, New Guinea, Loyalty Islands, and elsewhere, collected during the years 1895, 1896 and 1897", Vol 5(33). (University Press: Cambridge). 52 MEMOIRS OF THE OUEENSLAND MUSEUM MALE COMBAT IN THE BLACK WHIP SNAKE, DEMANSIA VESTIGIATA. Memoirs of the Queensland Museum 37(1): 52, 1994:- Male combat is well known in Australian elapids and boids (Shine, 1991), lt has been reported in many elapid genera - Austrelaps, Cryptophis (now Rhinoplocephalus), Deman- sia, Hemiaspis, Notechis, Oxyuranus, Pseudechis and Pseudonaja. These genera are characterised by male size superionty. Male combat is unknown in in which females are larger than males e.g. Acanthaphis and Vermicella, (Shine & Allen, 1980; Shine, 1991), Shine (1978) first reported this sexual combat correlation and has commented on its adaptive si (Shine, 1981, 1991). He summarised this as a ^,,.. male strategy for (i) forcing rival males to flee from the vicinity of a receptive female .. (ii) physically “carrying” rival males away from a female ... (iii) defending against homosexual courtship .. and (jv) gaining access to food ,..' (Shine, 1981), We report recent observations of male combat in Demansia vestipiata and present photographs of combat in this species. On June 13, 1994 two snakes were encountered | 1km south of the Starcke R., NEO (14751'S, 145702'E), in a grassy open forest dominated by Melaleuca viridifolia. (This is typical habitat for D. vestigiata on Cape York Peninsula), At about 4.00pm they were found in the open, in the wheel cut of the unsealed track from Cooktown to the Starcke R. They were observed and photographed (Fig. 1 A-C), but were not ex- amined to check sex. However, as mating has been observed by one of us (JC) to be relatively passive in this species; as the snakes were observed very closely (from a distance of about 5m) and were not copulating; as they were large (with TLs of approximately 70cm); and their behaviour was very similar to that reported for male combat in other elapid species, we are confident both snakes were males. We watched their ritual for close to 45 minutes. They moved, as a pair, about 20m along the road - all the time twisting, coiling, raising their forebodies both together and separately, and then falling back to begin the sequence anew, Male combat in D. vestigiata has been recorded before, briefly. Shine (1980) noted an observation by Neil Charles, near Cooktown , 22 September (year not cited). '.., The two males were plaited together in typical ritualised combat ... were captured and their sex confirmed ...'. It has also been photographed before by J. Stanisic, D. Potterand J. Chaseling, 21 September, 1988, near the Captain Billy Landing turn-off, Cape York Peninsula. Their photograghs have been lodged in the Queensland Museum Photographic Index (NP119-21). Demansia vestigiata is a species in which the males exceed females in size, Our observations conform with several strategies suggested by Shine (1981) toexplain this behaviour. The activity carried both snakes far from their starting point in the short penod they were observed closely. Shine (1980) observed *... no clear seasonal pattern (in reproduction) is evident ìn D- atra (now D. vestigiata) ...'. Our Winter obser- vation, considered with the only others known (in Spring) is consistent with this. Literature Cited Shine, R. 1978, Sexual size dimorphism and male combat in snakes. Oecologica 33: 269-278. 1980, Ecology of eastern Australian Whipsnakes of the penus Demansia. Journal of Herpetology 14(4); 381- 89. 1981, Mating and male combat in Australian Black snakes, Pseudechis porphyriacus, Journal of Herpetology 15(1): 101-107. 1991. ‘Australian snakes, A natural history’. (Reed Books Pty Lid: Sydney). dd Ag FIG. 1. A-C, Demansia vestigiata, males in combat, Starcke R., NEO. Shine, R. & Allen, S. 1980. Ritual combat in the Australian Copperhead, Austrelaps superbus (Serpentes, Elapidae). Victorian Naturalist 97: 188-190. J.A. Covacevich, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia; L. Roberts, Shipton's Flat, via Cooktown, Queensland 487 1, Australia; I. McKinna, James Cook Museum, Cooktown, Queensland 4871, Australia; 10 August 1994. ae oe > * y REDESCRIPTION OF NEPHRURUS ASPER GUNTHER, AND DESCRIPTION OF N. AMYAE SP. NOV. AND N. SHEAI SP. NOV. PATRICK J. COUPER AND ROBERT A.M. GREGSON Couper, P.J. & Gregson R.A.M. 1994 12 01; Redescnption of Nephrurus asper Günther, and description of N, amyae sp, nov, and N. sheai sp. nov, Memoirs of the Oueensiand Museum 37(1):67-81. Brisbane. ISSN 0079-8835. Three allopatric species, clearly defined by morphology and colour pattem, comprise the N, asper species complex. N. asper is confined to Queensland (10°53°S-25°21'°S), and has a distinctive, boldly banded colour morph occurring on Cape York Peninsula (extending south to Mt Surprise, 18°21'S); N. amyae sp. nov. to the south-central Northern Territory and N, sheai sp. nov. to the Kimberley region of Western Australia and the Arnhem Escarpment of the Northem Territory. All three species occur in open woodland habitats where they feed ona wide range of arthropod species. Males are in reproductive condition during the summer months and gravid females are present in the population from mid-summer to autumn for N. asper and N. amyae. N. sheai may be sexually active throughout the year. [_] Gekkonidae, Nephrurus, systematics, new species, distribution, open woodland, diet, reproduction, Patrick Couper, Oueensland Museum, PO Box 3300, South Brisbane, Oueensland 4101, Australia; Robert A.M. Gregson, Department of Psychology, Australian National University, Canberra, Australian Capital Territory 0200, Australia; 10 June 1994. Nephrurus asper was described from a single specimen (BMNH1946.8.23.34) from Peak Downs Stn, mideastem Queensland (22°56'S, 148°0S"E). Subsequently, specimens matching the type description have been recorded from most of northem Australia (Cogger, 1992). While the distinctive morphology of N. asper has ensured its nomenclatural stability (Cogger et. al., 1983), several authors have commented on geographical variation within the species. Storr (1963) noted differences in tail length between specimens from Western Australia and Queensland. Bauer (1990) observed that specimens from the south-central Northern Ter- ritory were larger than those in other parts of the range. Wilson and Knowles (1988) recognised the hroad-banded form of N. asper from Cape York Peninsula as distinct from other populations of this species, An examination of all N. asper held in the collections of major Australian museums revealed that the species, as currently recognised, could be divided into four geographically distinct groups (Amhem Escarpment, NT & Kimberley region, WA; south-central NT; Queensland & Cape York Peninsula) on the basis of colour, pattern, spinosity, and other scale characters. Pat- tem and spinosity have been used widely in gek- konid revisionary work (Couper et, al., 1993; Storr, 1988, 1989). They have been found to be useful in delineating the species in the N. asper complex, in a way which conforms closely with Statistical analyses. The current study is augmented by Kol- mogorov Smirnov Two -sample Tests and multi- variate discriminant analyses. These analyses, performed using body measurements and quan- titative scale counts only, provide an independent assessment of the morphological identity of the four geographic groups identified on colour pal- tern and/or degree of spinosity. The statistical analyses were conducted by one of us (RAMG) and the taxonomic analysis was conducted by the other (PJC) All meastirements Were taken using Mitutoyo electronic callipers. Abbreviations for body measurements arc as follows: snout-vent length (SVL); axilla to groin (AG); tail, tip to constric- lion at base (T); tail, tip to vent (TV); knob width (KW); forelimb, axilla to tip of longest digit (L1); hindlimb, groin to tip of longest digit (L2); neck length, axilla to posterior edge of ear (NL): head length, tip of snout to dorsal/anterior margin of ear (HL); head width, measured between ear openings (HW), head depth, from above eyes to ventral margin of maxillae (HD); snout, tip to anterior margin nf orbit (S); eye to ear, posterior margin of orbit Io anterior margin of ear (EE). Abbreviations for scale counts and caudal annuli are as follows: number of granular scales in direct contact with dorsal edge of rostral scale (R); number of interorbital scales counted across the narrow, bony bridge between the orbits, includes the enlarged series of scales on either side (I) number of supralabial scales counted to exclude granular scales towards angle of mouth (SL); 54 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1. Differentiation between groups using a KS2 Procedure 4^ from the 6 possible comparisons he- tween the 4 samples. N is the number of contrasts with KS2d y? statistic >6. The .05 level for y? is about 5.99, for the 01 level is about 9.21, forthe ,001 level is about 13.81. | GROUPS lee laso faos [eoo oar jaue mv fioo [sei [azs [sss Jose Iai [kw |650 [ra [oa [a30 faio [oas loso Jos | ea — EAA has foe p lem lan lr» Tie Tsar [se | | uw [o [803 [133 [7.16 [oai [563 | E ees ls pa fpu |2aa [920 [371 |sz | |NL [632 A 17 me me ri [za [zas fada aciem heata is aa | izi E a 0.79 30.13 3.79 9.45 a [issa |352x [6.04 [7.42 |020 [zga | 285 [732 m pa on ie las ne lim. ac Josz [23s [nes [137 [1.11 [moe | im le iy 4: de [al number of infralabial scales counted to exclude granular scales towards angle of mouth (IL), and number of caudal annuli counted between basal constriction and dista] knob (TA). OTHER ABBREVIATIONS USED: Queensland Museum (QM), Australian Museum (AM), Museum of Victoria (NMV), South Australian Museum (SAM), Western Australian Muscum (WAM) and the Northem Territory Museum (NTM), Cape York Peninsula (CYP), mideastern Queensland (MEQ), Queensland (OLD), North- ern Territory (NT), Western Australia (WA). Reproductive and dictary data were compiled by examination of specimens in which the body cavity had already been opened. STATISTICAL ANALYSES The following data analyses have been per- formed, using complete records on 149 animals. Some are infernal checks on consistency and are hence not reported in full here. The characters used in these analyses are the body measurements and scale counts defined above. Group | = the Arnhem Escarpment (NT) & Kimberley region (WA), group 2 = OLD, group 3 = south-central NT, and group 4 = CYP, 1. The Kolmogorov Smirnov Two-sample Test (KS2) (Kolmogorov, 1941; Smimov, 1939) on each of the 6 pairwise contrasts between groups on each of 17 characteristics, giving a total of 10 F values each with 2 d.f. This is a nonparametric procedure, using minimal prior assumptions about the form of frequency distributions of measurable characteristics (Walsh, 1965). It indicates which characteristics differentiate better than al some given probability level, if the characteristics are independent. This is not strict- ly true a priori, and hence conservative prob- ability levels may be used in deciding whatresults to accept. Tt also will fail to detect second-order measures, such as ratios or differences of charac- teristics, which can be non-linear and of use, unless these arc built into the data. The measures in mm have been divided by SVL, those based on scale counts have not. Bilateral measures have been pooled (Table 1), Discriminant analyses have also been run because they can utilise linear (but only linear) combinations of characteristics, and hence may separate groups which are not distinguished on single characteristics (Table 1). The discriminant analyses do, however, make some distributional assumptions approximating to normality and homoscedascity across groups. The role of the KS2 is also involved with the question of what characteristics 10 use in the discriminant analysis, if not all the characteristics which have heen measured are used. 2. The correlation matrix on the characteristics over the whole 149 complete records used. This showed that the rank of the data is much less than 17; this indicated, as expected, that there is redun- dancy in the morphological data. 3. Four discriminant function analyses (Rao, 1952): 3.1. Using all the characteristics, on raw measures, 2,2. Using a stepwise discnmunant analysis, on raw measures. 3,3, Using all the characteristics, on measures divided by SVL. 3.4. Using a stepwise discriminant analysis, on measures divided by SVL. The stepwise procedures are inbuilt to the SPS5x program used; because of the intercorrela- tions noted under part (2) above, the variables retained in a stepwise analysis will depend on those iniercorrelations and on the relative sizes of the four groups, but will be analogous to the DESCRIPTION OF NEPHRURUS 55 pOoOrra sect 2 Function 1 -4 -2 0 Group Centroids Group 4 Group 3 Group 2 Group | 2 4 6 FIG. 1.Plot of the individuals and group centroids on the 1st two canonical variates from the discriminant analysis. Group 1 = the Arnhem Escarpment (NT) & Kimberley region (WA), group 2 = QLD, group 3 = south-central NT, and group 4 = CYP. characteristics with larger y” values in Table 1. We present tables derived from (3.4) here; as there are only 11 animals in the smallest group, the number of variables in the discriminant analysis has to be reduced to below 10. The stepwise procedure is the most appropriate for this, retaining only those variables with dis- criminative power; it showed by F-tests that only eight variables were legitimately retained in the analysis. Information for the identification of new specimens when allocation is based on body measurements and scale counts only is not in itself sufficient (Tables 2, 3). Allocation to species must also take into consideration colour pattern and degree of spinosity. The relative values of the coefficients (Table 3) are of use, they are weights to multiply with corresponding measurements, and then sum the products and add the constant term in the column. As addition- al specimens become available it is preferable to incorporate them in a extended discriminant analysis, thus yielding new weights. The efficien- cy of allocation decisions using this discriminant analysis is summarised (Table 4); it will be noted that the relatively poor separation of group 2 (QLD) and group 4 (CYP) is compatible with the results from the KS2 analysis. RESULTS The plot of individuals and group centroids on the first two canonical variates from the dis- criminant analyses (Fig. 1) gives a picture com- TABLE, Standardised discriminant function coefficients and correlations with disenminant functions from the stepwise analysis with three canonical functions. Ic = coefficients for the first discriminant, Ir = its correlations with discriminating Variables, and so on, ferar [te fie |n |n _]me_|mr | mw [omw [a2 [su fars [706 [as | | 1453 470 [1.234 |-201 rı [32 |an [ae [i02 [a72 | m as us [s98 [a2 |-207 [ioi | IR [ais |20 [nia [a37 [ios liso | [se _ [sea [aeo |.367 [ase [inzi |122 | [TA |-1.750 | [ry dhs han his [2m Jins hz | patible with Tables one and four. Group 2 (OLD) and group 4 (CYP) are not significantly separable on external morphology, The recognition of group 4 was based on colour-pattern. The unifor- mity in morphology between groups 2 and 4 dovetails well with preliminary genetic data which show a low level of divergence between a broad-banded specimen from Heathlands, CYP and a specimen from the Capella region of MEQ (Couper & Donnellan, unpubl. data), All OLD material examined in this study, with the excep- tion of the broad-banded specimens from CYP, is consistent with the holotype of N. asper (BMNH14946.8.23.34) in both colour-pattern and scalation (see Remarks for W, asper) and is here assigned 10 this taxon, The broad-banded CYP specimens are regarded as a geographically dis- tinct colour morph of N. asper. The statistical analyses show that group | (Amhem Escarpment, NT & Kimberley region, WA) and group 3 (south-central NT) are morphologically distin- guishable from QLD and CYP combined as a single group (Table 1, Fig. 1) and also from each other, Groups | and 3 are here described as new, allopatric taxa. The recognition of these taxa is further supported by preliminary genetic data (Couper & Donnellan, unpubl, data), The ‘Atlas of Australian Resources’ (Anon, 1952-69) shows reasonable conformity between the distribution of the N. asper species complex and the occurrence of upland country and heavy, often stony soils. However, the zoogeographical implications of this distribution can not be as- sessed without first establishing a phylogeny for the genus and determining the dates of speciation events. Presently, there is no fossil record for Nephrurus and biochemical studies are incom- plete. MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS Nephrurus asper Gunther, 1986 (Fig. 2A,B; Statistical analyses groups 2,4) MATERIAL EXAMINED QUEENSLAND: QOMJ22179, OMJ24921, Bamaga, Cape York Peninsula (10°53'S 142°24°E); OMJ57652, 6.5km E of Heathlands airstrip (11°44,33'S 142°38.28"E); OMJ54644, Heaihlands road, 1km from main Cape road junction (1146'S 142'40'E); OMJ40165, Weipa to Aumikun, mid-way, N Cam Beagle Airstrip (13705'S 141°57'E); QMJ738, Kingsborough (16°55'S 145702'E), OMJ4960, Bul- lock Ck, via Cairns (16'55'S 145°46'E); OMJ57993, OMJ5&850, Elizabeth Ck, nr Bulleringa NP, N of Mt Surprise (I&'DD'21"S 145°S9°50"E); OMJ2125, ? Northern Rivers; OMJ44689, New Moon Stn (19 12'S 145744"E); OMJ3443, Charters Towers, Black Jack (2008'S, 1467137E); OMJ42015, Hughenden, approx 2km W of lona-Redeliffe Stn Hotnestead (20°59'S 145.33 B); OMJ44948, Egera Sin (20/59'S 14605 E); J143, Natal Downs (21703'S, 146°09'E); OMJ4525-6, Kuridala, S of Cloncurry (2171775 140 30/E); QMJ15565, 80km SW of Mackay (21 39'S 14&'42'E), OMJ31976, OMJ35040, Winton (22°23'S 143*02'E); OMJ5727, Lucknow Sin, W of Winton (22'43'5 140°55°E); OMJ1355, OMJ3552, Clermont (22 50'S 147°338"E); QMJ23699, Cork Sin via Winton (22 56'S 14218'E);, OMJ46720), Aramac Range, Winhaven Sin (22:5T'S 1457417E); OMJS608B, Capelia, 20km N on Clermont pond (22°59°S 147°53°E), OMJ36795, Moonggoo (23°10'S 148704'E); QMJ3767, Rock- hampton, Dalma, Waroula (23°20'S 150/32'E); OMJ9912, Longreach (23°26'S 144°|5°E), OMJ7983, Mary Vale Stn, Yalleroi via Blackall (23°35'S 146 58'E); OMJ36794, E of Comer Sin (23°37'S 148 740" E), OMJ342015-08, half way up the Blackdown Tableland escarpment (23°37'S 14910'E); OMJ8080, Bogantungan (23739', 147°18"}; QMJ741, Diamantina Lakes (23°46'S 14170R'E); QMJ31545, Winton fossil site (23°50°S 142^15'E); QMJ9132, Blackall (24°26 S 145°28"B); OMJ43356-7, Moura, 4km S of junction of Theodore & Gibihi Roads (24/38'S 149*59'F): OMJ7878, OMJ10526, Jundah (24°50°S 14304'E); QMJ4540, Dawson Valley, Castle Ck (24 750'S 150°20° E); OMJ6011, Weathersfield?; OMJ742, No locality data; AMR 128183, Batavia Downs, Wenlock R(12°S0'S 142740'E); AMR128796, Caims (16°55"S 145°46"E); AMR31773, Mi Fox nr Ingham (18°19°S 145°51"E); AMR113852, approx, 14.5km W of ‘Mt Cooper’, nr Si Paul's vine scrub (20°31°S 146755'E); AMR 14183, 210km S of Charters Towers (21°38"S 146°55'E); AMR63065, approx. 48km NW of Cler- font (22°25°S 147 23'E); AMR107703 N of Clermont (22/50'S 147'38'E), AMR120094, Mayne Junction Hotel, approx. 40km N of Diamantina Lakes Sin (2233'S 141°22'E); AMR15107, Yamala (23°35'S 148°22"B); AMR113116, Diamantina Lakes Stn, SW of Winton (23°41'S 141°11'E), AMR55786, Duaringa (23°43'S 149°40'E); AMR110544, AMR110562, DESCRIPTION OF NEPHRURUS 57 camp, t4km NE of Scott's Tank, Diamantina Lakes, NW of Windorah (23°45'S 141"40'E); AMR130721- 2, AMR130726, 20km W of Moura on Bauhinia Downs toad (24°37'S 149*47'E); NMVDT-D182 (Donald Thomson Collection), Lower Archer R (13°30'S 142"00'E); NMVD7527, Mi Cook, Cook- town (15728'S 145°15"E); NMVD2040, Queensland; SAMR14023, 22km NW of Cooktown (15°25'S 145704'E); SAMR12594, Cooktown (15°28'S 145715'E), SAMR15791, Charters Towers (20° 05'S 146°16°E); SAMR1771, Mt Coolon (21°23'S 147720'E); SAMR42600, 6km N of Diamantina Stn (23°43'S 141°08'E); SAMR42601-02, 4km N of Diamantina Stn (23°44'§ 141°08'E); SAMR42603, 85km W of Windorah (25°21'S 141°S0°E); WAMR55475-7, 16km W of Charters Towers (20 08'S 146°07'E); WAMR55552, WAMR55601-2, Fermoy Homestead, 88km S$ of Winton (23°10'S 143702'E); NTMR1144, Clermont (22°50'S 147°38'E); NTMR266, Rockhampton (23°22'S 150°32'E). DIAGNOSIS N. asper is medium-sized and moderately spinose. lt is distinguished from N. sheai by the colour pattern of its digits (digits unbanded ws digits strongly marked with altemating bands of brown and white); from N. sheat and N. amyae by its smaller size (max SVL 114mm vs 121mm X, sheai, 135mm N. amyae). lt is further distin- guished from N. amyae by the spinosity of the its rump and thighs (moderately spinose vs extreme- ly spinose). The arrangement of the basal scales surrounding the tubercules on the rump and thighs also separates N. asper and N. amyue. In N. asper the basal scales are uniform in size and less than half the height of the central scale. In N, amyae the basal scales are irregular in size and in most specimens examined, some of the basal scales are greater than half the height of the central scale. In the KS2 analysis N. asper (excluding broad- banded CYP specimens which are not sig- nificantly separable from other QLD populations of this species) and N. amyae show significant separation of the shape of frequency distributions for, KW, HW. S, NL, R, & 1. N. asper (excluding broad-banded CYP specimens) and N. sheai show significant separation of the shape of fre- quency disinbutions for; T, TV, KW, TA, NL, SL, IL & I (Table |). DESCRIPTION SVL(mm): 47-114 (N 74, mean 84.1). Propor- tions (WSVL): - LI - 36,8-47.8 (N 74, mean 43.0), L2 - 43.2-55.6 (N 73, mean 49.0), AG - 34.4-54.2 (N 74, mean 44.4), T- 11.4-22.2 (N 66, TABLE 3. Fisher linear discriminant function coeffi- cients, Animals are assignable to the species with the highest function, given their measurements corrected for SVL. fp Groups — mh bn [yw 84113 [sarao [78500 [s8174 a asas faisais pssi | 95.68 314.64 E Y IR [3754 |-12.84 [en oan [ae Jin | [Ta |-13506 |-837 |-28.78 [23392 | rv sas fare [sais |4&s7 | ei nm (nm hee Jas | mean 16.4), TV - 15.7-27.4 (N 66, mean 22.7), HL - 27.9-34.8 (N 73, mean 31.3), HW - 23.9- 30.6 (N 74, mean 27.7), HD - 10,7-15.8 (N 70, mean 13.8), S -9.2-12.7 (N 74, mean J1 1), EE - 10.0-15.0 (N 73, mean 12.3), NL- 10.4-21.1 (N 72, mean 15.5). Head. Large and deep, covered with small, round to hexagonal, juxtaposed scales. Posterior- ly bearing scattered tubereules intermixed with the smaller scales. Each tubercule consists of a high central scale, circled by a ring of smaller basal scales, Tubercules most promment on the nape. Dorsal skin co-pssified with skull. Nostril small, opening upwards and backwards. Eye large with vertical pupil. Ear vertically elongate, tympanum deeply recessed. Rostral scale small, with 8-17 (N 77, mean 12.5) scales in direct contact with dorsal edge, Inierorbital scales 7-12 (N 76. mean 8.9). Supralabial scales 11-17 (N 152, mean 13.5). Infralabial scales 10-17 (N 152, mean 13.5). Neck, Broad, less than half width of head. Body, Stout, dorsal and ventral surfaces with small granular scalation. Granular scales inter- mixed with larger conical tubercules on dorsal and Jateral surfaces. Mid-dorsal tubercules small, TABLE 4. Classification results for 149 animals. Actual group by HAY H group abi alc 58 MEMOIRS OF THE QUEENSLAND MUSEUM rump tubercules moderate. Basal scales sur- rounding rump tubercules uniform and less than half as high as the central scale. Rump tubercules larger than flank tubercules (45% material ex- amined) or subequal to flank tubercules (55% material examined). Rump tubercules smaller than nape tubercules (51% material examined)or subequal to nape tubercules (49% material ex- amined). Lower flanks and ventral surface with scattered, slightly raised rosettes which vary be- tween being pronounced or barely discernible, Limbs. Long and slender, bearing enlarged Wubercules on dorsal surfaces. Tubercules on thighs largest, with uniform basal scales less than half as high as the central scale. Digits short, cylindrical, undilated distally and terminating in a non-retractile claw. Third toe on hindlimb longest. Tail. Short, moderately depressed, constricted at base and terminating in a globular, kidney- shaped knob (KW 17.8-38.896T, N 66, mean 29,1), Usually four, rarely five or six longitudinal rows of tubercules present on dorsal and lateral surfaces. Uniform basal scales surrounding caudal mbercules less than half as high as the central scale. Caudal annuli 9-12 (N68. mean 10.4), PATTERN Body, In spirit, mid-dark brown, or mid-dark grey on dorsal surface. Many specimens have a broad black band on the nape, Seven - eight, rarely six, narrow irregular crossbands (crossbands broad and continuous in CYP specimens) present between head and hindlimbs. Crossbands range from obscure to bold and are continuous or broken into a series of spots, with each spot covering a tubercule. Fine black lines form reticulations which become obscure in larger specimens and can be difficult to see in darker individuals. Ventral surface cream, some- times faintly marbled with grey or brown. Limbs, Lacking irregular crossbands distally, Digits unbanded or only obscurely banded. Tail. As for body, with an obscure, broad, pale crossband on proximal half. Head. Sometimes slightly lighter than body, Covered dorsally and laterally with fine black reticulations, which form an intricate pattern, These reticulations are mosi prominent in juveniles and may fade in larger specimens. DISTRIBUTION From the drier parts of coastal MEQ through central and western districts, Also present on CYP (Fig, 3). The bocality given for QMJ2125 is '?Northern Rivers’, which usually refers Io an area in north- em NSW, The following entry in the register (OMJ2126) is a specimen of Saltuarius cornutus from the same locality, Neither of these species occur in NSW, nor are they sympatric elsewhere. N. asper typically occurs in dry open country while 5. cornutus is confined to rainforest habitats. Either the locality *?Northern Rivers’ should be regarded as an error or it refers to a region (unknown to the authors) of southern Cape York Peninsula where both rainforest and dry forests occur. HABITAT N. asper favours rocky substrates in open woodlands (Steve Wilson, Tim Hawkes, pers. comm.). lt has been recorded also from stony or compacting soil plains, where sheltenng sites are available (Ehmann 1992; Couper, unpubl, data); from low heath at Heathlands, CYP, and Darwin Stringybark open forest, at Aurukun (Cameron & Cogger, 1992). REPRODUCTIVE NOTES Clutch size - two eges. Egg-laying occurs in mid-late summer, Female WAMR55476 was gravid during late February, The two oviducal eggs measured 27.1 X 13.8mm and 25.2 X 14.0mm; OMJ54644 had a greatly enlarged vitel- logenic follicle in each ovary at the time it was killed in early February, 1992. The largest of these measured 9.67 X 7.28mm. A gravid female was brought to Wild World for identification in mid-April (T. Hawkes, pets. comm.). Male WAMRS55552 was in reproductive condition (epididymis enlarged and turgid) in late Novem- ber, DIETARY NOTES The gut of QMJ56088 contained two cricket legs (Orthoptera: Gryllidae). OMJ36795 con- tained a medium-sized centipede (Chilopoda: Scolopendridae) and an ant. OMJ44689 con- tained fragments of a medium-sized spider, and a medium-sized cockroach (Blattodea: Blattidae). Two faecal samples were collected at Heath- lands, CYP, in late January 1992. The first sample contained a large spider, a small centipede, a large coleopteran larva and small to medium-sized cockroaches (Blattodea). The second sample DESCRIPTION OF NEPHRURUS 59 FIG. 2. Nephrurus asper. A, Moura, QLD; B, Broad-banded form, Heathlands, Cape York Peninsula, QLD. (Photographs: S. Wilson). 60 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. Distribution of the Nephrurus asper species complex. N. amyae (diamonds), N. asper (circles), N. sheet (squares). contained legs from a medium-sized grasshopper (Acrididae), fragments of a large scorpion (Urodacus sp.) and fragments of a small phasmid. REMARKS A clear photograph of the holotype of N, asper (BMNH1946.8.23.34) was provided by Bauer, (1990), The narrow, irregular dorsal crossbands and moderate, uniform rump scalation of this specimen are consistent with the pattern and scalation of OMJ56088, a specimen recently col- lected only 14km from the type locality. Further, these specimens are morphologically/phenotypi- cally consistent with populations of spinose Nephrurus occurring in Queensland between 18°19'S & 25921'S. These are regarded here as N, asper. The broad-banded form on CYP is also included in N. asper because it cannot be sig- nificantly separated from this species by scalation features or body proportions, and preliminary genetic data show a low level of divergence be- tween a specimen from Heathlands, CYP and a specimen from the Capella region of MEQ (Couper & Donnellan, unpubl, data) The broad-banded CYP colour form is narrow- ly separated from the narrow-banded form at the southern limit of its range. Tim Hawkes (Cape York Herpetological Society) has provided colour transparencies of a broad-banded specimen from 6km east of Almaden (17721'S, 144°43’E) and a narrow-banded specimen from approximately 87km south, near the junction of the Kennedy and Gulf Redevelopmental High- ways (18°09, 144741”). This narrow-banded specimen is approximately 75km NE of the most southerly museum record of the broad-banded form (OMJ57993). Clearly, the range of both colour forms overlap latitudinally but, to date, they have not been found in sympatry. Nephrurus amyae Couper sp. nov. (Fig. 4; Statistical analyses group 3) Nephrurus asper, Cogger, 1992 (in part). MATERIAL EXAMINED HOLOTYPE: AMR 104458, Winnecke Goldfields, Gar- den Sin, N of Alice Springs, NT (23°17'S 134725'E). Found in mine tunnel, February, 1982, Don. M. Robin- son. PARATYPES: NORTHERN TERRITORY: QMJ51650, 15km SE of Glen Helen (23°31'S 132°21’E); OMJ53650, Hugh R, nr Alice Springs (23°49’S 133722'E); AMR49716, Mt Gillen, Alice Springs (23742'S 133°48'E); AMR50542, Alice Springs (23:42'S 133°53°E); AMR11965, Mt Gillen, Alice Springs (2343'S 133 48'E); AMR90198, Jay Ck, NT (23*50'S 133°29°E); AMR 10371, between Hale and Plenty Rivers, Central Australia (24725'S 136°10'E); NMVD538, Barrow Ck, (21/32'S 133753'E); NMVD51910-21, 28.9km S of Barrow Ck (21°45'S 133*40'E); NMVD55377, nr Alice Springs (23°42'S 133°52°E); NMVD12684-5, Central Australia; SAMR1892, Barrow Ck (21°32'S 133*53'E); SAMR38837, Mt Zeil (23°25'S 132°25'B); SAMR40561, approx 35km SE of Kings Ck Homes- tead (24*26'S 131°49 E); SAMR30523, No Locality data; NTMR441, Barrow Ck (21°32'S. 133753'E); DESCRIPTION OF NEPHRURUS 6) R14096, Mt Riddock Stn, Dulcie Ra. (22°30'S 135°25'E); NTMR33717, Winnecke Goldfields (23702'S 134°23°E); NTMR33716, Bond Springs Stn FS 133°52°E); NTMR12377, 16km N Alice Springs (23°34'S 133°53°E); NTMR33720, Alice Springs Hills, 19 Mile Bore (23°35'S 133°52'E); NTMR33722-3, Alice Springs, Wigleys Turnoff {23°36’S 133753'E); NTMR12380, 9km N Alice Springs (23°37'S 133°53°E); NTMR33721, Alice Springs, Charles R (23°39'S 133°51'E); NTMR5586, NTMR5969, NTMR1499}, Alice Springs (23°42'S 133°52'E); NTMR33724, NTMR33736, Alice Springs (23°42'S 133'53'E); NTMR5383, Alice Springs Power House (23°42’S 133'55'E); NTMR5466, Alice Springs, Mt Gillen (23°43'S 133 /48'E); NTMR 2458, Alice Springs, Emily Gap (23°45'S 133757'E); NTMR33719, Hermannsburg, Finke Crossing (23758'S 132°46"E); NTMR33715, Erichnuff Ra. (23759'S 132°38’E); NTMR33728, No data. DIAGNOSIS N. amyae is the largest and most spinose mem- ber of the N. asper species group, lts large size distinguishes it from other members of this group (max SVL 135mm vs 114mm N. asper, 121mm N. sheai). Ilis further distinguished from N. asper and N, sheai by the spinosity of the its rump and thighs (extremely spinose ys moderately spinose), The arrangement of the basal scales surrounding the tuhercules on the rump and thighs also separates N. amyae from N. asper and N. sheai. In N. amyae the basal scales are irregular in size and in most specimens examined, some of the basa] scales are greater than half the height of the central scale, In X. asper and N. sheaf the basal scales arc uniform in size and less than half the height of the central scale. in the KS2 analysis N. amyae and N. asper (excluding broad-banded CYP specimens which are not significantly separable from other QLD populations of this species) show significant separation of thc shape of freguency distributions for; KW, HW, 5, NL, R & I. N. amyae and N. sheai show significant separation of the shape of frequency distributions for; KW, HW, HL, S, NL, SL, IL, R & I (Table 1). DESCRIPTION SVL(mm): 50-135 (N 40, mean 100.3). Propor- tions, (SVL): - Ll - 38.1-52.4 (N 40, mean 42.7), L2 - 42.5-56.1 (N 40, mean 48.4), AG - 35.4-53.5 (N 40, mean 45.1), T - 12.3-17.5(N 37, mean 15.0), TV - 15.2-25.2 (N 37, mean 21.3), HL - 27.7-34.2 (N 38, mean 30.4), HW - 23.9- 30.6 (N 38, mean 26.4), HD - 10.5-15.1 (N 38, mean 13.2), S - 9.1-12.9 (N 39, mean 10.5), EE - 10.2-14.8 (N 39, mean 12.9), NL - 10.4-19.9 (N 37, mean 14.7). Head. Large and decp, covered with small, round to hexagonal, juxtaposed scales. Posterior- ly bearing scattered tubercules intermixed with the smaller scales, Each tubercule consists of a high central scale, circled by a ring of smaller hasal scales. Tubercules most prominent on the nape. Dorsal skin co-ossified with skull. Nostril small, Opening upwards and backwards. Eye large with vertical pupil. Ear vertically clongate, tympanum deeply recessed, Rosiral scale small, with 7-18 (N 39, mean 9.7) scales in direct contact with dorsal edge. Interorbital scales 6-11 (N 40, mean 8.0). Supralabial scales 11-17 (N 78, mean 13.5). Infralabial scales 11-17 (N 78, mean 13.4). Neck. Broad, less than half width of head. Body, Stout, dorsal and ventral surfaces with small granular scalation. Granular scales Inter- mixed with larger conical tubercules on dorsal and lateral surfaces. Mid-dorsal tubercules small, rump tubercules large (extremely pronounced) except in juveniles. Basal scales surrounding rump tubercules irregular, In many specimens, some are greater than half the height of the central scale, Rump tubercules larger than flank tuber- cules. Rump tubercules larger (han nape Luber- cules (90% material examined) or subequal to nape tubercules (10% material examined). Lower flanks and ventral surface with scattered, slightly saised rosettes which vary between being pronounced or barely discernible, Limbs. Long and slender, bearing enlarged jubercules on dorsal surfaces. Tubercules on thighs largest (extremely pronounced). Basal scales surrounding thigh tubereules irregular., In many specimens some are greater half the height of the central scale. Digits short, cylindrical, un- dilated distally and terminating in a non-retractile claw. Third toe on hindlimb longest, Tail. Short, moderately depressed, constricted at base and terminating in a globular, kidney- shaped knob (KW 21.3-35.0 ÆT, N 36, mean 27,6), Fonr longitudinal rows of tubercules present on dorsal and Jateral surfaces. Basal scales surrounding caudal tubercules less than half the height of the central scale. Caudal annuli 8-13 (N 37, mean 10.4). The measurements and scale counts for the holotype (AMR104458) are as follows; SVL 135.0mm, L1 54.1mm, L2 63.1mm, T 19.4mm, TV 23.3mm, HL 40.0mm, HW 34.4mm, HD 17.3mm, 5 12.8mm, EE 18.4mm, NL. 14,9mrn, KW 4.5mm, R 8,17. SL 15/15, IL 14/14, TA 10. 62 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4. Nephrurus amyae. Kings Ck Stn, southwestern NT, (Photograph: S. Wilson). This species has been illustrated previously as N. asper (e.g. Hoser, 1989: 74). PATTERN Body. In spirit, fawn to mid brown on dorsal surface. Many specimens with a black band present on the nape. Some specimens with seven to nine obscure, broken crossbands. These appear as a series of spots with each spot covering a tubercule. Others with no indication of banding or with obscure blotches along the vertebral line. Fine black lines form a reticulated pattern on the dorsum and upper lateral zone, These lines fade in larger specimens. Ventral surface cream. Limbs. Lacking irregular crossbands distally. Digits unbanded but sometimes obscurely mot- tled. Tail. As for body, with an obscure, broad, pale crossband on proximal half. Head. Often paler than body. Covered on top and sides with fine black reticulations which form an intricate pattern. These fade in larger specimens. ETYMOLOGY For Amy Couper. DISTRIBUTION Confined to the hill country of the south-central NT centred around Alice Springs (23°42’S, 133953'E), extending north to Barrow Ck (21932'S, 133953'E) (Fig. 3). HABITAT N. amyae favours open ground on rocky sub- strates in open woodland habitats. It may be found associated with a scattered 7riodia ground- cover. REPRODUCTIVE NOTES Clutch size two eggs. Females (NTMR33722- 23, NTMR5466) had small vitellogenic follicles on their ovaries between mid October - late December (< 5.5mm diameter). NTMR2458, col- lected in mid March, had a greatly enlarged fol- licle in each ovary (the largest of these follicles measured 14.58mm X 13.0mm). Bedford and Christian (1993) provide clutch data for a specimen from Gardens Station, 75km northeast of Alice Springs. Two eggs measuring 34.3mm X 16.6mm and 37.0mm X 16.1mm were laid on the 17/18 February 1992. These weighed 5.6g DESCRIPTION OF NEPHRURUS 63 and 5.8g respectively. The total clutch was 24,8% of the non-gravid female mass. Three males (NTMR33719, NTMR3372] & NTMR441) collected between early October - late November d to be in reproductive condition (epididymis enlarged and turgid), DIETARY NOTES The gut of NTMR33723 contained a large cen- tipede (Chilopoda: Scolopendridae), as did OMJ51650. OMJ53650 contained an ant head (Hymenoptera: Formicidae), the hind legs of a grasshopper (Orthoptera: Acrididae) and many medium-sized cockroach fragments (Blattodea: Blattidae), REMARKS N. amyae is one of Australia’s largest geckos in terms of SVL and weight (max SVL 135mm, AMRI(H458). It is exceeded in length only by Saliuarius cornutus (max SVL 145mm) and S, salebrosus (max SVL 141mm). However, despite its greatly reduced tail size, it is substantially heavier than both of these species and weighs a maximum of 65g (AMR104458). AMR10445& was compared in weight to a number of her large Australian gecko species held in the AM and QM collections. The results are as follows: Carphodactylus leavis OMJ54312(SVL 107mm. weight 26g); Cyrtodaerylus louisiadensis QMI38198 (SYL 130mm, weight 46g); Rhacodactylus australis QMJ39723 (SVL 110mm, weight 27g); Saltuarius cornutus OMJ48$179 {SVL 140mm, weight 49g), OMJ30991 (SVL 137mm, weight 57g), AMR42163 (SVL 145mm, weight 33g); £ salebrosus OMJ22288 (SVL 141mm, weight 57g) and S. swaini OMJ51640 (SVL 131mm, weight 43g), Nephrurus sheai Couper sp. nov, (Fig. 5; Statistica) analyses group 1) Nephrurus asper, Copger, 1992 (in part). MATERIAL EXAMINED HOLOTYPE: NTMR 11470, Bowerbird Camp, Magela Ck, NT (12°47°S 133°07'B), coll, 1. Archibald and J, Bywater, 01 September, 1983. PARATYPES; QMJ52872, March Fly Glen, Gibb R Ru, Kimberley, WA: QMJ57515, 10km SE of Oenpxlli, NT (12°22'S 133°07'E), AMR93181, Jabiluka Project ama, NT (12°29°S 132°54'E); AMR93182, Jubiluka Project area, NT (1273175. 132°57'Ek AMRSSO6R, Jabiluka Project area, NT (12°34'S 132755'E); AMR13403, “Katherine, NT (14°28'S 132° 16'E): AMR 12876, Manbullo Stn, NT (14731*S 13212'E), AMR72980, Jasper Gorge, NT (16°02'S 13040 E); AMR140279, Manning Gorge, Mi Barnett Sin, WA (16°39 31"S 125°55'37"E); NMVD4552, Pine Ck, NT (13°49"S 131°50°E):SAMR3597, Moolabulla, WA (18°12"S 127°30'E); WAMR13646, WAMR27374 Kalumburu, WA (14'1R'S 126°38"E}; WAMR47587, 8km N of Katherine, NT (14725'S 132°12'B); WAMR43153, WAMR77269, WAMR77581, WAMR77535 Mitchell Plateau, WA (14°49'S 125750'E); WAMR56423, King Edward R Crossing, WA (14752'S 126 IVE); WAMR50460, Drysdale R NP, WA (15°09"S 127°06'E); WAMR46782, Prince Regent R, WA (15719'5 125°35°E); WAMR73903, Ellenbrac Stn, WA (153757'S 127°04'B); WAMR60343, WAMR60947, Jasper Gorge 53km NW of Victoria River Downs Homestead, NT (16°02'S L30741'E); WAMR60362_ Jasper Gorge 53km NW of Victoria River Downs, NT (16°02'S 130°45'B); WAMR57123-7, Gibb R Crossing 47km N of Gibb River Stn, WA (16°06"S 126°31°E); WAMR352661, WAMR86927, Lake Argyle (Ord R), WA (16°07'S 128"44°E); WAMR83359, 24km NW of Mt Elizabeth Homestead, WA (16°11'S 126°00°E); WAMR70562, 3.5km at 288" from (new) Lissadell Homestead, WA (16°39'S 12R5317E); WAMR73904, Mt Barnett Stn, WA (1640'S 125°57'B); WAMR56466, Plain Ck. Beverley Springs Stn, WA (16'43'S 125°23°E); WAMRS3531-2, Galvans Gorge, WA (16"48'S 1257517E); WAMR64731,4.5km al 140° from Clancys Yard, WA (17°07'S [25°35'E); WAMR70551, 9.5km at 256° from Inglis Gap, WA (17°07°S, 125/05'E); WAMRS7118, Inglis Gap, King Leopold Range, WA (TOR'S (257TO'EN WAMR70552, 8.6km at 167° from Mt Amy (Napier Downs Stn), WA (17°14’S 124°54°E); WAMRS8671, 20km ENE of Tableland, WA (17°14'S 127°01"E); WAMR58636, 25km SE of Bedford Downs Homestead, WA (17°27'S 127°36'E); WAMR58647. 10km SE of Lansdowne Homestead, WA (17'41'S, 126°48'E); WAMR1340, Leopold Downs, WA (17°52°S 125725'E), WAMR12614, Cal- wynyardah, WA (18°00'S 124°47'E); WAMR70029, 43km at 316° from Fitzroy Crossing P.O., WA (18°00"S 125°30°E); WAMR48175, Moola Bulla, WA (18°11'S 127 30'E); WAMR26633, Halls Ck, WA (18°13"S 127°39'E); WAMR87092, 8km SW of Halls Ck, WA (TR 1678 127537' E); WAMR46116, Margaret River Homestead, WA (18°38'S 126°57'E),; WAMRS7082, 0.Skm N of Mary Pool, WA (18°43°S 126°53'E); NTMR12605, Nabarick, NT (1219'S 133°1L9°E); NTMR17822, 10km SE Oenpelli, NT (12°22'S 133°07'E); NTMR12493, Magela Ck, Bowerbird Camp, NT (12747'S 1393°OS*E); NTMR1 1464, Magela Ck, Bowerbird Camp, NT (I2747'S 133°07°B); NTMR4027, NTMR4272, Jim Jim Falls, NT (1371778 132°50°E); NTMR387, Katherine, 4.7m1 N, NT (14727'S 132'15'E); NTMR3759-51, Katherine, Akm N, NT (14 27'S 1327157E), NTMR267, NTMR297, Katherine, NT (14°28'S 132^1&'E); NTMR2377, Katherine, Lower Farm Rd, NT (14735'S 1321277E): NTMR33718, 4 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5. Nephrurus sheai. Manning Gorge, Gibb R Track, Kimberley region, WA. (Photograph: G. Harold). Wickham R, Victoria River Downs, NT (16'05'5 130°35"E); NTMR13485, Gregory NP, Bullita area, NT (16°07'S 130°26'E). DIAGNOSIS N. sheai is a large, moderately spinose member of the N. asper species group. N. sheai is distin- guished from N. asper and N. amyae by the colour pattern of its digits (digits strongly marked with alternating bands of brown and white vs digits not strongly banded with brown and white); also by size (max SVL 121mm vs 114mm N. asper, 135mm N. amyae). It is further distinguished from N. amyae by the spinosity of the its rump and thighs (moderately spinose vs extremely spinose). The arrangement of the basal scales surrounding the tubercules on the rump and thighs also separate N. sheai and N. amyae. In N. sheai the basal scales are uniform in size and less than half the height of the central scale. In N. amyae the basal scales are ir- regular in size and in most specimens examined, some of the basal scales are greater than half the height of the central scale, In the KS2 analysis N. sheai and N. amyae show significant separation of the shape of freguency distributions for; KW, HW, HL, S,NL, SL, IL, R & I. N. sheai and N. asper (excluding broad-banded CYP specimens which are not significantly separable from other OLD populations of this species) show significant separation of the shape of freguency distributions for; T, TV, KW, TA, NL, SL, IL, & I. (Table 1). DESCRIPTION SVL(mm): 42.3-121.4 (N 69, mean 82.8). Proportions,(%SVL); - L1 - 37,9-50.7 (N 69, mean 44.0), L2 - 45.0-59.1 (N 69, mean 49.8), AG - 35.2-52.4 (N 67, mean 43.9), T - 10.1-21.7 (N 68, mean 14.8), TV - 14.1-26.4 (N 68, mean 19.8), HL - 28.4-34.3 (N 68, mean 31.2), HW - 23.6-31.7 (N 69, mean 27.5), HD - 11.0-16.1 (N 69, mean 13.6), S - 9.4-13.0 (N 69, mean 11.0), EE - 10.9-15.1 (N 69, mean 12.8), NL - 8.6-18.0 (N 68, mean 14.6). Head. Large and deep, covered with small, round to hexagonal, juxtaposed scales. Posterior- ly bearing scattered tubercules intermixed with the smaller scales. Each tubercule consists of a high central scale, circled by a ring of smaller basal scales. Tubercules most prominent on the nape. Dorsal skin co-ossified with skull. Nostril small, opening upwards and backwards. Eye DESCRIPTION OF NEPHRURUS 65 large with vertical pupil. Ear vertically elongate, tympanum deeply recessed. Rostra] scale small, with 7-18 (N 69, mean 12.4) scales in direct contact with dorsal edge. Interorbital scales 8-15 (N 66, mean 10.5). Supralabial scales 12-20 (N 138, mean 15.8). Infralabial scales 11-20 (N 137, mean 16,1). Neck, Broad, less than half width of head, Body.Stout, dorsal and ventral surfaces with small granular scalation. Granular scales inter- mixed with larger conical tubercules on dorsal and lateral surfaces. Mid-dorsal tubercules small, rump tubercules small to large. Basal scales sur- rounding rump tubercules uniform, less than half as high as the central scale. Rump tubercules larger than flank tubercules (85% material ex- amined) or subequal to flank tubercules (15% material examined. Rump tubercules smaller than nape tubercules (41% material examined) or subequal to nape tubercules (59% material ex- amined). Lower flanks and ventral surface with scattered, slightly raised rosettes which vary be- tween being pronounced or barely discernible. Limbs. Long and slender, bearing enlarged tubercules on dorsal surfaces. Tubercules on thighs largest, with uniform basal scales Jess than half as high as the central scale. Digits short, cylindrical, undilated distally and terminating in a non-retractile claw. Third toe on hindlimb longest, Tail. Short, moderately depressed, constricted at base and terminating in a globular, kidney- shaped knob (KW 21.2-47.9 %T, N 67, mean 31.5). Four longitudinal rows of tubercules present on dorsal and lateral surfaces. Basal scales surrounding caudal 1ubercules uniform and less than half as high as the central scale, Caudal annuli 8-12 (N69, mean 9.7). The measurements and scale counts for the holotype (NTMR11470) are as follows; SYL 106.8mm, LI 45.3mm, L2 52.5mm, T 15.5mm, TV 214mm, HL 32.5mm, HW 29.8mm, HD 13.3mm, $ 10.9mm, EE 14.3mm, NL 17.4mm, KW 4.6mm, R 13,111, SL 18/17, IL 16/15, TA Il. This species has been illustrated previously as N. asper (e.g. Storr et. al, 1990: &2, pl.1; Schmida, 1985: 103). PATTERN Body. in spint, tan - greyish brown on dorsal surface. Many specimens have a broad black band on the nape, Seven - 1] narrow, irregular, cream crossbands present between head and hindlimbs. Crossbands range from obscure to bold and are continuous or broken into a series of spots, with each spot covering a tubercule. In most specimens fine black lines lie between the cream crossbands. These black lines are some- times interconnected to form a reticulated pattern. Ventral surface cream, sometimes faintly marbled with grey or brown, Limbs, Often with irregular crosshands on the distal half. Digits heavily banded with brown and white. Tail. As for body, with an obscure, broad, pale crossband on proximal half. Head. Tan, usually paler than the body, Covered dorsally and laterally with fine black reticulalions which form an intricate patter. These lines are most prominent in juveniles and may fade in larger specimens. ETYMOLOGY For Glenn Shea. DISTRIBUTION Confined to hill country from the Kimberley Region of WA to the Arnhem Escarpment in the NT (Fig. 3). HABITAT N. sheai favours rocky substrates in open Wood- Jand habitats. It is frequently associated with a scattered Triodia ground-cover (G. Harold, pers. comm.). The holotype (NTMR11470) was col» lected from a rocky outcrop in mid-high open forest with a grass understorey. REPRODUCTIVE NOTES Clutch size - usually two eggs, although a single egg clutch has been recorded (How et. al., 1990). Gravid females are present in the population in January - WAMR86927, WAMR27374; February- WAMR87092 and June - NTMR33718. WAMR70551 had a greatly en- larged vitellogenic follicle (12.42 x 10.83mm)in the ovary in May. Incubation of the eggs takes 115 days at a mean temperature of approximately 25°C (Ehmann, 1992). Gow, 1979 recorded the measurements of two eggs laid on 5 Dec 1973 (30 x16.5mm and 29x 15.5mm). Males were in reproductive condition (epididymis enlarged and turgid) in November - WAMR56423, NTMR267 and in Febmary - WAMR48175. How et al. (1990), record reproductive activily extending over at least nine months (spanning the October- April wet period) for N, sheai in WA and suggest that reproductive activity may occur throughout the year. 66 MEMOIRS OF THE QUEENSLAND MUSEUM DIETARY NOTES The gut of QMJ57515 contained moth or cater- pillar cuticle with dense, short hairs. QMJS2872 contained numerous termite heads (Nasutitermes sp.)- REMARKS AMR125387 was collected from Cadell Ck, near Hamilton, western OLD. This specimen has strongly banded toes, a diagnostic feature of N. sheai. lis occurrence in Queensland is not consis- tent with the overall distribution pattern of this species. Further, of the QLD material examined in this study ( 79 XN. asper}, AMR125387 is the only specimen to possess strongly banded toes. Because of these inconsistencies, this specimen has not been included jn the material examined for either N. sheai or N. asper. The status of this specimen is uncertain, We believe there has been an error with the collection data. KEY TO N. ASPER SPECIES COMPLEX Digits not strongly banded 2 Tube of he hump sec) thighs cxteniely pebaoanaed, na aura yn Tae ium a N, amyae aon o AUU Be Oh UO 58 8 Rh moderate ACKNOWLEDGEMENTS Rot meu AAG Go ID N. asper The authors wish to thank the Oueensland Museum and the Royal Geographical Society of Oueensland for supporting fieldwork associated with this project; Ross Wilkinson for assisting with the statistical computing analyses; Paul Horner, Lawrie Smith, Ross Sadlicr, Mark Hutchinson, John Coventry and the Donald F. Thomson Administration Committee for allow- ing access fo specimens in their care; Darren Storch for collecting additional matenal from CYP; Steve Wilson, Dave Knowles and Greg Harold for allowing the use of their photographs; Steve Wilson, Greg Harold, and Tim Hawkes (Cape York Herpetological Society) for generously sharing their field observations with us; Jeff Wright, Bruce Cowell & Garry Cranitch (Old Museum) for preparing the photographic plates; Kate Couper, Martin Cohen, Steve Wil- liams, Jeanette Covacevich and Martin Schultz for assistance in the field; Geoff Monteith for examining gut contents, Lauren Keim for assist- ing in the Laboratory; Glen Ingram and Steve Donnellan for their encouragement and advice and Jeanette Covacevich and Glenn Shea for suggesting improyements to the manuscript. LITERATURE CITED ANON. 1952-69. ‘Atlas of Australian resources’. (Department of National Development: Canber- ra), BAUER, A. M, 1990, Phylogenetic systematics and biogeography of the Carphodactylini (Rep- tilia:Gekkonidae). Bonner Zoologische Monographien 30; 1-217. BEDFORD, G, & CHRISTIAN, K. 1993, Egg size of the Prickly Knob-tailed Gecko (Nephrurus asper Gunther, 1876) with a preliminary comparison of investment per progeny among geckos. Dactylus 1(4):38-41, CAMERON, E.E. & COGGER, H. G. 1992, The her- petofauna of the Weipa region, Cape York Penin- sula, Technical Reports of the Australian Museum 7: 1-200. COGGER, H. G. 1992. ‘Reptiles and amphibians of Australia’, Sth ed, (Reed: Sydney). COGGER, H. G., CAMERON, E.E, & COGGER, H. M. 1983. Amphibia and Reptilia. Pp1-200. In Walton, D.W. (exec. ed.), “Zoological catalogue of Australia, Vol. 1'. (Australian Government Publishing Service: Canberra), COUPER, P. J, COVACEVICH, J. A, & MORITZ, C. 1993. A review of the leaf-tailed geckos endemic to castern Australia: a new genus, four new species, and other new data. Memoirs of the Queensland Museum 34(1): 95-124, EHMANN, H. 1992, ‘Encyclopedia of Australian animals: reptiles’. (Angus & Robertson; New South Wales). GOW, G. 1979, Notes on the biology of Nephrurus asper. Northem Territory Naturalist 1(2): 19-20. GUNTHER, A. 1876, Descriptions of new species of reptiles from Australia collected by Hr, Damel for the Godelfroy Museum. Journal Des Museum Godeffroy, Geographische, ethnographischc und naturwissenschaftliche Mitteilungen 5: 45-47. HOSER, R.T, 1989. ‘Australian reptiles & frogs,’ (Pier- son & Co: Sydney). HOW, R. A. DELL, J. & WELLINGTON, B.D., 1990. Reproductive and dietary biology of Nephrurus and Underwoodisaurus (Gekkonidac) in Westem Australia. Records of the Westem Australian Museum 14(4): 449-459, KOLMOGOROV, A. 1941. Confidence limits for an unknown distribution, Annals of Mathematical Statistics 12: 461-463. RAO, C.R, 1952. ‘Advanced staustical methods in biometric research,’ (New York: Wiley). SCHMIDA, G. 1985, “The cold-blooded Australians,’ (Doubleday Australia Pty Ltd; Sydney). SMIRNOV, N.V, 1939, On the estimation of the dis- crepancy between empincal curves of distribution DESCRIPTION OF NEPHRURUS 67 for two independent samples. Bulletin of Mathe- 1989. A new Heteronotia (Lacertilia: Gekkonidae) matics, University of Moscow 2: 1-16. from Western Australia, Records of the Western STORR, G.M. 1963. The gekkonid genus Nephrurus in Australian Museum 14(3): 269-273. Westem Australia, including a new species and STORR, G.M., SMITH, L.A. & JOHNSTONE, R.E. three new sub-species. Journal of the Royal 1990. ‘Lizards of Westem Australia, Vol. 3, geckos Society of Western Australia 46: 85-90. & pygopods.’ (Western Australian Museum: Perth). 1988. Three new Diplodactylus (Lacertilia: Gek- WALSH, J.E. 1965. ‘Handbook of nonparametric konidae) from the arid zone of Australia. Records statistics’, Vol. 2. (Princeton: Van Nostrand). of the Western Australian Museum 14(2): 217- WILSON, S. K. & KNOWLES, D. G. 1988. ‘Australia’s 223. reptiles’. (Collins Australia: Sydney). 68 MEMOIRS OF THE QUEENSLAND MUSUEM AN EARLY FLECKER RADIOGRAPH OF HYDROPHIS ELEGANS, AND NEW INFORMATION ON ITS FEEDING HABITS. Memoirs of the Queensland Museum 37(1):68.1994:- Dr Hugo Flecker (1884-1957) was a prominent naturalist - radiologist who spent much of his life in Cairns, NEO, His work is widely known. He made large, significant, botanical collections; founded the North Queensland Naturalist's Club; and published extensively on plant and animal injuries to humans. A host of plant and animal species bear the name fleckeri in recognition of the value of his contributions to natural history. Flecker was also a pìoneer in the field of radiotherapy and published on this, and on early bone development in humans (Clarkson, 1990; Pearn, 1994). It is not surprising to find that Flecker combined his major interests of natural history and radiology. He took at least 20 radiographs of native animals. These have recently been redis- covered (JRC). Amongst the series of prints (all original films appear to be lost) are radiographs of two species of sea snakes, Hydrophis elegans and Astrotia siokesii. That of the latter species is insignificant zoologically. However, that of the H. elegans specimen provides new data on the feeding habits of this species, and is of aesthetic interest (Fig, 1). Neither of the sea snake radiographs bears a date, However, as others in the series were taken in 1939, it seems reasonable to suggest those of the sea snakes date, at least roughly, from the same era. The prey species of many sea snakes of Australia's tropi- cal and subtropical waters are well known (e.g, Limpus, 1987). Most are bottom-feeders. Some have ultra-specialized diets (e.g. Emydocephalus annulatus, which feeds on only bleniniid and gobiid fish eggs from burrows in coral reefs). A few feed on a wide range of animals (e.g. Aipysurus laevis eats fish, prawns, crabs, worms and fish eggs from coral and rock reefs). H. elegans is reported to feed solely on very elongate fishes from soft bottom inshore waters (Limpus, 1987), Flecker's radiograph shows clearly three fish (the posterior-most specimen is a Herring, Family Clupeidae; the others are Cardinal Fish, Family Apogonìdae: R, McKay, pers. comm.) and a shrimp (Infraorder Caridea: J.Short, pers. comm.) in the gut of a specimen identified by him as H. elegans. This suggests that H. elegans may be a less highly specialised feeder than reported to date. Literature Cited Clarkson, J.R. 1990, Hugo Flecker and the north Queensland Naturalists’ Club, Pp. 171-178. In Short P.S. (ed.), “A history of systematic botany in Australasia’. Proceed- ings of a Symposium held at the University of Mel- bourne, 25-27 May, 1988. (Australian Systematic Botany Society Inc.; Melbourne). Limpus, C.J. 1987. Sea snakes. Pp. 195-204. In Covacevich, J., Davie, P, & Pearn, J. (eds), “Toxic plants and animals, A guide for Australia’. (Queensland Museum: Brisbane). Pearn, J. 1994. Hugo Flecker's contribution to medicine and natural history. The Medical Journal of Australia 16]: 63-70. J.A. Covacevich, Oueensland Museum, PO Box 3300, South Brisbane, Oueensland 4101, Australia; J.H. Peurn, Depart- ment of Child Health, University of Oueensland, Royal Children's Hospital, Herston, Queensland 4029, Australia; J.R, Clarkson, Queensland Herbarium (Mareeba office), Mareeba, Queensland 4880, Australia; 20 July 1994. ” FIG. 1, Radiograph of Hydraphis elegans, following ingestion of two fish and a shrimp, (Dr Hugo Flecker). TYPE SPECIMENS OF FROG AND REPTILE SPECIES, QUEENSLAND MUSEUM: RECENT ADDITIONS AND NEW INFORMATION J.A. COVACEVICH AND P.J. COUPER Covacevich, J.A. & Couper, P.J. 1994 12 01: Type specimens of frog and reptile species, Oueensland Museum: recent additions and new information. Memoirs of the Oueensland Museum 37(1):53-65. Brisbane. ISSN 0079-8835. Since publication of the last list of frog and reptile type specimens in 1981, types of 25 species of frogs and 78 species of reptiles have been deposited in the Queensland Museum reference collections. In addition, new data are to hand on types of two species of frogs and 14 species of reptiles listed in previous Queensland Museum type lists. [ |] Type specimens, frogs, reptiles, Queensland, Australia . J.A. Covacevich & P.J. Couper, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia; 20 August 1994. The tradition of listing type specimens in the frog and reptile reference collections of the Queensland Museum began over forty years ago (Mack & Gunn, 1953; Covacevich, 1971; Ingram & Covacevich, 1981). Since the last list went to press in June, 1980, there have been many additions to the type collec- tions; a ‘lost’ holotype has been found; some problems identified in previous lists have been solved; the currently recognised names of several have changed; and there are also some corrections of transcription and other errors. In recent years, several texts of Australia’s frog and reptile species have been published (e.g. Cogger, 1992). From these, it is tempting to see Queensland’s species as well known at least to the first level of description and documentation of distribution. How- ever, the rate of discovery and description of new species has been so rapid, that each of these texts has been incomplete at publication. For example, the review of Cogger (1992) is the most recent, and is extremely comprehensive. Yetit does not include one species of frog and 21 species of reptiles described from Queensland since its publication. Between June, 1980 and January, 1995, 22 species of frogs and 72 species of reptiles have been described from Queensland. Almost all are currently recognised. This list is presented in two parts. In the first, the names are arranged alphabetically by genus and species, in standard phylogenetic order. For each, we provide author/s, date, current status (if there is any variation), registration number, and any necessary explanatory comments. In the second, we document all changes which have been made with names or taxa listed in previous Queensland Museum type lists, in the same order and format. The following abbreviations have been used: - Australian Museum (AM); Queensland Museum (QM) and Queensland National Parks and Wildlife Service (QNPWS - now Queensland Department of Environment and Heritage). FROG AND REPTILE TYPE SPECIMENS DEPOSITED IN THE QUEENSLAND MUSEUM SINCE JUNE 1980 Class AMPHIBIA Family HYLIDAE Cyclorana manya Van Beurden & McDonald, 1980 Holotype QMJ34886; paratypes QMJ34887-88, QMJ36894-900. Data as for type description. Er- roneously, QMJ34889 was listed in the type descrip- tion instead of QMJ34887. The number OMJ34889 is for a specimen of Uperoleia lithomoda. Litoria electrica Ingram & Corben, 1990 Holotype QMJ38963; paratypes QMJ27240-43, QMJ38964, QMJ38973-74, QMJ38976-77, OMJ49227-28. Data as for type description. Litoria pallida Davies, Martin & Watson, 1983 Paratypes OMJ39256-58, OMJ41013-19. Data as for type description. Litoria revelata Ingram, Corben & Hosmer, 1982 Holotype OMJ28233; paratypes OMJ10896, OMJ12853, OMJ13156, OMJ19867-69, OMJ19872- 73, OMJ19887, OMJ19889-90, OMJ19892-94, OMJ19898, OMJ19900, OMJ19909-12, OMJ19915, OMJ19918, OMJ28232, OMJ30116-35, OMJ30137- 70 MEMOIRS OF THE QUEENSLAND MUSEUM 59, OMJ31465, QMJ31467-68, OMJ31971, QMJ35087, QMJ35105-06, QMJ35115-16, QMJ35539, QMJ35543. Data as for type description. Litoria xanthomera Davies, McDonald & Adams, 1986 Holotype QMJ42011; paratypes QMJ17109-10, QMJ25194-95, QMJ25258-60, QMJ25278, QMJ27105-06, QMJ35900-03, QMJ35910, OMJ35916, OMJ35919, QMJ35921, QMJ35943-44, QMJ35946, OMJ35950, OMJ35960, OMJ35962-63, OMJ35985-86, OMJ36011, OMJ36020, OMJ36024. Data as for type description. Nyctimystes oktediensis Richards & Johnston, 1993 Holotype OMJ56896; paratypes OMJ56897-98. Data as for type description. Family MICROHYLIDAE Asterophrys leucopus Richards, Johnston & Burton, 1994 Holotype OMJ58650. Data as for type description. Cophixalus bombiens Zweifel, 1985 Holotype OMJ42060; paratype OMJ42063. Data as for type description. Cophixalus crepitans Zweifel, 1985 Holotype OMJ28817; paratypes OMJ41643-53. Data as for type description. Cophixalus hosmeri Zweifel, 1985 Holotype OMJ42058; paratypes OMJ37281-83, OMJ56467-91. Data as for type description. Cophixalus infacetus Zweifel, 1985 Holotype OMJ42059; paratypes OMJ29728-29, OMJ41639-41. Data as for type description. Cophixalus mcdonaldi Zweifel, 1985 Holotype OMJ42064; paratypes OMJ42040-57. Data as for type description. Cophixalus monticola Richards, Dennis, Trenerry & Werren, 1994 Holotype OMJ58727; paratypes OMJ58728-33, OMJ58854-57, OMJ58871-74. Data as for type description. Cophixalus peninsularis Zweifel, 1985 Holotype OMJ42061; paratype OMJ42062. Data as for type description. Cophixalus tuberculus Richards, 1992 Paratype OMJ55246. Data as for type description. Family MYOBATRACHIDAE Mixophyes fleayi Corben & Ingram, 1987 Holotype OMJ26901; paratypes OMJ5198, OMJ26469-70, OMJ27859, OMJ29930-31, QMJ30545-48, OMJ32059, OMJ34101-03, QMJ34243-44, OMJ35461-65. Data as for type description. Pseudophryne covacevichae Ingram & Corben, 1994 Holotype OMJ52286; paratypes OMJ53870, OMJ53920-32. Data as for type description. Pseudophryne raveni Ingram & Corben, 1994 Holotype OMJ58433; paratypes OMJ1505-06, OMJ2955, OMJ12304, OMJ12310, OMJ12385, OMJ12867, OMJ18699-700, OMJ18702, OMJ18704- 05, OMJ18707-13, OMJ18716-23, OMJ18725-26, OMJ18729, OMJ19824, OMJ23855-56, OMJ23952, OMJ27399-402, OMJ27449, OMJ27451, OMJ27453, OMJ27483, OMJ27949, OMJ29259, OMJ29263, OMJ29277-79, OMJ29990-91, OMJ30085, OMJ30878, OMJ30882, OMJ31881, OMJ31886-89, QMJ32209, OMJ34160-61, OMJ35108, OMJ35801-06, QMJ36902, OMJ37032-33, OMJ37470, OMJ37543, OMJ39268, OMJ40141-49, OMJ40176, OMJ40214-15, OMJ40258, OMJ40442-46, OMJ40517-19, OMJ41959, OMJ42158- 59, OMJ42186, OMJ42220, OMJ42643-69, OMJ42671, OMJ42677, QMJ42691, OMJ42749, OMJ42751-53, OMJ42757, OMJ42760-64, OMJ43850-51, OMJ49439, OMJ50630, OMJ50632, OMJ50637-42, OMJ50644-56, OMJ52285, OMJ54402-05, OMJ55019, QMJ55021, QMJ55194, QMJ56789, QMJ57286, OMJ58434. Data as for type description. FROG AND REPTILE TYPE SPECIMENS 71 Rheobatrachus vitellinus Mahony, Tyler & Davies, 1984 Holotype QMJ42529; paratype QMJ42145. Data as for type description. Taudactylus pleione Czechura, 1986 Holotype QMJ42392; paratypes QMJ42137, OMJ42388-91, OMJ42422-23. Data as for type description. Uperoleia altissima Davies, Watson, McDonald, Trenerry & Werren, 1993 Holotype QMJ55301; paratypes QMJ19851-52, OMJ19855-587, OMJ51780-84. Data as for type description. Uperoleia capitulata Davies, McDonald & Corben, 1986 Holotype OMJ26428; paratypes OMJ26416, OMJ26427, OMJ39196, OMJ45956-60. Data as for type description. Uperoleia fusca Davies, McDonald & Corben, 1986 Paratypes OMJ10900-02, OMJ12294, OMJ12309, QMJ12323, OMJ12721, OMJ18817, OMJ18822, OMJ18825, QMJ18832, OMJ18837, OMJ19925, OMJ19927, OMJ19930, OMJ19936, OMJ19938-39, OMJ19941-43, OMJ19947, OMJ19952-53, OMJ19956-59, OMJ19961, OMJ19965-66, OMJ24088, OMJ27475, OMJ27477, OMJ27481, QMJ27563, OMJ27905, OMJ29280, OMJ30606, OMJ31556-62, OMJ31582, OMJ39315, OMJ40020, OMJ40315, OMJ40431, OMJ40433-35, OMJ40484, OMJ40486-87, OMJ40489-90, OMJ41533, OMJ42563-76, OMJ42578-614, OMJ45961-65. Data as for type description. Uperoleia littlejohni Davies, McDonald & Corben, 1986 Holotype OMJ45949; paratypes OMJ29874, OMJ38877-80, OMJ38883, OMJ38915, OMJ43154- 55, OMJ45950-54. Data as for type description. After providing a list of paratypes (p. 175), the authors list cleared and stained paratypes separately (p. 178). Here they fail to mention OMJ45952 and OMJ45954, both cleared and stained specimens already listed on p. 175. Further, OMJ35427 is erroneously listed as a cleared and stained paratype. This number is for a specimen of Ctenotus robustus. Uperoleia mimula Davies, McDonald & Corben, 1986 Holotype OMJ45943; paratypes OMJ19859, OMJ38271-72, OMJ40234-35, OMJ40247, OMJ42534-35, OMJ42537, OMJ45944-48. Data as for type description. Class REPTILIA Family CHELIDAE Elusor macrurus Cann & Legler, 1994 Holotype OMJ51275; paratypes OMJ51274, OMJ51468-70, OMJ54138-40. Data as for type description. Rheodytes leukops Legler & Cann, 1980 Holotype OMJ31701; paratypes OMJ31702-7. Data as for type description. Family GEKKONIDAE Bavayia septuiclavis Sadlier, 1988 Paratypes OMJ43985, OMJ44034. Data as for type description. Erroneously, OMJ44985 was listed in the type description instead of OMJ43985. The number OMJ44985 is for a specimen of Morelia spilota. Bavayia validiclavis Sadlier, 1988 Paratype OMJ43980. Data as for type description. Nephrurus amyae Couper (in Couper & Gregson, 1994) Paratypes OMJ51650, OMJ53650. Data as for type description. Nephrurus sheai Couper (in Couper & Gregson, 1994) Paratypes OMJ52872, OMJ57515. Data as for type description. 72 MEMOIRS OF THE QUEENSLAND MUSEUM Phylurus isis Couper, Covacevich & Moritz, 1993 Holotype QMJ53511; paratypes QMJ53480, OMJ53485-86, QMJ53512, OMJ53518, QMJ53591, OMJ53602-3. Data as for type description. Phyllurus nepthys Couper, Covacevich & Moritz, 1993 Holotype QMJ34058; paratypes QMJ32634-35, QMJ32669, QMJ32674-76, QMJ32695-96, OMJ32733, OMJ32740, OMJ34024-25, OMJ34076- 79, OMJ35128, OMJ50992-93, OMJ51098-101, 0MJ53330-32, OMJ53359-62. Specimen OMJ25411 is also a paratype of Phyllurus caudiannulatus Covacevich, 1975. Data as for type description. Phyllurus ossa Couper, Covacevich & Moritz, 1993 Holotype OMJ53444; paratypes OMJ53389-93, OMJ53414, OMJ53426-28, OMJ53443, OMJ53445- 47, OMJ53507, OMJ56311, OMJ56766-75, OMJ56791-92. Data as for type description. Saltuarius occultus Couper, Covacevich & Moritz, 1993 Holotype OMJ37040; paratypes QMJ37037-39. Data as for type description. Family PYGOPODIDAE Delma labialis Shea, 1987 Holotype QMJ45563; paratype QMJ30265. Data as for type description. Delma mitella Shea, 1987 Holotype QMJ32597. Data as for type description. Family AGAMIDAE Pogona brevis Witten, 1994 Holotype QMJ32292; paratypes QMJ38735, OMJ38760-1, OMJ46949. Data as for type description. Family SCINCIDAE Anomalopus brevicollis Greer & Cogger, 1985 Holotype QMJ42616; paratypes QMJ4561, OMJ33863, OMJ33870, OMJ34056, OMJ38740, OMJ41997-98, OMJ42454, QMJ46272. Data as for type description. Anomalopus gowi Greer & Cogger, 1985 Holotype OMJ42615; paratypes OMJ27617, OMJ31050. Data as for type description. Anomalopus mackayi Greer & Cogger, 1985 Paratypes OMJ8516, QMJ42433, OMJ42531. Data as for type description. Anomalopus swansoni Greer & Cogger, 1985 Paratype OMJ44236. Data as for type description. Calyptotis lepidorostrum Greer, 1983 Holotype OMJ33612; paratypes OMJ22067, OMJ22274, QMJ22472-73, OMJ23810, OMJ24133, OMJ24373, OMJ25397, OMJ29093, OMJ30229, QMJ30231, QMJ30236, OMJ31575, OMJ32653-55, OMJ32671, OMJ32673, OMJ32707-16, OMJ32721- 22, QMJ32734-38, OMJ33610, OMJ33613-15, 0MJ33737, OMJ33752, OMJ35309, OMJ35878-82, OMJ50545-48. Data as for type description. Calyptotis ruficauda Greer, 1983 Paratype OMJ26024. Data as for type description. Calyptotis temporalis Greer, 1983 Holotype OMJ32594; paratypes OMJ13718, OMJ25742-43, OMJ34089, OMJ35088. Data as for type description. Calyptotis thorntonensis Greer, 1983 Holotype OMJ28354. Data as for type description. FROG AND REPTILE TYPE SPECIMENS 73 Carlia parrhasius Couper, Covacevich & Lethbridge, 1994 Holotype QMJ57868; paratypes QMJ57867, OMJ57869. Data as for type description. Carlia pectoralis inconnexa Ingram & Covacevich, 1989 Paratypes OMJ25060, OMJ42496. Data as for type description. Carlia rubrigularis Ingram & Covacevich, 1989 Holotype OMJ29956; paratypes OMJ17826, OMJ17901, OMJ17906-07, OMJ22668, OMJ24649, OMJ24800, OMJ24807-08, OMJ25141, OMJ25143, OMJ25146, OMJ25198-200, OMJ25209, OMJ25211- 12, OMJ25240, OMJ25242, OMJ25245-50, QMJ25272, OMJ25293-94, OMJ25296-97, OMJ50335. Specimen QMJ25229 was listed as a paratype (Ingram & Covacevich, 1989). This specimen has now been re-registered as QMJ50335. The number OMJ25229 had been assigned also to a paratype of Lampropholis basiliscus. Data as for type description. Carlia storri Ingram & Covacevich, 1989 Holotype QMJ24656; paratypes QMJ17804, QMJ17895-96, QMJ17908, QMJ24639-40, OMJ24653-55, OMJ24657-58, OMJ24676, OMJ24683-85, OMJ24691, OMJ25312, OMJ25429, OMJ25600, OMJ26222, OMJ26256, QMJ26269. Data as for type description. Ctenotus allotropis Storr, 1981 Paratypes OMJ31847-8, OMJ34770. Data as for type description. Ctenotus aphrodite Ingram & Czechura, 1990 Holotype OMJ41814. Data as for type description. Ctenotus arcanus Czechura & Wombey, 1982 Holotype OMJ36925; paratypes OMJ437-38, OMJ440-42, QMJ1682, OMJ11030, OMJ12111-12, OMJ30266, OMJ30722-23, OMJ31863, OMJ34588, OMJ38695. Data as for type description. Ctenotus astarte Czechura, 1986 Holotype OMJ26499; Paratypes OMJ39580, OMJ40182-83, OMJ41603, OMJ41796. Data as for type description Ctenotus capricorni Storr, 1981 Holotype OMJ39470. Data as for type description. This specimen is listed in the type description as AM R65946. Ctenotus essingtonii brevipes Storr, 1981 Holotype OMJ39469. Data as for type description. Ctenotus eurydice Czechura & Wombey, 1982 Paratypes OMJ1618, OMJ15614, OMJ27526, OMJ39223. Data as for type description. Ctenotus eutaenius Storr, 1981 Holotype OMJ39467. Data as for type description. This specimen is listed as AM R93408 in type descrip- tion. Ctenotus hypatia Ingram & Czechura, 1990 Holotype OMJ42092. Data as for type description. Ctenotus ingrami Czechura & Wombey, 1982 Holotype OMJ34792; Paratype OMJ34791. Data as for type description. Ctenotus monticola Storr, 1981 Holotype OMJ39468. Data as for type description. This specimen is listed in the type description as AMR70937. Ctenotus nullum Ingram & Czechura, 1990 Holotype QMJ32424; paratypes QMJ24647, OMJ24705, OMJ37999-38001, OMJ41023-25, OMJ42736, OMJ42768-69. Data as for type descrip- tion. 74 MEMOIRS OF THE QUEENSLAND MUSEUM Ctenotus serotinus Czechura, 1986 Holotype QMJ43313; paratype QMJ40185. Data as for type description. Ctenotus terrareginae Ingram & Czechura, 1990 Holotype QMJ41996. Data as for type description. Ctenotus zebrilla Storr, 1981 Holotype QMJ39471. Data as for type description. This specimen is listed in the type description as AMR63316. Emoia atrocostata australis Brown, 1991 Paratype QMJ24732. Data as for type description. Eulamprus frerei Greer, 1992 Holotype QMJ47985; paratype QMJ39531. Data as for type description. Eulamprus sokosoma Greer, 1992 Holotype QMJ27702; paratypes QMJ15668, OMJ25915, QMJ27622-23, OMJ33843, QMJ34209, OMJ42506, OMJ42513, QMJ42516-17, QMJ42526, OMJ55403, OMJ55404. Data as for type description. Specimens OMJ55403 and OMJ55404 are listed in the type description as ONPWSN1787 and ONPWSN36821 respectively. Lampropholis adonis Ingram, 1991 Holotype OMJ35097; paratypes OMJ23805-08, OMJ23867, OMJ27725, OMJ27732, OMJ27820, OMJ27822, QMJ27825, QMJ27833, QMJ32644, QMJ32652, QMJ32701, QMJ32706 QMJ32754, QMJ32793, QMJ32795, OMJ33604, OMJ33607, OMJ33609, OMJ33701, QMJ33717-18, OMJ33720- 24, QMJ33736, QMJ35104, OMJ35122, OMJ35198, OMJ45297-98, OMJ46191, OMJ49563, OMJ49573, OMJ49591, OMJ49597, OMJ49746-48, OMJ49750- 51, OMJ49753, OMJ51319, OMJ51323. Data as for type description. Lampropholis amicula Ingram & Rawlinson, 1981 Holotype OMJ24333; paratypes OMJ22729, OMJ24330-2, OMJ24496-7, OMJ27523, OMJ30828, OMJ31308, OMJ32090, OMJ32519, OMJ37177-78, OMJ37288-89. Data as for type description. Er- roneously, OMJ34330-31 were listed in the type description instead of OMJ24330-31. Numbers OMJ34330-31 are for specimens of Lechriodus fletcheri. One of the paratypes, OMJ37287 has been transferred to the Australian Museum's herpetological collection, with the registration number AMR96587. Lampropholis basiliscus Ingram & Rawlinson, 1981 = Saproscincus basiliscus after Greer (1989). Holotype OMJ34409; paratypes QMJ11161, QMJ12145-47, OMJ12158-59, OMJ17435, OMJ17902-03, OMJ24648, OMJ24848, OMJ24918, OMJ25022-27, OMJ25029-45, OMJ25047-59, OMJ25061, OMJ25137-38, OMJ25204, OMJ25229, OMJ25257, OMJ25265, QMJ25289, OMJ25301, OMJ25308, OMJ25450, OMJ25825-75, OMJ26330- 32, OMJ26375-76, OMJ27135, OMJ27141, OMJ27258, OMJ29668, OMJ30810-11, OMJ32354, OMJ32602, OMJ32605, OMJ32758, OMJ32760, OMJ32766-71, OMJ32780-83, OMJ32794, OMJ34000-06, OMJ34036, OMJ34038-40, OMJ34042, OMJ34047, OMJ34062, OMJ34066, QMJ34069, OMJ34092-93, OMJ34095-100, OMJ34408. Data as for type description. Sadlier et al. (1993) treated S. basiliscus as a junior synonym of S. spectabilis. Following discussion of the type status of Mocoa spectabilis, Ingram (1994) showed that S. spec- tabilis was a senior synonym of S. galli and not of S. basiliscus as proposed by Sadlier et al. (1993). Lampropholis caligula Ingram & Rawlinson, 1981 Paratype OMJ38704. Data as for type description. Lampropholis coggeri Ingram, 1991 Holotype OMJ27133; paratypes OMJ12205, OMJ14009, OMJ14022, OMJ14092, OMJ18006, OMJ21408, OMJ25139, OMJ25201-03, OMJ25217, OMJ25230, OMJ25241, OMJ25243-44, OMJ25251- 55, QMJ25271, OMJ25285, OMJ25299, OMJ25300, OMJ25330, OMJ26301, OMJ27008-11, OMJ27130- 32, OMJ27134, OMJ29622, OMJ39858, OMJ39864, QMJ39872, OMJ40536, OMJ40663, OMJ41733, OMJ42276-17, OMJ42294, OMJ48170, OMJ48172, OMJ48210. Data as for type description. FROG AND REPTILE TYPE SPECIMENS 75 Lampropholis colossus Ingram, 1991 Holotype QMJ49687; paratypes QMJ27549, OMJ27550, OMJ30655, OMJ30656, QMJ30657, OMJ30659-60, OMJ46095-97, OMJ49689, OMJ49692. Data as for type description. Lampropholis couperi Ingram, 1991 Holotype OMJ49575; paratypes OMJ25741, OMJ25745, QMJ27723, QMJ30293, OMJ30827, OMJ33605, OMJ34162, OMJ40153-57, OMJ42156, OMJ42426, OMJ43956, OMJ47950-51, OMJ49660, OMJ49669, OMJ49752, QMJ51663. Data as for type description. Lampropholis czechurai Ingram & Rawlinson, 1981 — Saproscincus czechurai after Greer (1989). Holotype OMJ34402; paratypes OMJ12148-50, OMJ25227, QMJ27072-73, OMJ31201, OMJ31204- 06, OMJ34403, OMJ34405-07. Data as for type description. Lampropholis mirabilis Ingram & Rawlinson, 1981 Holotype OMJ24439; paratypes OMJ4404, OMJ24339-40, OMJ24416, OMJ24424, OMJ24435, OMJ24437-38, OMJ24440-41, OMJ24528, OMJ27615, OMJ32555. Data as for type description. Lampropholis robertsi Ingram, 1991 Holotype OMJ43911; paratypes OMJ31194-200, OMJ39490-91, OMJ39855-57, OMJ40033, OMJ40036-39, OMJ40041, OMJ40609-10, OMJ41706-08, OMJ43912, OMJ43918, OMJ43958, OMJ43964, OMJ46193, OMJ47097, OMJ47956, OMJ47959, OMJ48295, OMJ49648, OMJ49659, OMJ51405-06, OMJ51948. Data as for type descrip- tion. Leiolopisma jigurru Covacevich, 1984 — Bartleia jigurru after Hutchinson et al., (1990). Holotype OMJ40040, paratypes OMJ39492-99. Data as for type description. Leiolopisma zia Ingram & Ehmann, 1981 = Cautula zia after Hutchinson et al., (1990). Holotype OMJ30563; paratypes OMJ26025, OMJ27787-88, OMJ27793, OMJ27855-58, OMJ30213, OMJ30555-62. Data as for type descrip- tion. Lerista aericeps aericeps Storr, 1986 Paratypes OMJ26502, OMJ34137, OMJ39572. Data as for type description. Lerista cinerea Greer, McDonald & Lawrie, 1983 Holotype OMJ40097; paratypes OMJ40094-96, OMJ40098-100. Data as for type description. Lerista colliveri Couper & Ingram, 1992 Holotype OMJ16181; paratypes OMJ16182-83, OMJ33123-28, OMJ45648, OMJ46266-67. Data as for type description. Lerista emmotti Ingram, Couper & Donnellan, 1993 Holotype OMJ53959; paratypes OMJ9038, OMJ50066, OMJ50068-69, OMJ51217-18, OMJ51529, OMJ51629-30, OMJ52585-87, OMJ53958, OMJ53960, OMJ54144-46, OMJ54292, OMJ54491-93. Data as for type description. Lerista ingrami Storr, 1991 Holotype OMJ32396; paratypes OMJ20644-51, OMJ20653. Data as for type description. Lerista storri Greer, McDonald & Lawrie, 1983 Holotype OMJ39480; paratype OMJ39481. Data as for type description. Lerista vittata Greer, McDonald & Lawrie, 1983 Holotype OMJ40102; paratypes OMJ40101, OMJ40103-04. Data as for type description. 76 MEMOIRS OF THE OUEENSLAND MUSEUM Lerista zonulata Storr, 1991 Holotype OMJ54137; paratypes OMJ31223, OMJ38756, OMJ46268, OMJ47103. Data as for type description. The holotype is listed in the type descrip- tion as AMR63747. Lygisaurus rococo Ingram & Covacevich, 1988 Holotype OMJ46014; paratype OMJ42068. Data as for type description. Lygisaurus sesbrauna Ingram & Covacevich, 1988 Holotype OMJ24664; paratypes OMJ24620, OMJ24630-32, OMJ25599, OMJ25602, OMJ25615, OMJ25990, OMJ25993, OMJ26204-06, QMJ26236, OMJ26259, OMJ28057, OMJ32512, OMJ32516, OMJ34461-63, OMJ34465, OMJ34578, OMJ37508, OMJ37510, OMJ37513-20, OMJ37522-26, QMJ37530-31, OMJ38092, OMJ38299, OMJ42113. Data as for type description. Lygisaurus tanneri Ingram & Covacevich, 1988 Holotype OMJ32352; paratypes OMJ20609-11, OMJ22380, OMJ22789, OMJ24117-18, OMJ27093- 96, OMJ32358-59, OMJ32362-64, OMJ42771-72. Data as for type description. Lygisaurus zuma Couper, 1993 Holotype OMJ55760; paratypes OMJ53397, OMJ55761-67, OMJ56278-80, OMJ56765, OMJ56786, OMJ56874-76, OMJ56883. Data as for type description. Menetia koshlandae Greer, 1991 Holotype OMJ50554; paratype OMJ45800. Data as for type description. Menetia sadlieri Greer, 1991 Holotype OMJ24448. Data as for type description. This specimen is also a paratype of Menetia timlowi Ingram, 1977. Nangura spinosa Covacevich, Couper & James, 1993 Holotype OMJ55424; paratypes OMJ56029, OMJ56031, OMJ57246-47. Data as for type descrip- tion. Ophioscincus cooloolensis Greer & Cogger, 1985 Holotype OMJ31573; paratypes OMJ27381-85, OMJ31574, OMJ31578, OMJ40092, OMJ40223-26. Data as for type description. Sphenomorphus cracens Greer, 1985 — Glaphyromorphus cracens after Greer (1989). Holotype OMJ42714; paratypes OMJ1535-36, OMJ31022, OMJ31029-31, OMJ31043-49, OMJ42070-74, OMJ46261-65. Data as for type description. Family VARANIDAE Odatria keithhornei Wells & Wellington, 1985 — Varanus keithhornei - new synonymy Holotype OMJ31566. Data as for type description. This specimen is also the holotype of Varanus teriae Sprackland, 1991. Varanus telenesetes Sprackland, 1991 Holotype OMJ1190. Data as for type description. Varanus teriae Sprackland, 1991 — Varanus keithhornei (Wells & Wellington, 1985). Holotype OMJ31566; paratypes OMJ35450-51. Data as for type description. This specimen is also the holotype of Odatria keithhornei Wells & Wellington, 1985. Family TYPHLOPIDAE Ramphotyphlops chamodracaena Ingram & Covacevich, 1993 Holotype OMJ40233; paratypes OMJ28082, OMJ31963, QMJ39673, OMJ41550, OMJ51980. Data as for type description. FROG AND REPTILE TYPE SPECIMENS 77 Ramphotyphlops silvia Ingram & Covacevich, 1993 Holotype QMJ27387; paratypes QMJ8521, OMJ23620, QMJ27386, OMJ31576-77, OMJ31579, OMJ35872, OMJ43785, OMJ46128. Data as for type description. Family BOIDAE Liasis stimsoni orientalis Smith, 1985 = Antaresia stimsoni orientalis after Kluge (1993). Paratypes OMJ390, OMJ5963, OMJ7154, OMJ10256-58, OMJ10274, OMJ10444, OMJ28432, OMJ29032, OMJ32284-85, OMJ35685, OMJ39087. Data as for type description. NEW INFORMATION ON SPECIES INCLUDED IN PREVIOUS QUEENSLAND MUSEUM TYPE LISTS Class AMPHIBIA Family MYOBATRACHIDAE Mixophyes balbus Straughan, 1968 Thirteen specimens of this species are listed as in the Australian and the Queensland Museums (Straughan, 1968). Covacevich (1971) noted that the Straughan collection held in the Queensland Museum contained *M. balbus - 10 unsexed specimens, unregistered, found with tadpoles in jar labelled New England sp. nov. M. balbus, but with no other data.’ Cogger (1979) listed six M. balbus paratypes in the Australian Museum. Assuming they are paratypes, there should be only seven paratypes in the Oueensland Museum. Corben & Ingram (1987) examined the ten specimens referred to by Covacevich (1971) and noted 'these specimens are now registered as OMJ45785-45794. We have examined them and found that three (OMJ45785-7) are M. fasciolatus and the other seven (OMJ45788-94) are M. balbus.” These seven specimens are probably the missing seven paratypes, and are labelled so in the collection. Mixophyes iteratus Straughan, 1968 Two paratypes were apparently lodged in the Queensland Museum from Tweed River, Mount Warming, NSW (Straughan, 1968). Covacevich (1971) noted the Straughan collection in the Queensland Museum con- tained ‘M. iteratus - 2 unsexed specimens, QMJ18851, Lynch's Creek, Kyogle, N.S.W.; 1 unregistered specimen without data.’ Corben & Ingram (1987) noted “OMJ18851 from Lynch’s Creek, Kyogle' did not agree with the given locality of “Tweed River, Mount Warning. The other specimen (now registered as OMJ45796) had no accompanying data.” The status of these two specimens thus remains uncertain. Class REPTILIA Family GEKKONIDAE Gehyra catenata Low, 1979 Ingram & Covacevich (1981) erroneously listed OMJ28794 as a paratype. This number is for a specimen of Gehyra dubia. Oedura cincta de Vis, 1888 — Oedura marmorata Gray, 1842 after Cogger (1957). Specimen OMJ226 was listed by Covacevich (1971) as a '? syntype’. Wells & Wellington (1985) have designated this specimen as the lectotype. Phyllurus caudiannulatus Covacevich, 1975 Paratype QMJ25411. This specimen is also a paratype of P. nepthys Couper, Covacevich & Moritz, 1993. Family SCINCIDAE Ablepharus boutonii clarus Storr, 1961 = Cryptoblepharus virgatus Garman, 1901 after Cogger et al. (1983). Paratypes QMJ30920-21. The type description alludes to ‘cotypes’ and ‘paratypes’, without registration num- bers. Close to thirty years after his description, Storr identified these specimens as paratypes (Storr, in litt., 23 March 1990). Heteropus bicarinatus Macleay, 1877 = Carlia bicarinata after Ingram & Covacevich (1989). Neotype QMJ27717, designated by Ingram & Covacevich (1989). Heteropus blackmanni de Vis, 1884 78 MEMOIRS OF THE QUEENSLAND MUSEUM = Carlia vivax (de Vis, 1884) after Ingram & Covacevich (1989). Lectotype QMJ19985; paralectotypes QMJ7773?, OMJ19968-84, OMJ19986-90 Lectotype designated by Ingram & Covacevich (1989). In their discussion of the status of material associated with the type specimen history of Carlia vivax, they refer to OMJ19986-90 twice. The second reference is erroneous , as OMJ19986-99. See Covacevich (1971) for detail of doubtful type status of OMJ7773. Heteropus lateralis de Vis, 1884 — Carlia pectoralis pectoralis (de Vis, 1884) after Ingram & Covacevich (1989). Lectotype OMJ234, designated by Ingram & Covacevich (1989). Heteropus mundus de Vis, 1884 = Carlia munda after Ingram & Covacevich (1989). Neotype OMJ15654, designated by Ingram & Covacevich (1989). Lygisaurus foliorum de Vis, 1884 Neotype OMJ23660, designated by Ingram & Covacevich (1988). Menetia timlowi Ingram, 1977 Paratype OMJ24448. This specimen is also the holotype of Menetia sadlieri Greer, 1991. Mocoa delicata de Vis, 1888 = Lampropholis delicata after Mather (1990). Neotype QMJ45765, designated by Mather (1990). Mocoa spectabilis de Vis, 1888 = Saproscincus spectabilis after Wells & Wellington (1985). The type status history of specimens of this taxon is complicated. Covacevich (1971) treated QMJ244, OMJ255, OMJ19742-3 as syntypes, while noting that measurements of OMJ244 fitted those in the type description. Wells & Wellington (1985) designated OMJ19742 as the lectotype. Specimens OMJ244, OMJ255 and OMJ19743 thus became paralectotypes. Sadlier et al. (1993) recognised the work of Wells & Wellington (1985), and recommended nomenclatural changes based on it. Ingram (1994), asserted Covacevich's (1971) recognition of four syntypes was erroneous and that the type description (de Vis, 1888) was based on one specimenonly, the holotype, OMJ244. His action thus overturned the nomenclatural change recommended by Sadlier et al. (1993). Myophila vivax de Vis, 1884 = Carlia vivax after Ingram & Covacevich (1989). Neotype QMJ24176, designated by Ingram & Covacevich , 1989. Rhodona allanae Longman, 1937 = Lerista allanae after Greer (1976). The holotype of this species, *... Described from three specimens ...’, was designated (OMJ6180) by its author (Longman, 1937). However, registration numbers for the two paratypes were not listed. The question of which specimens are the paratypes has already been addressed twice, with different opinions. Covacevich (1971) found three specimens in jars with a tag marked ‘Rhodona allanae Lgmn. Paratypes’. This tag has been checked, and is in Longman’s handwriting (I. Filmer, pers. comm.). The three specimens are QMJ6040, QMJ6238 and OMJ6308. The last mentioned is incomplete, lack- ing head and some body. Assuming Longman would have mentioned a damaged specimen in his description if it were part of his type series, yet acknowledging the possibility that it may have been a type as the Longman handwritten tag sug- gested, she listed QMJ6040 and QMJ6238 as paratypes and OMJ6308 as a ‘? paratype’, signify- ing ‘possible type status’. Erroneously, she did not mention QMJ6179, stored separately from the former three specimens, but identified in the register ‘paratype’, also in Longman’s handwrit- ing. Codret & Ingram (1992) commented on this decision ‘... there is no doubt about OMJ6040, both the dates of registration of QMJ6238 and QMJ6308 (28 June, 1937 and 30 November, 1937 respectively) are after the date of publication of Longman’s paper (24 June, 1937) and thus unlike- ly to have been examined for the paper. Actually the second paratype must be QMJ6179, which was registered with the holotype and is noted as ‘paratype’ in the register.’ Data for these specimens from the register and from the tag found by Covacevich (1971), all in Longman’s hand are: FROG AND REPTILE TYPE SPECIMENS Reg. number Labelled Locality/donor/other and Date ‘paratype’? relevant data | | ar [Register| — O OMJ6040 Retro Station, Capella, OMJ6179 Retro Station, Capella, 19 May, 1937 y Old; Mrs P.C. Allan Retro, Capella, O; Mrs P.C. Allan; 'on exhibition’ Retro, Capella, Old; - Mrs P.C. Allan; ‘1/2 OMJ6238 28 June, 1937 OMJ6308 30 Nov., 1937 not invariably their true dates of publication; many type specimens have been registered after their descriptions have been published, a situation easily possible where registration numbers are not published (as is the case here); and the registration date of OMJ6179 (19 May, 1937) is less than five weeks earlier than the date of publication which appears on the type description (24 June, 1937). As a five week turn around time between submis- sion of manuscript and publication of a journal is extremely rapid by even present highly mechanised standards, it seems reasonable to sug- gest that this specimen must have been registered by Longman after his paper had gone to press. Further, the registration date of OMJ6238 is only four days after the alleged date of publication of the paper (28 June, 1937 vs 24 June, 1937). Article 72 B vii of the International Code of Zoological Nomenclature (3 ed., 1985) states ‘The mere citation of “Type” or equivalent expression, in a list of types, or in a catalogue in a museum, or on a label is not to be construed alone as evidence that a specimen is or is fixed as any of the kinds of types...”. (italics ours). Recommen- dation 72 B states ‘External evidence admitted. - If an author, in establishing a nominal species - group taxon, does not explicitly state what specimens constitute the type series, evidence in addition to published evidence may be taken into account (e.g., labels by the original author and specimens known to have been in appropriate col- lections at the appropriate time).’ ... If dates of publication and registration are con- sidered along with type designation by Longman either in the register or in a jar, three specimens are possible candidates for status as two paratypes. (Specimen QMJ6308 can be excluded from further consideration because of a late date of registration and as an incomplete specimen, despite its presence with the Longman paratype label). Of the three candidates for type status (QMJ6040, QMJ6179,QMJ6238),it seems reasonable to exclude QMJ6238 as a possible paratype because, according to the register it was 79 ‘on exhibition’ (Longman’s handwriting). This specimen, once removed from exhibition, could easily have been placed in the jar containing QMJ6040 and OMJ6179, the paratypes, along with Longman’s original paratype label. We thus confirm the correction by Couper & Ingram (1992) of Covacevich’s (1971) decision, but for reasons which differ from theirs, Family ELAPIDAE Denisonia rostralis de Vis, 1911 = Simoselaps warro (de Vis, 1911) after Mack & Gunn (1953). Holotype QMJ193. The holotype was not located by Covacevich (1971) and was listed ‘presumed lost’ by Cogger et. al. (1983) but has been discovered in the Queensland Museum collection. LITERATURE CITED BROWN, W.C. 1991. Lizards of the genus Emoia (Scin- cidae) with observations on their evolution and biogeography. Memoirs of the California Academy of Sciences 15: 1-94. CANN, J. & LEGLER, J.M. 1994. The Mary River tor- toise: anew genus and species of short-necked chelid from Queensland, Australia (Testudines: Pleurodira). Chelonian Conservation and Biology 1(2): 81-96. COGGER, H.G. 1957. Investigations in the gekkonid genus Oedura Gray. Proceedings of the Linnean Society of New South Wales 82: 167-79. 1979. Type specimens of reptiles and amphibians in the Australian Museum. Records of the Australian Museum 32: 164-210, 1992. ‘Reptiles and amphibians of Australia’. Sth ed. (Reed: Sydney). COGGER, H.G., CAMERON, E.E. & COGGER, H.M. 1983.‘Amphibia and Reptilia Vol. 1. Zoological Catalogue of Australia’. (Australian Government Publishing Service: Canberra). CORBEN, C.J. & INGRAM, G.J. 1987. A new barred river frog (Myobatrachidae: Mixophyes). Memoirs of the Queensland Museum 25: 233-237. COUPER, P.J. 1993. A new species of Lygisaurus de Vis (Reptilia: Scincidae) from mideastem Queensland. Memoirs of the Queensland Museum 33: 163-166. COUPER, P.J., COVACEVICH, J.A. & LETHBRIDGE, P. 1994. Carlia parrhasius, a new Queensland skink, Memoirs of the Oueensland Museum 35: 31-33. COUPER, P.J., COVACEVICH, J.A. & MORITZ, C. 1993. A review of the leaf-tailed geckos endemic to eastern Australia: a new genus, four new species, and other new data. Memoirs of the Oueensland Museum 34: 95-124. COUPER, P.J. & GREGSON, R.A.M. 1994, Redescnption of Nephrurus asper Günther, and description of N. amyae sp. nov. and N. sheai sp. 80 MEMOIRS OF THE QUEENSLAND MUSEUM nov. Memoirs of the Queensland Museum 37(1): 67-81. COUPER, P.J . & INGRAM, G.J. 1992. A new species of skink of Lerista from Oueensland and a re-ap- praisal of L. allanae (Longman). Memoirs of the Oueensland Museum 32: 55-59. COVACEVICH, J. 1971. Amphibian and reptile type- specimens in the Queensland Museum. Memoirs of the Queensland Museum 16: 49-67. 1975. A review of the genus Phyllurus (Lacertilia: Gekkonidae). Memoirs of the Queensland Museum 17: 293-303. 1984. A biogeographically significant new species of Leiolopisma (Scincidae) from north eastern Queensland. Memoirs of the Queensland Museum 21: 401-411. COVACEVICH, J.A., COUPER, P.J. & JAMES, C. 1993. A new skink, Nangura spinosa gen. et sp. nov., from a dry rainforest of southeastern Queensland. Memoirs of the Queensland Museum 34: 159-167. CZECHURA, G.V. 1986. A new species of Taudac- tylus (Myobatrachidae) from southeastern Queensland, Australia. Memoirs of the Queensland Museum 22: 299-307. 1986. Skinks of the Ctenotus schevilli species group. Memoirs of the Queensland Museum 22: 289- 297. CZECHURA, G.V. & WOMBEY, J. 1982. Three new striped skinks, (Ctenotus, Lacertilia, Scincidae) from Queensland. Memoirs of the Queensland Museum 20: 639-645. DAVIES, M., MARTIN, A.A. & WATSON, G.F. 1983. Redefinition of the Litoria latopalmata species group (Anura: Hylidae). Transactions of the Royal Society of South Australia 107: 87-108. DAVIES, M., MCDONALD, K.R. & ADAMS, M. 1986. A new species of green tree frog (Anura: Hylidae) from Queensland, Australia. Proceed- ings of the Royal Society of Victoria 98: 63-71. DAVIES, M., MCDONALD, K.R. & CORBEN, C. 1986. The genus Uperoleia GRAY (Anura: Lep- todactylidae) in Queensland, Australia. Proceed- ings of the Royal Society of Victoria 98: 147-188. DAVIES, M., WATSON, G.F., MCDONALD, K.R., TRENERRY, M.P. & WERREN, G. 1993. A new species of Uperoleia (Anura: Leptodactylidae: Myobatrachinae) from northeastern Australia. oe of the Queensland Museum 33: 167- 4. DE VIS, C.W. 1884. A concept of the genus Heteropus. Proceedings of the Royal Society of Oueensland 1: 166-173. 1888. A contribution to the herpetology of Oueensland. Proceedings of the Linnean Society of New South Wales 2: 811-826. 1911. Descriptions of snakes apparently new. An- nals of the Oueensland Museum 10: 22-25. GREER, A.E. 1976. A new generic arrangement for some Australian scincid lizards. Breviora 267: 1-19. 1983. The Australian scincid genus Calyptotis, de Vis: resurrection of the name, description of four new species, and discussion of relationships. Records of the Australian Museum 35: 29-59. 1985. A new species of Sphenomorphus from north- eastern Queensland. Journal of Herpetology 19: 469-473. 1989. ‘The biology and evolution of Australian lizards’. (Surrey Beatty & sons: Chipping Nor- ton). 1991. Two new species of Menetia from north- eastern Queensland, with comments on the generic diagnoses of Lygisaurus and Menetia. Journal of Herpetology 25: 268-272. 1992. Revision of the species previously associated with the Australian scincid lizard Eulamprus tenuis. Records of the Australian Museum 44: 7-19. GREER, A.E. & COGGER, H.G. 1985. Systematics of the reduce-limbed and limbless skinks currently assigned to the genus Anomalopus (Lacertilia: Scincidae). Records of the Australian Museum 37: 11-54. GREER, A.E., MCDONALD, K.R. & LAWRIE, B.C. 1983. Three new species of Lerista (Scincidae) from northern Queensland with a diagnosis of the wilkinsi species group. Journal of Herpetology 17: 247-255. HUTCHINSON, M.N., DONNELLAN, S.C., BAVERSTOCK, P.R., KRIEG, M., SIMMS, S. & BURGIN, S. 1990. Immunological relation- ships and generic revision of the Australian lizards assigned to the genus Leiolopisma (Scincidae: Lygosominae). Australian Journal of Zoology 38: 535-554. INGRAM, G.J. 1977, Descriptions of three small lizards - two of them new. Genus Menetia (Lacertilia, Scincidae) in Queensland. Victorian Naturalist 94: 184-187. 1991, Five new skinks from Queensland rainforests. Memoirs of the Queensland Museum 30: 443- 453. 1994. The holotype of Macoa spectabilis de Vis, 1888. Memoirs of the Queensland Museum 35: 34, INGRAM, G. & CORBEN, C. 1990. Litoria electrica: a new treefrog from western Queensland. Memoirs of the Queensland Museum 28: 475- 478. 1994, Two new species of broodfrogs (Pseudo- phryne) from Queensland. Memoirs of the Queensland Museum 37: 267-272. INGRAM G.J., CORBEN, C. & HOSMER, W. 1982. Litoria revelata: a new species of tree-frog from eastern Australia. Memoirs of the Queensland Museum 20; 635-637. INGRAM, GJ., COUPER, P.J. & DONNELLAN, S.C. 1993. A new two-toed skink from eastern Australia. Memoirs of the Queensland Museum 33: 341-347, INGRAM, G. & COVACEVICH, J. 1981. Frog and reptile specimens in the Queensland Museum, with a check- FROG AND REPTILE TYPE SPECIMENS 81 list of frogs and reptiles in Queensland. Memoirs of the Queensland Museum 20: 291-306. 1988. Revision of the genus Lygisaurus de Vis (Scin- cidae: Reptilia) in Australia. Memoirs of the Queensland Museum 25: 335-354. 1989. Revision of the genus Carlia (Reptilia, Scin- cidae) in Australia with comments on Carlia bicarinata of New Guinea. Memoirs of the Queensland Museum 27: 443-490. 1993. Two new species of striped blindsnakes. Memoirs of the Oueensland Museum 34: 181-184. INGRAM, G.J. & CZECHURA, G.V. 1990. Four new species of striped skinks from Oueensland. Memoirs of the Queensland Museum 29: 407-410. INGRAM, G. & EHMANN, H. 1981. A new species of scincid lizard of the genus Leiolopisma (Scincidae: Lygosominae) from southeastern Queensland and northern New South Wales. Memoirs of the Queensland Museum 20: 307-310. INGRAM, G. & RAWLINSON, P. 1981. Five new species of skinks (genus Lampropholis) from Queensland and New South Wales. Memoirs of the Queensland Museum 20: 311-317. KLUGE, A.G. 1993. Aspidites and the phylogeny of pythonine snakes. Records of the Australian Museum, Supplement 19: 1-77. LEGLER, J.M. & CANN, J. 1980. A new genus and species of chelid turtle from Queensland, Australia. Contributions in Science Natural History Museum of Los Angeles County 324: 1-18. LONGMAN, H.A. 1937. Herpetological notes. Memoirs of the Queensland Museum 11: 165-168. LOW, T. 1979. A new species of gecko, genus Gehyra (Reptilia: Gekkonidae) from Queensland. Victorian Naturalist 96: 190-196. MACK, G. & GUNN, S.B. 1953. De Vis’ types of Australian Snakes. Memoirs of the Queensland Museum 13; 58-70. MACLEAY, W. 1877. The lizards of the ‘Chevert’ Ex- pedition. Proceedings of the Linnean Society of New South Wales 2: 60-69. MAHONY, M., TYLER, M.J. & DAVIES, M. 1984. A new species of the genus Rheobatrachus (Anura: Leptodactylidae) from Queensland. Transactions of the Royal Society of South Australia 108: 155-162. MATHER, P.B. 1990. Electrophoretic and morphological comparisons of Lampropholis delicata (Lacertilia: Scincidae) populations of eastem Australia, and a resolution of the taxonomic status of this species. Australian Journal of Zoology 37: 561-574. RICHARDS, S.J. 1992. A new species of microhylid frog (genus Cophixalus) from the Star Mountains, central New Guinea. Science in New Guinea 18: 141-145. RICHARDS, S.J., DENNIS, A.J., TRENERRY, M.P. & WERREN, G.L. 1994. A new species of Cophixalus (Anura: Microhylidae) from northern Oueensland. Memoirs of the Oueensland Museum 37(1): 307- 310. RICHARDS, S.J. & JOHNSTON, G. R. 1993. A new species of Nyctimystes (Anura: Hylidae) from the Star Mountains, Papua New Guinea. Memoirs of the Oueensland Mustum 33: 73-76. RICHARDS, S.J., JOHNSTON, G.R. & BURTON, T.C. 1994. A remarkable new asterophryine microhylid frog from the mountains of New Guinea. Memoirs of the Oueensland Museum 37(1): 281-286. SADLIER, R.A. 1988. Bavayia validiclavis and Bavayia septuiclavis, two new species of gekkonid lizard from New Caledonia. Records of the Australian Museum 40: 365-370. SADLIER, R.A., COLGAN, D.J. & SHEA, G.M. 1993. Taxonomy and distribution of the scincid lizard Saproscincus challengeri and related species in southeastern Australia, Memoirs of the Queensland Museum 34: 139-158. SHEA, G.M. 1987. Two new species of Delma (Lacertilia: Pygopodidae) from northeastern Queensland and a note on the status of the genus Aclys. Proceedings of the Linnean Society of New South Wales 109: 203- 212. SMITH, L.A. 1985. A revision of the Liasis childreni species-group (Serpentes: Boidae). Records of the Westem Australian Museum 12: 257-276. SPRACKLAND, R.G. 1991. Taxonomic review of the Varanus prasinus group with descriptions of two new species. Memoirs of the Queensland Museum 30: 561-576. STORR, G.M. 1961. Ablepharus boutonii clarus, a new skink from the Esperance district, Western Australia. Westem Australian Naturalist 7: 176-178. 1981. Ten new Ctenotus (Lacertilia: Scincidae) from Australia. Records of the Western Australian Museum 9: 125-146. 1986. A new species of Lerista (Lacertilia: Scincidae) with two subspecies from central Australia. Records of the Western Australian Museum 13: 145-149, 1991. Revision of Lerista orientalis (Lacertilia: Scin- cidae) of northem Australia. Records of the Western Australian Museum 15; 413-417. STRAUGHAN, I.R. 1968. A taxonomic review of the genus Mixophyes (Anura, Leptodactylidae). Proceedings of the Linnean Society of New South Wales 93: 52-59. VAN BEURDEN, E. & MCDONALD, K.R. 1980. A new species of Cyclorana (Anura:Hylidae) from northern Oueensland. Transactions of the Royal Society of South Australia 104: 193-195. WELLS, R.W. & WELLINGTON, C.R. 1985. A clas- sification of the Amphibia and Reptilia of Australia. Australian Journal of Herpetology, Supplementary series 1: 1-161. WITTEN, G.J. 1994. Taxonomy of the agamid genus Pogona (Reptilia: Lacertilia). Memoirs of the Queensland Museum 37(1): 329-343, ZWEIFEL, R.G. 1985. Australian frogs of the family Microhylidae. Bulletin of the American Museum of Natural History 182: 265-388. 82 MEMOIRS OF THE QUEENSLAND MUSEUM NEW PHASCOGALE TAPOATAFA (BRUSH-TAILED PHASCOGALE) RECORDS FROM CAPE YORK PENINSULA, AUSTRALIA. Memoirs of the Queensland Museum 37(1):82. 1994:- Phascogale tapoatafa is a species rarely seen, and sometimes difficult to find, even by spotlight- ing and trapping (Traill & Coates, 1993). Recent work (Sharpe, 1993) has shown that ‘.,. in prime habitat, one in- dividual every 35ha is the norm. `... It is widely, but patchily, distributed in coastal areas of Australia with an average annual rainfall of between 500 and 2000mrn (Cuttle, 1982; Cuttle in Strahan, 1983). In Oueensland, it is well Known in the southeast, but specimens or reliable sightings of P. tapoatafa in other parts of the state are scant. There are records from the central coastal area (Rockhampton, 23722'S 150732'E: In- gram & Raven, 1991); and from northem Cape York Penin- sula (Coen, 13°52’S 143°12’E; Lockhart R. area 13°04'S 143°24"E: Cuttle, 1982; Dixon & Huxley, 1985); and from skeletal remains in limestone caves at Chillagoe, on southeastern Cape York Peninsula, 177098 144931'E: Tresize, 1970. There is also one doubtful record from the western coast of Cape York Peninsula (Edward R. 14*39'S 142703'E: D. Thomson, 1929, p.29 in Dixon & Huxley, 1985). This apparent sight record conflicts with data in the same reference (p.14): `u Phascogale tapoatafa ... not uncommon in places on east coast; not seen on western fall of central range. ’... Some recent specimen and sight records shed light on the occurrence of this elusive species on Cape York Peninsula: Nr McLeod Ck, Mclvor R. Rd, 14°16'S 144727'E, killed by cat, specimen not retained, I Sept 1976, L. Kuhneman; Iron Ra,, 12°43’S 143°17'E, Jun 1978, G. Czechura, sight only; Mt. Molloy, 2.5km N of Rifle Ck, 16°40'S 145°20°E, 1 Aug 1986, L. Moore & F. Crome, sight only; Emerald Ck on Cairns-Mareeba Rd, 16°58'S 145*28'E, 17 Jun 1992, J. Grant, sight only; Lamb Ra., Emu Ck Rd, 17°03'S 145928'E, 5 Aug 1993, D.L. Storch, sight only; Lamb Ra., Emu Ck Rd 17°04'S 145°30°E, 5 Aug 1993, D.L. Storch & N. Goudberg, sight only, QMJM10591, dead on road, 20km W of Cooktown on Mclvor R. Rd, 15726'S 145°03'E , Nov 1993, L. Roberts; QMJM10579 dead on road, 8km S of McIvor R., Mclvor Rd 15920'S 145704'E, 13 Jun 1994, J. Covacevich, L, Roberts; Bubardt Pt, Lockhart R, 12°58°S 143930'E, no date recorded, W. Butcher, in Leung et al., 1994, One of the two Melvor Rd specimens (OMJM10591) was found on a road which runs between a narrow band of riverine rainforest and open woodland, The second (OMJM10579) was found dead on that road in sparse open woodland. Both Lamb Ra, observations were made in open forests on soils derived from granite. Ground cover is sparse in the area. Data on vegetation are not available for other localities at which specimens have been collected or observed. Literature Cited Cuttle, P. 1982. Life history strategy of the dasyund marsupial Phascogale tapoatafa.Pp.13-22. In Archer, M. (ed.), ‘Carnivorous marsupials’. Vol. 1. (Royal Zoological Society of New South Wales: Sydney). Dixon, J.M. & Huxley, L. (eds). 1985. ‘Donald Thomson's mammals and fishes of northern Australia’. (Thomas Nelson Australia: Melbourne). Ingram, G.J. & Raven, R.J. (eds). 1991, ‘An atlas of Oueensland's frogs, reptiles, birds and mammals’. (Board of Trustees, Oueensland Museum, Brisbane). Leung, L.K., Venables, B. & Pritchard, J. 1994. Terrestrial vertebrate fauna survey of the Iron Range area, Cape York Peninsula, 1993-1994. A report to the Queensland Department of Environment and Heritage.Un- published, Sharpe, L. 1993. The return of a little predator, Australian Natural History 24(7): 22-29. Strahan, R. (ed.). 1983. “The Australian Museum complete book of Australian mammals’. The National Photographic Index of Australian wildlife. (Angus & Robertson: Sydney). Tresize, J. 1970. Bone deposits in Chillagoe Mungana Caves. Journal of the North Queensland Naturalists Club 37(152): 2-4. Traill, B.J. & Coates, T. 1993, Field observations on the Brush-tailed Phascogale Phascogale 1opoatafa (Mar- supialia: Dasyuridae). Australian Mammalogy 16(1): 61-66. J.A. Covacevich, Queensland Museum, Box 3300, South Bris- bane, Queensland 4101, Australia; L. Roberts, Shipton's Flat, via Cooktown, Queensland 4871, Australia; D.L. Storch, Queensland Department of Environment and Heritage, Box 2066, Cairns 4871, Australia; S. Van Dyck, Queensland Museum, Box 3300, South Brisbane, Queensland 4101, Australia; 10 September 1994. A NEW SPECIES AND A NEW RECORD OF HEPATOPORUS FROM NORTH- WESTERN AUSTRALIA (CRUSTACEA: DECAPODA: XANTHIDAE) P.J.F. DAVIE AND P.A. TURNER Davie, P.J.F. & Turner, P.A, 1994 12 01: A new species and a new record of Hepatoporus from northwestern Australia (Crustacea: Decapoda: Xanthidae). Memoirs of the Queensland Museum 37(1):83-86, Brisbane. ISSN 0079-8835, Hepatoporus asper sp,nov., is described from the North-West Shelf, Western Australia, It is separated from other Hepatoporus by its carapace regions being separated by deep channels lined with mushroom shaped tubercles. H. guinotae (Zarenkov, 1971) is recorded for the first time from Australian waters, greatly extending its range from west Africa and the Red Sea. It is suggested that H. distinctus (Takeda & Nagai, 1986) is of uncertain validity and may prove to be a junior synonym of H. guinotae. | ] Crustacea, Decapoda, Brachyura, Xanthidae, Euxanthinae, Hepatoporus, Australia, new species, distribution. P.J.F. Davie & P.A.Turner, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia; 2 May 1994, Two species of the xanthid Hepatoporus were identified in dredged material from the North- West Shelf, collected by the CSIRO aboard the R.V. ‘Soela’. One, H. guinotae (Zarenkov, 1971) marks a considerable range extension and is a new record for Australia; the other is a new species, H. asper. Hepatoporus was established by Serêne (1984) io include Carpoporus orientalis Sakai, 1935, from Japan, and C. guinotae Zarenkov, 1971, from the Red Sea and the westem Indian Ocean. Sakai (1935) and Zarenkov (1971) had both noted that the closest relative to their species was Car- poporus papulosus Stimpson, 1871, a West At- lantic species. Takeda & Nagai (1986) independently recognised the generic differences between C. papulosus and the two Indo-West Pacific species, and described the new genus Carpoporoides for them, as they did not know of the work of Seréne (1984). Takeda & Nagai (1986) also described a new species, Car- poporoides distinctus, from Koza, Japan. Later, Takeda (1986) recognised Carpoporoides as a junior synonym of Hepatoporus. Hepatoporus now contains four species: H. orientalis (Sakai, 1935), H. guinotae (Zarenkov, 1971), H. distincrus (Takeda & Nagai, 1986) and H. asper sp. nov. Measurements given in the text are of the carapace breadth (measured at the widest point) followed by length. ABBREVIATIONS: G1, gonopod 1; QM, Queensland Museum, Brisbane; P1-P5, pereiopods 1-5. SYSTEMATICS Family XANTHIDAE MacLeay, 1838 Sub-family EUXANTHINAE Alcock, 1898 Hepatoporus guinotae (Zarenkoy, 1971) (Fig. 1A, B) Carpoporus guinolae Zarenkov, 1971; 191, tig. 86. Hepatoporus guinotae: Seréne, 1984: 74, 75, fig. 40, pl. 10, d-f; Takeda, 1986; 51. MATERIAL EXAMINED OMWI5071, 9 (8.3 x 6.3mm), R.V. 'Soela', Stn OIBISNT, 22.2.1983, 52m, 20°01.4'S, 116*57.3'E, North-West Shelf, Western Australia; QMW14778, 3(3.1x 2.6mm), R.V. ‘Soela’, Sin 02B02S, 22.4.1983, 43m, 19°56.9°S, 117°53.7'E, North-West Shelf, Western Australia. REMARKS The hepatic cavities of the two specimens show different states of development which may be age rather than sex related. The small male has only weakly developed cavities that make only slight impressions on the anterolateral margins, whereas those of the larger female are close to the state seen in Serêne's illustration of a male from Kenya (Seréne 1984: pl. 10d,e,f). The posterior part of the hepatic cavity is not as deeply ex- cavated posteriorly as in Serêne's specimen, but is much more deeply excavated than on the holotype of H. distinctus. Since the depth and shape of the hepatic cavity is the only sig- nificant character separating H. guinotae and H. 84 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 1. A, B, Hepatoporus guinotae (Zarenkov), QMW 15071, 9(8.3 x 6.3mm); C, D, Hepatoporus asper sp. nov., OMW19896, & holotype (9.1 x 8.7mm). distinctus Takeda & Nagai, 1986, we would be surprised if this later species proves to be validly separable; a greater range of material is needed to decide the matter. DISTRIBUTION Red Sea (type locality, Zarenkov, 1971); Madagascar, Kenyan Coast (Seréne, 1984); and northwestern Australia. Bathymetric range: 108m (Seréne, 1984) and 42-52m (this paper). Hepatoporus asper sp. nov. (Figs 1C, D; 2A, B) MATERIAL EXAMINED HOLOTYPE: QMW19896, â (9.1 x 8.7 mm), R.V. ‘Soela’, Stn 05B03BT, 26.10.1983, 40m, 19*55'S, 117°56.0°E, North-West Shelf, Western Australia. DESCRIPTION Carapace. Carapace heptagonal in outline, length 0.97 width. Dorsal surface formed by a pavement of abutting mushroom-shaped tubercles, eroded in appearance, with prominent raised pair of sub-conical, sub-median gastric prominences and moderately inflated branchial regions. Cardiac region uniformly convex, smoother but less raised than branchials. Lateral margins with 2 marked concavities, anterior hepatic and lateral branchial; dorsal surface broadly excavated obliquely behind posterior cavity. Gastro-cardiac regions separated from branchials by deep sulci; sulci forked anteriorly around hepatic regions; sulci with marginal mushroom shaped tubercles. Deep median sulcus separating frontal lobes, extending posteriorly to between gastric prominences. Deep, often inter- connecting, pits on most regions. Intestinal region bears 7 small, obtuse teeth at its edge. Frontal width 0.37 X width of carapace; deflexed, pitted; bilobed, with inner projections forming basal circular hole. Postero-lateral mar- gins concave, Antennular fossae broad, oblique; basal seg- ment of antennae deeply pitted; epistome and anterior pterogostomial region pitted; posterior A NEW RECORD OF HEPATOPORUS 85 FIG. 2, First male gonopod of Heparoporus asper sp. nov., holotype, OMW19896. A, abdominal view; B, enlarged view of apex. half of pterogostomial and sub-hepatic regions coarsely granular. Outer edge of orbit with several short, simple setae on inside edge; orbital margin relatively smooth, with few sparse, Jow, obtuse lobes. Third maxillipeds. Surface coarsely pitted, some joining to form irregular longitudinal chan- nels. Ischium c, 2 xX length of merus. Proximal half of exopod smooth, separated from pitted distal portion by smooth oblique ndge. Thorax and abdomen. Thoracic stemites deep- ly eroded. Abdominal segments 3 - 5 fused, sur- face formed by pavement of abutting mushroom-shaped tubercles. Chelipeds. Equal, short, length subequal carapace length. Fingers with tips crossing when closed. Upper surface of dactyl tuberculate proximally, becoming smooth distally. Outer sur- face of prodopus appearing coarsely granular, with more or less coalesced low, mushroom-like tubercles; ventral surface with small, smooth tubercles grading into smaller more sparsely dis- persed granules on inner surface. Upper and outer surface of carpus similar to palm; inner and dorsal surface granular. Ventral surface of merus smooth; inner surface sparsely granular; row of c. 4 large tubercles along the inner disto-ventral angle of merus; laterally projecting oval concave lamella present disto-medially. Inner-dorsal angle of carpus and merus bearing long plumose setae, Ischium and coxa with less dense setae. Ambulatory legs, Short, P2 0.8 x length of cheliped. Meri of P2 and P3 completely hidden below lateral extensions of carapace, trigonal in cross-section; carpi and prodopi sub-trigonal; dactyli cylindrical, sparsely granular, with plumose setae and acute chitinous tips. Outer surface of carpi and prodopi similar to outer sur- face of chelae, Outer surfaces of meri smooth except for P5. Three to five tubercles on the ventro-proximal margins of men, reducing in size from P2-P5. Legs with marginal plumose setae, longest on dorso-proximal margins of meri. Gl(Fig. 2A, B). Stout; curved distally, slowly tapering. c. 18 sub-terminal fine, plumose, setae not extending beyond apex; small sub-distal lobe present on inner face; numerous stout, proximally directed, conical setae present on inner and upper faces, several with distinct sockets. REMARKS For the purpose of this comparison, Hepatoporus distinctus is treated as indistin- guishable from H. guinotae for reasons given under that species. H. asper differs most con- spicuously from other species of the genus by the following characters; 1, H. asper possesses a concave lateral branchial cavity posterior to the hepatic cavity; 2, the dorsal surface of the carapace in H., asper has the regions well separated by deep channels lined with mush- room-shaped tubercles whereas both H. guinotae and H. erientalis have a much smoother, punctate, or minutely granular surface, with only shallow depressions separating the regions; 3, H. 86 MEMOIRS OF THE QUEENSLAND MUSEUM asper bears only two gastric prominences whereas H. guinotae bears large tubercles on the supraorbital, gastric, and branchial regions, and the gastric, epibranchial, and cardiac regions of H. orientalis are ‘protuberant’ (Sakai, 1935); 4, the chelipeds and walking legs in H. asper bear flattened, often coalesced, fungiform tubercles, giving a coarsely tuberculate appearance whereas on H. guinotae and H. orientalis, they are merely granular; 5, the third maxillipeds of H. asper are covered by small, densely packed, often intercon- necting pits whereas those of H. guinotae and H. orientalis are only granular; 6, the thoracic ster- num of H. asper has deep erosions that occupy a large portion of each sternite; 7, the abdomen of H. asper bears low, fungiform tubercles, that of H. guinotae is only granular. Sakai (1935) gives no indication of the nature of the sternum and abdomen in H. orientalis; 8, the G1 of H. asper (Fig. 2A, B) is distinctly different from that il- Justrated by Seréne (1984: fig. 40) for H. guinotae. The new species bears c. 18 subrer- minal setae that barely reach past the tip. In contrast, the pleopod of H. guinotae bears seven setae that extend well beyond the tip. Also the inner face of the G1 of H. asper bears a broad, subterminal lobe, which is absent in H. guinotae, The G1 is not known for H. orientalis. ETYMOLOGY From the Latin asper = rough or uneven, refer- ring to the deeply pitted and channelled dorsal surface of the carapace. DISTRIBUTION & HABITAT Only known from northwestern Australia, Dredged from 40m. ACKNOWLEDGEMENTS The present paper is part of an ongoing study of Australian Xanthoidea supported by a grant from the Australian Biological Resources Study. LITERATURE CITED ALCOCK, A. 1898. Materials for a Carcinological Fauna of India. No. 3. The Brachyura Cyclometopa. Part I. The Family Xanthidae. Jour- nal ofthe Asiatic Society of Bengal 67 (2):67-233. MACLEAY, W.S. 1838. On the brachyurous decapod Crustacea brought from the Cape by Dr Smith. Pp. 53-71. In Smith, A. (ed.), 'Hlustrations of the Annulosa of South Africa; being a portion of the objects of nalural history collected during an ex- pedition into the interior of South Africa, under the direction of Dr Andrew Smith, in the years 1834, 1835, and 1836; fitted out by “The Cape of Good Hope Association for Exploring Central Africa'”. (London). SAKAI, T. 1935. New or rare specics of Brachyura, collected by the “Misago” during the zoological survey around the Izu-Peninsula, Science Reports of the Tokyo Bunrika Daigaku, B, 2(32): 63-88. SERENE, R. 1984. Crustacés Décapodes Brachyoures de l'Ocean Indien Ocidental et de la Mer Rouge, Xanthoidea: Xanthidae et Trapeziidae. Avec un addendum par Crosnier, A.; Carpiliidae et Menip- pidae. Faune Tropicale. Office de la Recherche Scientifique et Technique Outre-Mer Paris 24: 1-400. STIMPSON, W. 1871. Preliminary report on the Crus- tacea dredged in the Gulf Stream in the Straits of Florida, by L.F. de Pourtales, assistant United States Coast Survey, Part I. Brachyura, Bulletin of the Museum of Comparative Zoology, Harvard 2(2): 109-160. TAKEDA, M. 1986. Carpoporoides Takeda et Nagai, 1986, a synonym of Hepaïoporus Serêne, 1984 (Crustacea, Decapoda, Brachyura). Proceedings of the Japanese Society of Systematic Zoology, Tokyo 33; 51, TAKEDA, M. & NAGAI, S. 1986, Establishment of a new genus on two Japanese crabs, Zoological Science, Tokyo 3(3): 547-550, ZARENKOV, N.A. 1971. On the species composition and ecology of the Decapoda in the Red Sea. Pp. 155-203. In Vodyanitzkii, V.A.(ed.), 'Benthos of the Continental Shelf of the Red Sea’. (Naukova Dumka: Kiev). MID-DEVONIAN CHONDRICHTHYAN SCALES FROM THE BROKEN RIVER, NORTH QUEENSLAND, AUSTRALIA ALISON M. DE POMEROY De Pomeroy, A.M. 1994 1201: Mid-Devonian chondrichthyan scales from the Broken River, north Queensland, Australia. Memoirs of the Queensland Museum 37(1):87-114. Brishane, ISSN 0079-8835, Chondrichthyan scales from mid Emsian to earliest Frasnian of the Dosey-Craigie Platform, Broken River region, northern Queensland include three new form genera, Gondwanalepis, Notiolepis, and Aussilepis, each represented by new species: C. grossi, N. dienemos, and A. lukaso. Also present are Cladolepis sp, cf. C. gunnelli, and scales tentatively referred to Ohiolepis sp. The diverse fauna includes acanthodian, crossopterygian, palaconiscoid and thelodont scales; placoderm platelets; onychodontid, palaeoniseoid and indeterminate chondrichthyan teeth; a dipnoan toothplate; and bone fragments of various affinities] Chondrichthyan, Devonian, Eifelian, Givetian, Queensland. Alison M. De Pomeroy, Cenire for Ecostratigraphy and Palaeobiology, School of Earth Sciences, Macquarie University, New South Wales 2109, Australia; 19 April 1994. The Broken River Group of north Oucensland, covering approximately 320km?, is dated Jate Early to earliest Late Devonian; the biochronol- ogy of the area is described by Mawson & Talent (1989). The Dosey-Craigie Platform, from where the new material was collected, is the southernmost of two shallow marine mixed car- bonate and siliciclastic shelf sequences in the area (Fig. 1). Numerous sections of the Dosey-Craigie Plat- form have been sampled over recent years, leached with acetic acid, and examined for con- odonts by the Mawson-Talent team of the Mac- quarie University Centre for Ecostratigraphy and Palaeobiology (MUCEP); abundant microver- tebrate remains have been recovered from these residues, A diverse fauna has been recognised; acanthodian, chondrichthyan, crossopterygian, palaeoniscoid and thelodont scales; placoderm platelets; onychodontid, palaeoniscoid and in- determinale chondrichthyan teeth; è dipnoan toothplate; and bone fragments of various af- finities. A description of the chondrichthyan scales is given here; descriptions of other taxonomic groups are in progress. Conedont determinations (Mawson & Talent, 1989) for the mid-Emsian to Jate-Givetian interval give precise ages for horizons containing the scales. Fish remains from the Dosey-Craigie Platform occur in several stratigraphic units: the Papilio Formation (shales with subordinate siltstones and nodular limestones - Givetian), the Spanner Limestone Member of the Papilio Formation (bioclastic, well-bedded and frequently nodular limestone - wercus Conodont Zone), the Stanley Limestone Member of the Mytton Formation (bioclastic limestone - late Givetian to earliest Frasnian, late hermanni-cristaius 10 early asym- metricks conodont zones), the Lomandra Lime- stone (mostly calcarenites and calcisiltites - Emsian-Eifelian, serotinus to costatus conodont zones), the Bracteata Formation (mudstones and lithofeldspathic sandstones, late Emsian serotinus-patulus conodont zones), and the Dosey Limestone (calcarenties and calsiltites, late Hifelian-early Givetian, kockelianus-ensen- sis conodont zones), Mawson & Talent (1989, fig. 2) summarised the stratigraphic relationships between these units, The Papilio Formation and associated Spanner Limestone Member contain by far the most abundant fish microfossils. These sediments were laid down in deeper water than, for example, the Lomandra and Dosey Lime- stones, formed under shallow water conditions or possibly sometimes exposed (Mawson & Talent, 1989). The chondrichthyan scales described herein occur in thirteen sectyuns from the Dosey-Craigie Platform succession - SD15, SDI28, SD130, SD131, SD146, SD164, SD170, SD190, SD192, SD196, SD204, SD210, and SD216 (Fig. 2). Table 1 lists the geographical location of the sections, All the scales occur in horizons dated kockelianus to hermenni-cristalus conodont zones, with two forms extending into the Frasnian asymmetricus Conodont Zone (Fig.3). In addition to possible changes in the fauna through time, this distribution was probably influenced by facies differences (see above). Most taxa have a range spanning all or part of vareuy Conodont Zone, the age of horizons in the richly fossiliferous Papilio Formation. ss MEMOIRS OF THE OUEENSLAND MUSEUM 1982 and Xenacanthus sp. from the Early Car- boniferous. Two new an- tiarchs, Wurungulepis denisoni and Nawagiaspis wadeae, were described by z ‘ casucs 7 === Young (1990) from the F nee) == === Broken River Group of the Sas. <==" a eee southern (Dosey-Craigie) = (ee | rtrd sever Torna from = === reporte SEE un the immediate area: an early phoebodont tooth from the Papilio Formation; Cheiracanthoides comptus Wells, 1944 scales, onychodontid teeth, palaeoniscoid remains and new shark scales from Fish Hill; turiniids and scales resembling nikoliviids from the Broken River Group; and endemic turiniids, buchanosteid and possibly rhenanid scales, nostolepid scales and platelets and onychodontid teeth from the pesavis-sulcatus con- odont zones of the underly- ing Martins Well Limestone. Remains of chondrich- thyans and other groups have been reported from the surrounding region. Turner (1991: fig. 5 i, j) noted and illustrated shark scales and teeth in Middle and Late Devonian limestones FIG, 1. Palaeogeographic and lithofacies relationships of the Broken River Group from the Broken Riverarea, È rk A — J ey ' Ae ANa y mOrago, — calcareous — g— - (— — argillaceous. faces ar ~ {Papilio Fm) __ = — =; T yr Dosey / —- . h ae g km py L 1 Mwdstane Ll Sandstone during Givetian times (from Mawson and Talent, 1989: fig. 3). and reported (Turner, , 1993) endemic thelodonts and No chondrichthyan scales have been formally Turinia australiensis described from the Early or Middle Devonian Gross, 1971 from Lochkovian horizons of the of the Broken River Group, although scalesand Broken River Embayment. teeth have been found in acid-leached residues Early and Middle Devonian chondrichthyans (Turner, 1991; 1993). Turner (1982) described have been reported in the literature from other and illustrated shark teeth from the Late areas of Australia and overseas; forms described Devonian and Early Carboniferous from the as chondrichthyan are summarised (Tables 2, 3). northern (Pandanus) platform: Thrinacodus lt should be noted, however, that the chondrich- ferox Turner, 1982, Phoebodus cf. P. politus thyan affinities of some of these forms have sub- Newberry, 1889, and three species of Sequently been questioned. Protacrodus, all dated as “probably Famen- Localities and sections bear the prefix “SD” for nian” - and Stethacanthus themasi Turner, Storm Dam (Fig. 2). Specimens are housed in the MID-DEVONIAN CHONDRICHTHYAN SCALES 89 palaeontological collections of the Queensland Museum (OMF). SYSTEMATIC DESCRIPTION Subclass CHONDRICHTHYES Infraclass ELASMOBRANCHII REMARKS The scales described below are interpreted as chondrichthyan by the presence of neck canal openings and a bony diamond-shaped base, char- acters considered diagnostic for the group (Turner, 1991). It is further assumed for the present that the taxa described below belong within the Elasmobranchii, because of overall similarity of scale morphology to that of articu- lated remains known to belong to elasmobranchs, such as Antarctilamna prisca Young, 1982. Gondwanalepis gen. nov. ETYMOLOGY From Gondwana, and the Greek ‘lepis’ = scale. DIAGNOSIS Crown subrhombic or rounded subtriangular, bears eight short parallel ridges deeply dissecting the anterior edge. Posteriorly, crown overhangs base a short way. Neck indented al posterior. Base diamond-shaped or subrectangular, flared into a narrow rim around edges, and gently convex, flat, or gently concave. Six to twelve elliptical open- ings in posterior neck area. REMARKS Gondwanalepis is presumed to be a chondrich- thyan because the scales have a diamond-shaped, flat or gently convex base, several neck canal openings at the posterior, and low, narrow. con- Mytlnn Fonmalien- mudstone and aranitee, with colic Stanley Marta d Papilio Formation mudstones with subordinate nodular limestone; includes Spanner Limastahe Membar Dosey Formation Stormy Mill Sandstono-aranites with) sybardinsle rudites Lomandrs Limestone mudstones with minor arenites and rare zarbnnates = Braeam- Formatia FIG. 2. Broken River Group in the Dosey-Craigie Platform area showing location of stratigraphic sections from which chondrichthyan scales have been recovered (after Mawson & Talent, 1989), 90 MEMOIRS OF THE OUEENSLAND MUSEUM TABLE 1, Geographic Iocalit;es of the sections mentioned in the text. | Section | Geographic Localit (North) - Section commences in small gully tributary to Bracteata Creek at WANDO VALE 558388, across small divide to next gully to SW and extending down it to what is approximately axis of Dosey Syncline. SD128 | SDI30 | (South) - Continuation o! ding section, traversing across other limb of Dosey Syncline, going down section | io contact between Storm Hill Sandstone and Dosey Limestone; section ends at ANDO VALE 558384. Section through Papilio Formation, commencing in top of Dosey Limestone in a Bully at WANDO VALE N 566396, crossing it and terminating over next gully to SE at WANDO VALE 568395, approximately 2 km NNE of Storm Dam. Section through Dosey Limestone and Papilio Formation, commencing at WANDO VALE 561392, ly 1.6 km NNE of Storm Dam. |SD131 uu | ing i Creek to terminate at high escarpment of Mytton Formation. Section measured in Lomandra Creek, through Bracteata Formation, Lomandra Limestone, Storm Hill SD170 |Sandstone, Dosey Limestone and basal beds of Papilio Formation, commencing at WANDO VALE 609398, approximately 6,5 km NE of Storm Dam. spiôn | Spot locality in base of Papilio Formation, at WANDO VALE 600420, approximately 300 metres SSW of junction of Dosey and Lomandra Creeks, and approximately 5.7 km NE of Storm Dam. spiu> |Section through WANT Member of Mytton Formation, west of Pages Creek to top of ridge, | sp204 Section through Dosey Limestone and Papilio Formation, commence 596400, approximately 4.3 km NE of Storm Dam and aligned down terminate at high escarpment of Mytton Formation. ing in Camp Gully at WANDO VALE amp Gully, crossing Dosey Creek to Section through to 7m of Dosey Limestone and through Papilio Formation, commence: WANDO VALE 60140] approximately, 4.8 km NE of Storm Dam and aligned down n GB Gully at Gully, crossing Dosey approximate: gh Dosey Limestone and Papilio Formation, commencing at WANDO VALE 560391, commencing at WANDO VALE 543415, approximately 4 km NNW of Storm Dam, Section throw approximately 1.5 km NNE of Storm Dam. Section throug Lomandru Limestone, Storm Hill Sandstone and Papilio Formation, commencing at WANDO VALE 574405, approximately 3.2 km NE of Storm Dam. Section through Papilio Formation, commencing at head of gully at WANDO VALE 533360, approximately 2.2 km SW of Siom Dam. Section through Papilio Formation, commencing at head of gully, tributary to Storm Dam Creek at WANDO VALE 556376, approximately 1 km ESE of Storm Dam. ND216 Section through Spanner Member of Papilio Formation, commencing at WANDO VALE 546422, approximately 6 km NNE of Storm Dam. edi ventric ridges around the posterior margin of the short, backwardly-pointing, slightly overlapping crown. Scale morphology ts known in atleast 18 ribs (Gross, 1973: pl,33, figs 13-15), whereas the genera of Early and Middle Devonian chondrich- crown of Gondwanalrpis has short, parallel, thyans (Tables 2, 3), but the scales described rounded ribs at the anterior. Scales of below cannot be readily referred to any of these. Protacrodus, also discussed by Gross (1973), Wells (1944) assigned scales to three new differ from Gondwanalepis in having a low, flat genera - Cladolepis, Ohiolepis, and Deirolepis - crown, highly convex base, and distinct furrow and also illustrated scales of genera Crenacan- where the base joins the neck area (Gross, 1973: thas, Cladoselache, and Cladodus (Wells, 1944: pl. 32, figs 3-20; pl. 33, figs 1-12). figs 6,7; pl. 3, figs 2-21). Seales of Scales of Polymerolepis were initially con- Gondweinalepisare like none ofthese. Cladvlepis sidered heterustracan by Karatajite-Talimaa scales have a flat, thin crown ornamented by long (Obruchev & Karatajiite-Talimaa. 1967), but arc curved ridges with shorter, overlapping ndges likely tò belong to an Early Devonian shark anteriorly; the crown of Ohiolepis scales is (Turner & Murphy, 1988). Scales illustrated by covered with numerous anteriorly grooved spines Obruchev & Karatajiite-Talimaa (1967) and (see further discussion below); Deirolepis scales © Turner & Murphy (1988) have the crown heavily have a long neck and thin base; and scales of ornamented from anterior to posterior with many Crenacanthus, Cladeselache, and Cladodus all deep ridges, in most specimens parallel, but lack the anterior parallel ridges present on the sometinies radial; this is quite different from the crown of Gondwanalepis. crown omamentation of Gondwanalepis. Gondwanalepis scales arc unlike those of Her- From the Emsian Receptaculites Limestone, cynolepis in their crown ornamentation. The Giffin (1980) figured two scales referred to crown uf Hercynolepis scales is covered with Karatajiite-Talimaa’s thelodont Skamolepis. MID-DEVONIAN CHONDRICHTHYAN SCALES 91 These are now considered Io be scales of a new chondrichthyan (Turner, 1993). The scales bear no resemblance to Gondwanalepis scales in either shape or ornamentation. Ellesmereia, erected by Vieth (1980), has ndges extending from the anterior right to the posterior margin of the crown, in contrast to the short anterior ridges of Gondwanalepis, and the neck of Ellesmereia is more constricted. Gondwanalepis differs from scales of An- tarctilamna Young, 1982 in having ridges only at the anterior of the crown, in being generally thicker in lateral view, and in lacking a con- stricted neck and concave, cup-shaped base, Pruemolepis, when erected by Vieth-Schreiner (1983), was placed in the acanthodian order Climatiida. Mader (1986) transferred the genus to the chondrichthyan subclass Elasmobranchii on the basis of histology. However, Pruemolepis scales are now thought to be acanthodian branchial scales (Valiukevicius, pers. comin,, 1993). Pruemolepis scales have a thicker crown, a more constricted neck, and less conspicuous crown ornamentation than do the scales of Gondwanalepis, Gondwanalepisis unlike the genera from China described by Wang (1984): Gualepis, Chan- golepis, and Peilepis. Gualepis, unlike Gondwanalepis, is characterised by a constricted neck and a dentate posterior margin. Gualepis- like scales have been recovered from the late Early Devonian Cravens Peak Beds of the Geor- gina Basin (Turner, 1993: fig. 8.4g,h). Chan- golepis, although having a similar lateral view to Gondwanalepis, differs by having a strongly con- vex central rib on the crown forming a long posterior cusp. The crown of Peilepis has anterior flutings and is posteriorly bifurcated, and the flat base has a large elliptical pulp opening. Scales of the genera erected by Mader (1986), Arauzia, Iberolepis and Lunalepis, differ from those of Gondwanalepis in their crown omamen- tation. The crown of Arauzia scales bears one or several thick, stellate, posteriorly-inclined, pointed tubercles. Both Jerolepis and Lunalepis scales have parallel ridges extending to the posterior margin of the crown. However, the lateral view of one Lunalepis scale illustrated by Mader (1986: pl. 4, fig. 9b) is similar to the lateral view of some scales of Gondwanalepis. Gondwanalepis grossi sp.nov. (Figs 4; 5; 6A-C) ETYMOLOGY For Walter Gross (1903-1974). MATERIAL HOLOTYPE: Scale QMF26084 (Fig. 4A-C), OTHER MATERIAL: Figured scales QMF26085 - 26093 and 124 other scales. LOCALITY AND HORIZON Scales occur at eleven localities in the area (SD15, SD128, SD130, SD131, SD146, SD164, SD170, SD196, SD204, SD210, SD216 - Fig, 2), in horizons of the Papilio Formation and its as- sociated Spanner Limestone Member, and the Dosey Limestone, ranging in age from kock- elianus to hermanniscristatus conodont zones. DIAGNOSIS As for genus (this is the only species). REMARKS The Jateral view of Gondwanalepis scales is similar to that of both Changolepis tricuspidus Wang, 1984 from the Early Devonian of south-west DEVONIAN | MIDDLE | rae oa G7 HE Wi EISE oO suekyiyoupuoyz Gandwanalepis grossi Natiolepis dienemos var. 1 var.2 vat.3 Aussilepis lukasa Ohiolepis sp. Cladolepis sp. ct. C. gunnelll Chondrichthyan A Chondrichthyan B FIG, 3. Relative ranges of chondrichthyan taxa, 92 MEMOIRS OF THE QUEENSLAND MUSEUM China (Wang, 1984; fig. 12f) and Maplemillia costata Gross, 1973 from the Late Devonian of Iowa (Gross, 1973: pl. 30 fig. 1d). In all three cases, the gently convex base is flared into a rim around the edge, the neck is deeper al the back, and the crown slopes up and back from the anterior of the base, with no anterior edge on the crown. But Gondwanalepis differs from Chan- golepis and Maplemillia m the crown omamen- tation, neither of these genera has short, parallel ridges on the crown. The short, deep, parallel ridges on the crown of Gondwanalepis are similar to the ornamentation on some acanlhodian scales, e.g. Cheiracam- thoides comptus Wells, 1944 illustrated by Giffin (1980: fig. 5), or Cheiracanthoides sp. cf, com- ptas illustrated by Boucot et al, (1989: fig. 19). However, in Cheiracanthoides scales the crown is more pointed postenorly, and the base more convex. The original generic diagnosis for Cheiracanthoides of Wells (1944) includes several characters Which could be confused with these of Gondwanalepis. Cheiracanthoides scales have radiating ridges usually extending at least halfway to the posterror corner of the crown, which has a well-defined antenor edge (Wells, 1944: fig. 3). However, the grooves between the ridges are not deep enough to notch the anterior margin deeply, the scales have a well-developed neck, and there are no neck canals. In contrast, scales of Gondwanalepis have parallel ndges which are shorter than in Cheiracantholdes, the crown has no antenor edge, and thc ridges con- tinuc down over thc front of the scale, where they nòlċh the margin deeply. The neck is not well- developed, being indented only at the posterior. In addition, Gondwanalepis scales always lack the concentric ridges on the base, which are typi- cal of acanthodian scales (Gross, 1973). MEASUREMENTS The scales vary in length between 0.3mm and 0.8 mm, and in height between 0.3mm and 0.7 inm. The width of most scales is between 0.6mm and 0.8mm, but the range extends from 045mm to 1.1m1n. The Jength/width ratio ranges froin 0.5 in the particularly wide, high scales with the subrectangular, concave base, to 1.0 in the rela- tively longer scales with the diamond-shaped, convex base, Approximately 75% of the scales in the available sample have width greater than length. The remaining scales have equal width and length. DESCRIPTION Morphology. The crown bears eight short sub- parallel ridges. The ridges deeply dissect the anterior edge of the crown, extending down al- most 1o the flared rim around the base. In some particularly wide specimens, with a gently curved anterior margin, the ridges are extremely deep, and give the edge of the scale a scalloped ap- pearance (Fig. 4M-O). The anterior edge of the scale is variably curved, ranging from ap- proximately 90° to 150°. Only 4% of the scales in the available sample have the anterior edge gently curved (Fig. 4M-O); 60% of the scales have the anterior edge moderately curved (Fig, 4A-C, G-I); the remaining scales (36%) have a more sharply curved anterior margin (Fig. 5A-C). The central ridges extend back from the anterior margin for about one-third the length of the crown, The outer ridges continue as low, narrow curved ridges parallel to the lateral and posterior margins of the crown (Fig. 4A). There may be up to four of these concentric ridges in the posterior section. The anterior margin of the crown is high; the crown slopes slightly upwards posteriorly and extends only a short way beyond the posterior of the base (Fig. 4B,E). The neck is not clearly defined, although all scales are high, particularly those that are wider than they are long. The neck area is indented posteriorly, and shows up to 12 elliptical canal openings (Fig. 5M,6C), A single row of small circular openings around the lower part of the FIG, 4. Gondwanalepis grossi gen, el sp.nov. A-C, holotype, scale OMF26084 from SD164/19; D-F, scale QMF26085 from $D216/106,1; G-1, scale OMF26086 from SD164/18; J-L, scale QMF26087 from $D128/212 (50 paces north); M-O, scale OMF26088 from SD128/210, A, crown view, x60; B, lateral view, x 95; C, basal view. x 60; D, crown view, x 50; E, lateral view, x 75; F, basal view, x 45; G, crown view, x 45; H, lateral view, x 70; I, basal view, x 45; J, crown view, x 60; K, lateral view, x 90; L, basal view, x 60; M, crown view, x 75; N, lateral view, x 90; O, latero-basal view, x 90. FIG. 5. Gondwanalepis grossi gen.cu sp.nov. A-C, scale QMF26089 from SD204/174; D-F, scale OMF26090 from SD164/18; G-1, scale OMF26091 from SD210/9; J-M, scale QMF26092 from SD204/168, A, crown view, x 55; B, lateral view, x 65; C, basal view, x 50; D, crown view, x 50; E, lateral view, x 90; F, basal view, x 60; G, crown view, x 60; H, Jateral view, x 75; 1, basal view, x 50: J, crown view, x 40; K, lateral view, x 60; L, basal view, x 40; M, posterior view, x 50. MID-DEVONIAN CHONDRICHTHYAN SCALES 93 94 MEMOIRS OF THE QUEENSLAND MUSEUM MID-DEVONIAN CHONDRICHTHYAN SCALES B5 anterior neck area is Visible in some spectmens (Fig. 6A). The base js most commonly diamond-shaped (Fig. 4C, F, I,L), but may be subrectangular (Fig. 4N-O). These represent the ends of a continuous range of morphological types. The diamond- shaped bases are usually gently convex, or some- times flat, and flared into a narrow rim around the edges. The subrectangular bases are gently con- cave. All the scales are high relative to the crown area, but this is particularly noticeable in those specimens with the concave, subrectangular base (Fig, 4N-O), Histology, The base of cellular bone extends in an inverted cone shape high into the scale (Figs 11A.B). This tissue contains osteocytes aligned both concentrically and radially (Fig. 11B). The crown appears Io consist of discrete increments added anteriorly and posteriorly to a central ini- tial element (arrow in Fig. 11C). This type of apposed growth is typical of complex chondrich- thyan scales (Zangerl, 1981), in contrast to the concentric growth pallern in most acanthodian scales (Dension, 1979); one exception is the acan- thodian Nostolepis robusta (Brotzen), 1934 described by Gross (1971a) - these scales show some apposition of crown elements. The postenor section of the crown consists of cellular material (Fig. 11D) similar in appearance to the Stranggewebe in acanthodian climatiid scales such as Cheiracanthoides comptus Wells, 1944 (e.g. Gross, 1973: fig. 5b,c). This nostolepid-type histology typical of climatiids is also seen in some early cladolepid chondrichthyans (J. Vergoossen, pers. comm., 1994), DISCUSSION It is possible that differently shaped scales are of different ages, or occur on different parts of the body. The wide, short scales (Fig, 4M) have a similar width range to the longer scales (Figs 4A,D,5G), Complex shark scales grow by adding new growth clements around the margins (Zangerl, 1981); it is conceivable that the scale depicted in Fig. 4M is a young scale that could have grown further by additions at the anterior and posterior (see discussion of histology above), resulting in a scale with a more curved anterior margin, longer anterior ridges on the crown, and a longer posterior section to the crown (Figs 4A-C, 4G-I, 5G-T), Notiolepis gen, nov. ETYMOLOGY From the Greek 'notios' = southern, and 'lepis” = scale. DIAGNOSIS Crown oval or subrectangular, bearing four to eight parallel anterior ridges, Crown joins direct- ly onto base anteriorly, and slopes up towards posterior. Neck not indented, and has six to twelve canal openings along posterior. Base rhombic or suboval, flat or concave, with no rim around the edge. REMARKS The scales of Notiolepis are distinguished from those of Gondwanalepis, described above, by the following characters: the crown is relatively longer compared with the width; the antenor ndges on the crown are less pronounced; the neck has the same thickness as the base, rather than being indented as in Gondwanalepis ; and the base lacks a rim around its edge. These seem to be consistent differences, even though there is variation in crown shape within each genus. which may indicate scales from different parts of the body. On the available small sample, Notiolepis is therefore considered to be a separate: taxon, although this assessment may change as more material becomes available. Nottolepis gen. nov, as defined here may be distinguished from previously described genera inthe type of crown omamentaiion; the six genera FIG, 6. A-C, Gondwandlepis grossi gen.ct sp, nov., scale QMF26093 from SD204/174; D-L. Notiolepis dienemos gen.etsp nov. Var. 1. D-F, holotype, scale QMF26094 from SD15/81; G-I, scale OMF26095 from SD204/119.5; EL, scale OMF26096 from SD204/174. A, lateral view, x 75, B, crown view, x 50; C, posterior view, x 65; Dy crown view, x 45; E, lateral view, x 65; F, basal view, x 50; G, crown view, x 50; H, lateral view, x 65; I, basal view, x 50; J, crown view, x 90; K, lateral view, x 90; L, basal view, x 90. FIG. 7. A-J, Notiolepisdienemos gen.ct sp.nov. Var. 2; K-M, Cladolepissp.cf. Cgunnelli, A-D, scale QMF26097 from $D128/212+ (150 paces N) - specimen broken during SEM photugraphy; E-G, scale OMF26098 from SD128/217; H-J, scale QMF26099 from SB 128/202.3, K-M, scale OMF 26100 from SD 190/60, A, crown view. x 35; B, basal view, x 60; C, posterolateral view, x 75, D, detail of broken edge, x 275; E, crown view, x 70; F, lateral view, x 75; G, basal view, x 70; H, crown view, x 55: 1, lateral view, x 60; J, basal view, x 55; K, crown 96 MEMOIRS OF THE OUEENSLAND MUSEUM MID-DEVONIAN CHONDRICHTHYAN SCALES 97 98 MEMOIRS OF THE QUEENSLAND MUSEUM described and illustrated by Wells (1944); Cladolepis, Ohiolepis, Deirolepis, Ctenacan- thus, Cladoselache, and Cladedus, Hercynolepis and Protacrodus described and illustrated by Gross (1973); Polymerolepis figured by Obruchev & Karatajiiie-Talimaa (1967) and Turner & Murphy (1988); 'Skamolepis' (Giffin, 1980; Turner, 1993); Ellesmereia and Pruemolepis (Vieth, 1980; Vieth-Schreiner, 1983); Anrarctilamna (Young, 1982); the three genera from China, Gualepis, Changolepis, and Peilepis (Wang, 1984); and the Spanish lberolepis and Lunalepis (Mader, 1986). No scales of these genera have the short, parallel ribs on the anterior section of the crown, with the posterior part of the crown smooth, as in Notiolepis, Notiolepis also differs from most of these other chondrichthyan genera in lateral and basal views. Nofiolepis scales have a similar lateral view to scales of Crenacanthus cosiellanis Traquair, 1884, Peilepis solida Wang, 1984, and Tberolepis aragonensis Mader, 1986; the dif- ferences between Notiolepis and these genera are discussed below. Notiolepis dienemos sp.nov. (Figs 6D-L; 7A-J; 8A-F) ETYMOLOGY From the Greek 'dienemos' = windswept, referring to the appearance of the crown, MATERIAL HoLoryre: Scale QMF 26094 (Fig. 6D-F), OTHER MATERIAL: Variety 1: Figured scales, QMF26095 & 26096, and 24 other scales. Variety 2: Figured scales, OMF26097-9, and seven other scales. Variety 3: Figured scales, OMF2510|-2, and five other scales. LOCALITY AND HORIZON All specimens were recovered from the Papilio Formation, Spanner Limestone or Stanley Lime- stone. Variety 1 occurred at five localities (SD15, SD128, SD204, SD210, SD216 - Fig- 2) in honzons ranging in age from mid- emnsensis to hermanni-cristatus conodont zones; Variety 2 at two localities (SD128, SD210) in varcus Con- odont Zone; and Vanety 3 at five localities (SD15, SD128, SD192, SD204, SD210) in kock- elïanus, lower and middie varcus, and asym- metricus conodont zones, DIAGNOSIS As for genus (only species). MEASUREMENTS Variety 1: scale length varies between 0.5mm and 0.9mm; width ranges from 0.4mm to 0.7mm; height ranges from D,2mm to 0.3mm. The length/width ratio is 1.17 to 1,29, Variety 2: length of the crown ranges from 0.7mm to 0.9mm, and width from 0.5mm to 0.6mm. The length/width ratio ranges from 1.4 to 1.6. Variety 3: the crown length is 0.6mm to 0.7mm, the width is 0.7mm to 0.8mm, and the length/width ratio is 0.86 to 0.87. DESCRIPTION Morphology. Three varieties of scales have been grouped together on the basis of their similar lateral views (Figs 6E,H,K; 7F,1; 8B,E). In Variety 1, (Fig. 6D-L), the crown is oval shaped, with both anterior and posterior edges gently curved. The crown bears four to seven long parallel ridges, usually starting at the anterior edge and extending posteriorly approximately halfto two-thirds the length of the crown. In some specimens, the base extends slightly anteriorly, to give a smooth anterior margin to the scale in dorsal view (Fig. 61). In these specimens, the parallel ridges do not extend right to the antenor margin of the scale, and the crown extends further beyond the base posteriorly than in the specimens lacking the anterior basal extension, The base is diamond- shaped, and either flat or gently con- cave, Up to 12 neck canal openings are found at the posterior. In Variety 2 (Fig. 7A-J) the crown is subrectan- gular, thin, and slopes steeply upward from a fairly straight anterior edge to a gently rounded posterior margin, The crown bears four to seven parallel ridges, extending from just behind the anterior edge, back to approximately mid-way towards the posterior (Fig. 7E,H). The neck area is not indented. The base is a concave, narrow semi-diamond or semi-oval shape, and, like the crown, is straight along the anterior margin (Fig. 7B,G). The general appearance of the scale is wide, flat and high, with only the base area ap- pearing to have much volume. Specimen QMF26097 was broken during SEM photog- raphy, revealing numerous pulp canals within the crown (Fig. 7D), typical of the composite type of chondrichthyan scale described by Zangerl (1981), where each new growth element has its own vascular supply. Variety 2 scales mainly differ from those of Var. 1 in their base. The base of Var, | scales is flat or gently concave, and diamond-shaped. The MID-DEVONIAN CHONDRICHTHY AN SCALES 99 TABLE 2. Reported chondrichthyans from the Early and Middle Devonian of Australia. LT aan y | Late Giv. -early Fras. Lochkovian 'Skamolepis' sp. denticles, endacraniade Turner, 91; 93 BE" — Turner, 91:93 pr cf. Rhyolite, Mt. Dam Limestone Réceptheutites Rece, picnics t Late Emsian teeth, fin spines, WORN sp. Bunga Beds | Ohiolepis sp- Jerula Fm., Trundle Turner, 93 a Em——— a Pete LAS etal., 85 H serotinuslpatulus Zones - early Eifelian THL fs erbonus - scales, dermal Young, 82; Long, 91; Beds, Dulladerry Tumbong si | |Tomer, lTumer,93 | ene wdn seman | dehiscens - serotinus laticostatus) Giffin, 80 Jesse Limestone scales Turner, 93 Mt. Frome Limestone ine Ms | Possible occurrences: pe Bunga Beds base of Var. 2 scales is les ie deeply concave, and a narrow half-diamond or semi-oval shape, with the anterior edge of the base fairly straight. In Variety 3 (Fig. 8A-F) the crown is fairly straight across the anterior, but rounded at the posterior (Fig. 8D). The front of the crown slopes up steeply from the base, then levels out and slopes up more gradually towards the posterior margin (Fig. 8B). Seven short parallel ridges occur on the steeply sloping anterior edge of the crown. The posterior part of the crown is flat, thin and unornamented. The neck is not constricted. The base is a flat, narrow rhombic shape (Fig. serotinus - patulus Zon a_^) Trundle Beds ?Late Lochkovian — Turner, 91; 93 Mid Devonian Turner, 93 articulated or semi- articulated cartilage Long, 91 remains ve, anda BC.F), Six to eight ~F). Six to eight elliptical neck c neck canal openings occur on the underside of the crown, just behind the base. The overall appearance of these scales is wide, flat, and low. The crown is wider than it is long. Variety 3 scales differ from Var. 1 and Var, 2 in the shape of both the base and the crown. The base here is flat and narrowly rhombic, in contrast with the diamond-shaped flat or gently concave base of Var. 1, and the narrow half-diamond- or semi-oval-shaped deeply concave base of Var. 2. The crown of Var. 3 scales is straight at the anterior edge, widely rounded at the posterior edge, and bears seven short parallel ridges only at the very front of the scale. The crown of Var. 1 and Var, 2 bears parallel ridges extending much further back towards the posterior. The angle of inclination of the crown varies between in- dividual scales but is similar in all three varieties of Notiolepis. However, each of the three varieties has the length/width ratio falling within a distinctive range, reflecting a general difference it overall scale shape between the three varieties: 1.17-1,29 for Var. 1, 1.4-1,6 for Var. 2, and 0.86-0.87 for Var. 3. Histology. Scales consist of a conical bony base (Fig. 12A), containing osteocytes and fine radial fibres (Fig. 12B). The cellular crown material is attached to the posterior edge of the basal cone (Fig.12A). The crown, like that of Gondwanalepis, appears 1o grow by appesition of discrete elements (Fig. 12B), and these crown elements also consist of material similar in ap- pearance to Strangeewebe (Fig. 12C). DISCUSSION Notiolepis scales are similar in lateral view to the scales of the chondrichthyans Cienacanthus costellatus Traquair, 1884 (Reif, 1978: fig. la), Peilepis solida Wang, 1984 (Wang, 1984: figs 14, 15), and Jberolepis aragonensis Mader, 1986 (Mader, 1986: pl. 4, fig. 2c). Noriolepis scales differ from C. cestellatus Traquair, 1884 scales by lacking the constricted neck and flared base, and having less pronounced, but a greater number of, anterior ridges on the crown. The P. solida Wang, 1984 scales have an anteriorly protruding base, a large basal cavity, a posteriorly bifurcated crown, and three flutings on the anterior crown surface, all of which are absent in Neriolepis. The base of J. aragonensis Mader, 1986 is longer and flatter than that of Notiolepis, and the crown ornamentation is different. Specimens of Netiolepis with the concave base (Var. 1) resemble those of Antarctilamna prisca Young, 1982 in basal view (Young, 1982: fig. 4c). However, Notiolepis Var. 1 scales differ in having a more gently concave base, lacking a constricted neck, and having parallel ridges in the anterior part of the crown rather than concentric MEMOIRS OF THE QUEENSLAND MUSEUM ridges over the whole crown forming short cusps along the posterior margin. Variety 2 scales also have a basal view similar to that of A. prisca Young, 1982, except that the concave cup-like base is narrower. In lateral view, Noriolepis scales may perhaps be confused with some thelodont scales, for ex- ample Turinia sp. (Young et al., 1987: fig. 5h) or Canenia grossi Vieth, 1980 (Vieth, L980; pl, 3, fig. 3c). However, the Notiolepis scales lack the thick, rounded base encircling the lower cdge of the scale, the large basal cavity and the single pulp canal opening typical of thelodont scales, In addition, the posterior part of thc crown is thinner and flatter, and the crown bears ndges only at the antenor edge, not laterally as is the case with the Turinia sp. scales figured by Young et al. (1987), Scales of C. grossi Vieth, 1980 have a round, gently convex base in ventral view, and three pronounced ndges covering the length of the crown and extending into three denticles at the posterior margin in dorsal view. Notiolepis scales, in contrast, have a rhombic, flat or con- cave base, ridges only in the anterior part of the crown, anda smoothly rounded posterior margin, Aussilepis gen. nov. ETYMOLOGY From the local colloquial word Aussie, and the Greek 'Iepis' = scale. DIAGNOSIS Crown subcircular, extends beyond base posteriorly. Eight thick, deep ridges extend from anterior edge to approximately middle of crown, Neck shallow, slightly indented at anterior and posterior. Approximately six to ten small open- ings in posterior part of neck. Base diamond- shaped, convex. REMARKS The scales of Aussilepis resemble those of Gondwanalepis in the deep anterior ridges on the crown, and in the narrow rim around the margin of the base, this latter feature distinguishing these two genera from Notiolepis. However, Aussilepis FIG. 8, A-F, Notielepis dienemos genet sp.nov, Var. 3; G-M, Ohiolepix sp. A-C, scale QMF26101 from SD128/212; D-F, scale OMF26102 from SD204/174; G, scale OMF26105 from SD128/144.2; H, scale QMF31828 from SD204/174 upslope; I, scale OMF26104 from SD216/99.8; J, broken scale QMF31829 from SD204/174 upslope: K.L., scale QMF26103 from SD130V262.5; M, scale OMF31830 from SD204/174 upslope. A, crown view, x 60; B, lateral view, x 85; C, basal view, x 60; D, crown view, x 50; E, lateral view, x 95; F, basal view, x 50; G, crown view, x 25; H, basal view, x 45; L crown view, x 35; J, crown view, x 60; K, crown view, x 45; L, lateral view, x 60; M, crown view, x 45. MID-DEVONIAN CHONDRICHTHYAN SCALES 101 MEMOIRS OF THE QUEENSLAND MUSEUM 102 MID-DEVONIAN CHONDRICHTHY AN SCALES differs from Gondwanailepis in the following fea- tures: the posterior part of the crown is smooth, thin, and lacks the narrow ndges parallel to the terior margin; the crown overhangs the base urther posteriorly; the base is more convex: and the outer ridges of the crown are less thickened (Fig. 9B,E.H) in contrast to Gondwanalepis (Figs 4B,K; 5E,K). Aussilepis scales are distinguished from Noriolepis scales by the differences in neck and base; Aussilepis has a slightly indented neck and convex base, while in Noriolepis the neck and base are the same size, with the base being either flat or concave. The base of Notiolepis scales occupies a smaller proportion of the length of thc scale than with Aussilepis scales. These scales have been assigned toa new genus because their morphology is unlike that of pre- viously described scales (Tables 2, 3). As with Gondwanalepis and Notiolepis, it is the erown omamentation that differs most from previously described genera, The crown of Aussilepis has short, deep ridges and furrows at the anterior margin, while the posterior section of the crown in unornamented (except for occasional low markings, similar to growth rings, parallel to the posterior margin Fig. 9J). Cladolepis scales (Wells, 1944) have a flat, thin crawn ornamented by long curved ridges with shorter, overlapping ridges anteriorly. The crown of Ohiolepis scales is covered with numerous anteriorly grooved spines (Wells, 1944), Deirolepis scales (Wells, 1944) have a long neck and thin base. Scales of Ctenacanthus, Gladoselache, and Cladodus, as figured by Wells (1944), all lack the anterior parallel ridges present on the crown of Aussilepis, The crown of Hercynolepis scales is covered with short, backwardly-pointing, slightly overlapping ribs (Gross, 1973). Protacrodus scales have a low, flat crown, deeply convex base, and distinct furrow where the base joins the neck area (Gross, 1973), Scales of Polymerolepis have the crown omamented with many deep parallel or radiating ridges (Obruchev & Karatajiite-Talimaa, 1967; Turner & Murphy, 1988). Scales described as 'Skamolepis' by Giffin (1980) differ from those of Aussilepis in general shape and omamentation. Scales of Ellesmereia (Vieth, 1980) have ridges extending from the anterior right to the posterior margin of the crown, and the neck of Ellesmereia FIG. 9. Aussilepis (ukaso gen.et sp.nov, A-C, holo 18 more constricted, Aussilepis lacks the con- stricted neck, the concave, cup-shaped base, and distinctive curved ridges on the crown of An- tarctilamna (Young, 1982), and is thicker tn lateral view. Pruemolepis scales (Vieth- Schreiner, 1983; Mader, 1986) have a thicker crown, a more constricted neck, and less con- spicuous crown ornamentation than those of Axs- silepis, Gualepis (Wang, 1984) is characterised by a constricted neck and a dentate posterior margin. Changolepis (Wang, 1984) has a strong- ty convex central rib on the crown forming a long posterior cusp. The crown of Peilepis (Wang, 1984) has anterior fÌutings and is postenorly bifurcated, and the flat bese has a large elliptical pulp opening. Scales of both Jberolepis and Lunalepis (Mader, 1986) have parallel ndges ex- tending to the postenor margin of the crown. Aussilepis lukaso sp.nov. (Fig. 9) ETYMOLOGY From the languages of the Koori (original in- habitants of Australia) ‘luk’ = like, and ‘kaso’ = another, referring to the similarity between the lateral view of these scales and that of some other taxa. MATERIAL HOLOTYPE: Scale QMF26106 (Fig. 9A-C). OTHER MATERIAL: Figured scales, OMF26107-9 and 75 other scales. LOCALITY AND HORIZON Scales occur at eight Iecalities (SD15, SD128, $D146, SD164, SD196, SD204, SD210, SD216 -Fig. 2), tn horizons of the Papilio Formation and its associated Spanner Limestone Member dated from ensensis to hermanni- cristatus conodont zones. DIAGNOSIS As for genus (this ts the only species). MEASUREMENTS Scales range in length from 0.5mm to 0.9mm; in width from 0.4mm to 0.9mm; and in height from 0.3mm to 0.45mm. The length/width ratio ranges from 1.0 to 1,33. , scale OMF26106 from SD204/174; D-F, scale QMF26107 from §D204/174; G-I, scale OMF26108 from SD204/174; J-L, scale QMF26109 from SD210/30; A, crown view, x 50; B, lateral view, x 70; C, basal view, x 45; D, crown view, x 50; E, lateral view, x 60; F, basal view, x 50; G, crown view, x 45; H, Jateral view, x 60; I, basal view, x 45; J, crown view, x 60; K, lateral view, x 75; L, basal view, x 60. 104 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 3. Reported eee from the Early and Middle Deve Devonian from overseas. me Nu WW——— feat Setene fin spines, a prisca Aztec Siltstone, Antarctica Givetian scales, Young, 82; 89; 91 FFF | Cin hin Ln Mi Dn Mass [Gre Artarenana sp. Sais yne [ma Sutera | Bolivacanthus sagitalis | Catavi Fiy., Bolivia Late Early - Mid. Devonian in ament | Cagnier et al., 88 Santa Rosa Fm., Bolivia phan Pragian Lelievre et al., 93 Chang golepis tricuspidus Xitun Member, Sth China Earl [Early Devonian | Wang Nianzhong, 84 | eect indian, 0S esyd Devon [sates oes. 75 1 Lauch AR German scale Vieth - Schreiner, 83 = Cincinnati Arch region, US Wells, 44 ‘ladies, — ——OeO Mader, 86 Wells, 44 Cincinnati Arch region, US _ |Mid. Devonian __|scale_|Wells.44_| Ctenacanthus sp. fin spine__ |Janvier & Melo, 92 Late Givetian - Earl Erma y = Lelievre et al., 93 Wang Nianzhong, 84 Her olepis meischneri ee” C [pty Donan [tn | Mader, 86 |iberolepisaragonensis _ |Spain [Early Devonian scales |Mader86 | edu andwyo dach lw tens ly brent E en ee ES" ee aes he — |ia— [Mt — Sotenich Trough, German Bifelian [scales | SA i Lauch Fm., Germany |EarlyEifelian [scales _| Vieth-Schreiner, 83 eo Mader, 86 .fobmathensis__|Soteich Trough Germany [Bary Eitetin [seses | Friman, 83 Wang Nianzhong, 84 Ohiolepis sp. ormon RS cuba & Emsian - Eifelian a |. Ørvig, 69 Ohiolepis newberryi MID-DEVONIAN CHONDRICHTHYAN SCALES TABLE 3. Continued. Peilepis solida | Phoebodus floweri | Indiana, US Ph. bryanti Cincinnati Arch regìon, US Iowa, US Polymerolepis whitei Dneister Range, Podolia | Protacrodus sp. Germany & US |Pruemolepissp. | is Sp. Possible occurrences: Indet scale Pleuracanth Indet shark scale DESCRIPTION Morphology. In these scales the subcircular crown extends posteriorly beyond the base such that the length of the base is approximately half to two-thirds the length of the crown (Fig. 9C,F.I,L). The parallel ridges on the crown become thinner and shallower posteriorly, and disappear at about the middle of the crown. The crown has no anterior rim; rather the ridges extend down the anterior edge of the crown and neck, ending just above the flared joint of neck area and base (Fig, 9K). The neck is shallow, in some specimens slightly deeper posteriorly (Fig. 9B,E), with only very slighi Xitun Member, South China En Devonian cae A Cincinnati Arch region, US IMid, Devonian fieeh | Wells, 44 Early Mid, Devonian lleih | Mid. Devonian heeh |wels,4 _ | Early Mid. Devonian koth (ar73 Early Devonian y e 67 Simpson Park Range, Nevada |Lochkovian (delta Zone) EL Tane & Murphy, Protacrodus wellsi [Iowa us Barly Devonian scales [Gross73 | Harz & Morocco [Early Devonian [scales [Gross 73 | Mid. - Late Devonian Tne Givetian - Early Frasnian ee wellsi eef. Cenaer_| nA Emsian - Eifelian 4 83 Spats s oeM —— tag Devonian scales Mader, 86 . n Sicasica & Belen Fms., ; Pucapampella rodrigae Bolivia Eifelian [Vesispitsbergen |Emsian - Eifetian Bokkeveld Gp., South Africa | Late Mid. Devonian Chondrichthyan ? Catavi Fm., Bolivia Late Early- Mid. Devonian | spines | Simpson Park Range, Nevada | Lochkovian (delta Zone) Fees AD sp.nov. & Aztec Siltstone, Antarctica Young, 89; 91 Chondrichthyan __|Talengitsequence [Ems | lieïewectal, 93 105 Wang Nianzhong, Gross, 73 Zanperl, 81 Lelievre et al., 93 Veith - Schreiner, scales synarcual | Lelievre et al., 93 fin element | Lelievre et al,, 93 tooth impressions Oelofsen, 81 Gagnier et aL, 88 Turner & Murphy, BR rphy anterior and posteriorindentationsin lateral view. The neck area is not indented laterally. Six to ten small round neck canal openings are located in the posterior neck area (Fig. 9E). The convex base is diamond-shaped, and flared into a slight nm around the perimeter, where it connects with the neck (Fig. 9B,E,H). In some specimens the base extends anteriorly beyond the crown, producing a smooth anterior margin on the scale (Fig. 9A,D). Such scales are deeper in a dorso-ventral direction. Histology. The conical bony base contains both fine radial fibres (Fig. 13A) and osteocytes (Fig. 106 13B). The crown shows no evidence of con- centric growth but, like both Gondwanalepis and Noriolepis, consists of apposed increments of Stranggewebe-like material (Fig, 13B). DISCUSSION The basal view of Aussilepis lukaso ts similar to that of the Late Silurian shark Elegestolepis grossi Karatajitte-Talimaa, 1973 (Karatajiite- Talimaa, 1973: pl. 3, figs 3b, 4b). However, the crown omamentation of the two types of scale is quite different; the ridges on Elegestolepis are longer, thinners, and more numerous than thosc on Aussilepis. Some of the scales, in lateral view, are similar to illustrated acanthodian scales, for example Nostolepis striata Pander, 1856 (Gross, 1947: pl, 25 fig. 7c; Denison, 1979: fiz. 171, N. arctica Vieth, 1980 (Vieth, 1980: pl. 5, figs 4b, 5b, 7b), Cheiracanthoides comptus Wells, 1944 (Gross, 1973: pl. 27, fig. 2c), or Cheiracanthoides sp. cf. compius (Boucot ei al., 1989: fig, 194), Apart from the absence of neck canal openings, these scales differ from Aussilepls in various mor- phological details, Nostolepis scales have a posteriorly pointed crown omamented with con- verging midges, commonly paired, with a median depression (Denison, 1979; Forey et al., 1992), However, the crown of Aussilepis scales is rounded posteriorly, and extends further beyond the base posteriorly than in the acanthodian scales, The base of Aussilepis ts less strongly convex than the base of Nostolepis scales (Gross, 1947: pl. 26; Forey et al, 1992: fig. 12a). In scales of Cheiracanthoides comptus Wells, 1944, the grooves between the ridges are not deep enough to notch the anterior margin (Wells, 1944), In Aussilepis scales, the anterior edge of the crown is always deeply notched (Fig. 9K), as is also the case with Gendwanalepis. Scales of C. comptus Wells, 1944 have a constricted neck, unlike those of Aussilepis, and a nearly flat crown. Anssilepis scales have a crown that slopes down anteriorly to meet the flared rim joining the base and neck; there is no anterior rim to the crown of Aussilepis as there is in Cheiracan- thoides. Scales of C. comptus Wells, 1944 have a ‘thick, inflated’ base (Wells, 1944), described by Gross (1973) as “highly domed”, The base of Aussilepis scales is convex, but more gently so than in figured scales of C. comptus Wells, 1944, and appears smooth, lacking the distinct con- centric striations characteristic of acanthodian scales (Gross, 1973), As with Nostolepis striata Pander, 1856 (Denison, 1979: fig. 17f), the crown MEMOIRS OF THE QUEENSLAND MUSEUM of C. compius Wells, 1944 extends posteriorly beyond the base a shorter distance than in Aiur- silepis. Ohiolepis Wells, 1944 TYPE SPECIES Ohiolepis newberryi Wells, 1944 Ohiolepis sp, (Fig, 8G-L) REMARKS The genus Ohiolepis was established by Wells (1944) for scales from the Middle Devonian bone beds of Ohio, Indiana, and Kentucky. His two species, O. newberryi and O. stewartae, were subsequently united by Gross (1973), wha ascribed differences in shape between the two species to scales coming from different parts of the body. The taxon has also been reported from Australia (Schultze, 1968; Giffin, 1980; Turner, 1982; Pickett elal, 1985; Turner, 1993; Young. 1993), China (Wang, 1984), and Germany (Orvig, 1969; Friman, 1983; Vieth-Schreiner, 1983); see Tables 2, 3. Turner (1993) questioned the chondrichthyan affinities of some of these records, suggesting that some might be placoderm scales. Two new species have been erected since Gross's 1973 amalgamation: 'Ohiolepis' frohnrathensis Friman, 1983 from the early Eifelian of Germany, and O,? xitunensis Wang, 1984 from the Early Devonian of China. All other occurrences have been assigned either to O, new- berryi or simply to the genus. Scales assigned to the genus and Figured in the literature vary in morphology, Wells (1944) es- tablished as generic characters a flat or convex base; 4 broad crown omamented by numerous spines that are cnamelled, anteriorly indented or grooved, with an anterior sloping face, acute posteriorly inclined tips, and resting directly on the base; and a narrow, spine-free strip around the anterior cdges and sides of the scale. However, Wells (1944:38) also remarked that the spines of O), newberryi Wells, 1944 may be blunt and club- shaped; this is inconsistent with his diagnosis. Wells described his less common second species, O. stewariae, as having spines similar to those of O. newberryi Wells, 1944, but fewer in number, and arranged in a single transverse series with the smallest spine in the centre. Gross (1973) placed the two species in O. new- berryi Wells, 1944 by combining Wells's two MID-DEVONIAN CHONDRICHTHYAN SCALES groups of specific charac- ters, but made no mention of the spines being blunt or club-shaped, or having an anterior groove. However, his illustrations show scales with both pointed (Gross, 1973; pl. 30, figs 8-21) and blunt-tipped (pl. 31, fig. 8) spines, and both with (pl. 30, figs 8-21) or without (pl. 31, figs 6,7) an anterior groove. Most of Gross's figures (1973: pl. 30, figs 8-19,21; pl. 31, figs 3-7) show examples similar to Wells's original O. new- berryi type, with numerous closely- packed spines, each having an anterior groove and a posieriorly- directed acute tip. Other examples (Gross, 1973: pl. 31, figs 1, 2) have the spines more widely-spaced and conform more closely to Wells's original descrip- tion for O. stewartae. In- deed, all specimens of Ohiolepis described since 1973 have widely-spaced spines (Giffin, 1980: fig. 4a,b; Friman, 1983: figs 1,2; Vieth-Schreiner, 1983: pl. 4, fig. 36; Wang, 1984: figs 16,17; Pickett et al., 1985: cover photo K). On this evidence, it seems best for the moment to provisionally retain Wells's two species, with O. newberryi only known from North America, and examples from elsewhere showing more resemblance to O. stewartae. In the absence of precise locality data, it is possible that the material analysed by Gross came from several different horizons. The concept of the genus Ohiolepis is further confused because one of Friman’s (1983) species characters for. ‘Ohiolepis’ frohnrathensis is in- consistent with Wells’s (1944) genus diagnosis: 'Ohiolepis' frohnrathensis Friman, 1983 scales lack an anterior groove on the spines. One option is to consider anterior grooves on the spines a generic character applying to most scales in a sample, but accepting some variation in sca]es from different regions of the body. Friman's specimens (1983: fig. 2b-e) have spines on the crown not dissimilar to those found on dermal denticles from the branchial region of An- tarctilamna prisca Young, 1982 (fig. 2c,d; pl. 87, 107 FIG. 10. A-C, Chondrichthyan A - fam.,gen. et sp. ndet., scale QMF26110 from SD15/192; D-G, Chondrichthyan B - fam. gen. et sp. indet., scale OMF26]1 11 from SD204/174. A, crown view, x 32; B, lateral view, x 56; C, basal view, x 32; D, crown view, x 48; E, lateral view, x 72; F, basal view, x 48; G, posterior view, x 68. figs 9,10), so this morphotype might occur in many different shark taxa. Wang's (1984) O. ? xitunensis was provisional- ly referred to Ohiolepis. Listed differences be- tween O. newberryi Wells, 1944 and his new species included shape of the denticles, and shape and structure of the base. MATERIAL Six scales; OMF26103-5, QMF31828-30 (Fig. &G M). LOCALITY AND HORIZON The scales occur in four sections of the Broken River Group (SD128, SD130, SD204 and SD216 - Fig. 2), in horizons of the Papilio Formation and the associated Spanner Limestone Member aged middle and upper varcus Conodont Zone. This age is younger than other occurrences of Ohiolepis; both @rvig (1969) and Friman (1983) consider the genus to be characteristic of Em- sian/Eifelian boundary beds in Europe, North America, and Australia. MEASUREMENTS The scales range in length from 0.7mm to 2.5mm, and in width from 0.7mm to 1.6mm. 108 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 11.Gondwanalepis grossi gen.et sp.nov., longitudinal section QMF261 12 of scale from SD204/174 upslope. A, full view, x 30; B, detail of base, x 60; C, detail of posterior part of scale, x 60; D, detail of posterior part of crown, x 180. DESCRIPTION AND DISCUSSION With only six scales, the examples from Broken River can only be provisionally compared with Ohiolepis. Each has a crown completely covered with numerous small, conical, closely-packed, posteriorly inclined tubercles or spines, without an anterior groove (Fig. 8G,1I,J,K,M). The spines radiate from the centre anterior margin (Fig. 8G,M) and cover the crown, although one specimen (Fig. 8M) has a small spine-free area in the central anterior region of the crown. Broken spines (arrows in Fig. 8M) show a central cavity. The scales have a subcircular to subrectangular outline, a very shallow neck, and a flat or slightly concave unornamented base (Fig. 8H). The crown of these new scales has spines finer and more closely-packed than those in the il- lustrations of Ohiolepis by Gross (1973), Giffin (1980), Friman (1983), Vieth-Schreiner (1983), Pickett et al. (1985), and ?Ohiolepis-type scales of Liszkowski & Racki (1993). However, one illustration by Wells (1944: fig. 7f) shows small, closely-packed spines, similar to the Broken River specimens. The spines of the Broken River scales overlap (Fig. 8J,M), in contrast to the con- dition noted by Gross (1973). The profile of the scales is low and flat, similar to Giffin’s (1980) description of the Ohiolepis scales from Taemas. Gross (1973) commented that it is hard to differentiate between the crown MID-DEVONIAN CHONDRICHTHY AN SCALES and the neck; this is the case with the Broken River scales. The spines have a round or oval cross-section, without an anterior groove, thus resembling Friman's (1983) species ‘Ohiolepis’ frehnrathensis, as discussed above. Friman (1983) described the omamentation of ‘Ohiolepis' frohnrathensis as asymmetrical, but according to Gross (1973), the spines on the crown are arranged both concentrically and in diagonal rows, with the earliest formed, smallest spines located in the centre, and later ones added around the edges. In most illustrated specimens in the literature, the spines appear to be ap- proximately symmetrical, subparallel and point- ing posteriorly; this is also the case in the Broken River specimens. The Broken River scales have a subcircular to subrectangular outline, and a flat or slightly con- cave base. According to Wells (1944), the base of O. newberryi may be flat or convex. The base of 'Ohiolepis' frohnrathensis Friman, 1983 is al- ways anteriorly convex (Friman, 1983), while that of O. ? xitunensis Wang, 1984 is flat (Wang, 1984). Cladolepis Wells 1944 TYPE SPECIES Cladolepis gunnelli Wells, 1944 Cladolepis sp. cf. C. gunnelli (Fig. 7K-M) Cladolepis gunnelli Wells 1944: 36,37; pl, 3, Figs. 2-7; pl. 8, fig. 4; fig, 6a,b,d. Cladolepis omata Wells 1944; fig. 6h. Cladolepis sp. Wells 1944; fig. 6c,i. Cladolepis gunnelli Gross 1973: 97-99; pl. 31, figs. 11-17; pl. 32, figs, 1,2. ?Cladolepis gunnelli Gross 1973; pl. 31, figs 9.10. Cladolepis? gannelli Vieth-Schreiner 1983; 151,152; pl. 3, fig. 25. Cladolepis cf. gunnelli Mader 1986: pl. 7, fig, 10. REMARKS Cladolepis was erected by Wells (1944) from material from the bone beds of the Cincinnati Arch Region, from horizons subsequenily dated by conodonts as kockelianus Zone or older (Klap- per & Johnson, 1980). The Broken River scales resemble Cladolepis gunnelli Wells, 1944 in shape and crown ornamentation, bul differ in having a thick, sloping crown rather than the flat thin crown of C. gunnelli Wells, 1944, The base 109 is ‘convex and acanthodianlike’, as Wells (1944) described some of his specimens. The crown omamentation is also similar to that of An- tarctilamna prisca Young, 1982, but that form has a completely different base. MATERIAL Seven scales - one intact (QMF26100), three broken at posterior, three severely broken or abraded, LOCALITY AND HORIZON The scales occurred in three sections (SD190, SD192, SD210) fram horizons of the Papilio Formation and the Stanley Limestone Member of the Mytton Formation, dated middle and upper varcus, hermanni-cristatus and asymmetricus conodont zones. MEASUREMENTS The only unbroken scale (QMF26100) measures 1.9mm long, 2.2mm wide and 0.7mm high. All other specimens have the posterior scc- tion broken. DESCRIPTION The rounded rhombic or subdiamond-shaped crown slopes gently up from the anterior and flattens out towards the posterior (Fig. 7L). The posterior margin is slightly pointed. The crown is covered in curved, concentric low ridges that parallel the lateral margins of the scale, and con- verge posteriorly, The central anterior part of the crown has short, curved, overlapping ridges; these short central anterior ridges and the anterior pn of the longer side ridges are grooved (Fig. K). The neck area is shallow to nonexistent, and is not indented. The base is diamond-shaped and gently convex (Fig. 7L,M). The anterior part of the base extends beyond the crown, producing a flat, unornamented rim along the anterolateral margins of the scale (Fig. 7K). The crown over- hangs the base posteriorly. Where the base joins onto the neck area, a rim is formed, especially at the posterior (arrow in Fig. 7L). DISCUSSION The crown ornamentation resembles that of Antarctilamna prisca Young, 1982 with con- centric ridges converging posteriorly, and grooves on the anterior part of the ridges. The Broken River scales differ from those of A. prisca Young, 1982 by having a large, diamond-shaped, convex base and shallow, broad neck; A. prisca Young, 1982 scales have a deeply concave, cup-shaped base and a constricted neck 110 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 12. Notiolepis dienemos gen. et sp. nov., SD204/174 upslope. A, longitudinal section QMF31912 of scale, x 45; B, longitudinal section QMF 31913 of scale, x 45; C, longitudinal section QMF31914, detail of posterior part of crown, x 90. MID-DEVONIAN CHONDRICHTHY AN SCALES TH (Young, 1982). The Broken River scales also possess several short, flat, curved ridges in the central anterior section of the crown, Such ridges are absent i some scales of A. prisca figured by Young (1982: pl. 87, fig. 7; lext-figs 2b, 4d); however, one illustrated scale (Young, 1982: pl. 87, fig. 6) shows (wo short anterior ridges, similar to those presentin Cladolepis. Enlarged photographs of the holotype of A. prisca Young, 1982 show several scales in the dorsal fin area with the short, overlapping, grooved anterior ridges characteristic of Cladolepis scales, so this type of scale varialion can occur on different parts of the body. Scales referred to Antarcrilamna sp. by Forey et al. (1992) from the Pragian or early Emsian of Saudi Arabia have superficially similar crown ùr- namentation to the Broken River specimens. They differ from both the Broken River and A. prisca Young, 1982 scales in having finer or- namentation, a larger number of the shorter anlerior ridges, and by having the anterior area of shorter, flatter ridges more clearly delineated from the rest of the crown. The scales from both Broken River and Saudi Arabia are a similar size, approximately 2mm; this is slightly larger than the A. prisca Young, 1982 scales from Antarctica. The thick, convex base of the Broken River specimens is similar to that of Ohiolepis new- berryi Wells, 1944 and Protacrodus wellsi Gross, 1973, but the crown ornamentation is different. Scales from a possible species of Cladolepis have been reported from the gigas Zone Mos- tyn Vale Formation near Keepit, NSW (Turner, 1993}, Overseas, the genus is known from the early Eifelian Lauch Formation of the Wollen- bach member (Vieth-Schreiner, 1983), the Middle Devonian of Indiana (Gross, 1973), the Cincinnati arch region (Wells, 1944), and the middle Givetian to early Eifelian Holy Cross Mountains of Poland (Liszkowski & Racki, 1993). Cladodont teeth are known in Australia, including the Broken River area (Turner, 1982), Írom Late Devonian and younger horizons. Chondrichthyan A - fam.,gen. et sp. indet. (Fig. 1OA-C) MATERIAL Figured scale QMF 261 10 and two other scales. LOCALITY AND HORIZON Scale QMF26110 is from section SD15, in the middle vareus Conodont Zone Papilio Forma- tion. The other scales occurred in the Papilio Formation of section SD131, dated middle varcux Conodont Zone, and the Spanner Limestone of section SD216, dated Aermanni-cristatus Con- odont Zone. REMARKS This chondrichthyan scale is not similar to any described form. An unusual feature is a thickened central ridge on the underside of the posterior section of the crown. Examination of more specimens will be required before the scales can be confidently assigned to a genus. MEASUREMENTS The scales measure 0.4mm to 0.75mm in length, 0.65mm to 1.2mmm in width, and 0.2mm to 0.4mm in height. The length is approximately twice the height, and the width is approximately three times the height. DESCRIPTION The crown is wider than long, and slopes up towards the posterior margin. The crown is gently curved at the front, and more pointed at the back. The anterior half of the crown has approximately 12-14 shallow, subparallel ridges, not clearly defined (Fig. 10A), Two or three narrow, low, closely-spaced ridges parallel the posterior mar- gins of the crown. The neck is shallow at the anterior, deeper and clearly indented at the posterior (Fig. 10B). The subtriangular base is flat or gently concave and flared into a thin trim around the edges (Fig. 10B,C), and, like the crown, 15 wider than it 18 long. Chondrichthyan B - fam.,gen, et sp, indet. (Fig. 10D-G) MATERIAL Figured scale OMF26111 and four uther scales, LOCALITY AND HORIZON The scales occurred in three sections (SD128, SD204 and SD210 - Fig. 2), in bonzons of middle varcus Conodont Zone age. REMARKS The anterior rim and ridges on the crown are similar to these found on scales of the acan- thodian Cheiracanrhoides compius Wells, 1944 (e.g. Vieth-Schreiner: 1983, pl: 4, fig, 32), How- FIG. 13. Aussilepis lukaso gen. et sp.nov., SD204/174 upslope. A, longitudinal section QMF26113 of scale showing base and central part of crown, x 60; B, longitudinal section QMF261 14, detail of posterior part of scale, showing base and crown, x 80. ever, these scales from Broken River do not have the shallow neck and deep rounded base typical of acanthodian scales (Turner, 1991). Rather, they have neck canal openings, a relatively high neck, anda flat, diamond-shaped base, distinctive MEMOIRS OF THE QUEENSLAND MUSEUM of shark scales (Turner, 1991; Turner & Young, 1987). MEASUREMENTS Dimensions of the one intact scale are length 0.6mm, width 0.8mm, and height 0.15mm. DESCRIPTION The crown is oval-shaped, thin, and flat (Fig. 10D-G). Seven subparallel ridges extend from the rim at the anterior edge to approximately halfway along the scale; the outer ridges extend into con- centric ridges parallel to the posterior margin of the scale (Fig, 10D). The neck is fairly deep, not indented, and has circular canal openings around the posterior edge (Fig. 10G). The base is a nar- row diamond shape, elongated across the width of the scale, and is flat (Fig, 10F). ACKNOWLEDGEMENTS Sincere thanks to Ruth Mawson and John Talent, who provided their material for study, and have given yaluable advice, support and en- couragement at all stages of this project. Special thanks also to Gavin Young and Sue Tumer for reviewing the manuscript and suggesting many improvements, and for providing much assis- tance and encouragement over a long penod; Theresa Winchester-Sceto for reviewing the manuscript, giving helpful advice and encourage- ment, and allowing me to read her unpublished manuscript; and Glenn Brock, who prepared Fig- ure 3, helped with the tables, and has given much useful advice. I thank Michael Engelbretsen, Coral Gilkeson, Annabelle Gregor, Jenny Nor- man, Mark Hocking and Ned Stephenson for assistance with SEM photography, developing and printing, Dianne Hughes for discussion of histology and use of her photographic equipment, Ken Rousell for drafting Figures 1 and 2, and Judy Davis for prompt assistance in the drafting office. Peter Jell's willingness to accept this manuscript at the last moment for publication is much appreciated. This is a contribution to UNESCO/TUGS:IGCP328-‘Palaeozoic microvertebrates”. LITERATURE CITED BLIECK, A. & GOUJET, D, 1991, Les vertêbrês du Dévonien inférieur d' Arville et de Nonceveux. Annales. Société Géologique du Nord |(2me serie): 67-78. BOUCOT, A.J, McCLURE, H.A., ALVAREZ, F. ROSS, J.R.P, TAYLOR, D.W,, STRUVE, W., MID-DEVONIAN CHONDRICHTHY AN SCALES SAVAGE, N.N. & TURNER, S. 1989. New Devonian fossils from Saudi Arabia and their blogeographical affinities. Senckenbergiana lethaea 69: 535-597, DENISON, R. 1979. Acanthodii. li Schultze, H.-P- text), “Handbook of Palaeoichthyology', Vol $, (Gustav Fischer Verlag: Stuttgart). FOREY, P.L, YOUNG, V.T. & McCLURE, H.A. 1992. Lower Devonian fishes from Saudi Arabia. Bulletin of the British Museum of Natural History (Geology) 48: 25-43, FRIMAN, L. 1983, Ohiolepis-Schuppen aus dem un- teren Mitteldevon der Eifel (Rheinisches Schiefer- gebirge). Neves Jahrbuch fiir Geologie und Palaontologie, Monatshefte 1983(4):; 228-236. GAGNIER, P-Y., TURNER, S., FRIMAN, L.. SUAREZ-RIGLOS, M. & JANVIER, P, 1988. The Devonian vertebrate and mollusc fauna from Seripona (Dept. of Chuquisaca, Bolivia). Neues Jahrbuch fiir Geologie und Paläontologie, Ab- handlungen 176: 269-297. GIFFIN, E.B. 1980, Devonian vertebrates from Australia. Postilla 180: 1-15. GROSS, W. 1947. Die Agnathen und Acanthodier des Obersilurischen Beyrichicnkalkes. Palacon- tographica A 96: 92-161, 1971a. Downtonische und Ditlonische Atan- thodicr-Reste des Ostseegebictes. Palicom 1ngraphica A 136:1-82. 1971b. Unterdevonische Theloduntier- und Acan- thodier-Schuppen aus Westauseralien Palliontologischen Zeitschrift 45; 97-106. 1973Kleinschuppen, llossenstacheln und zähne von fisches aus Europäischen und Nordamerikanis- chen bonebeds des Devons. Palacontographica A 142: 51-155. JANVIER, P. & MELO, J.H.G. 1992. New acanthodian and chondrichthyan remains from the Lower and Middle Devonian of Brazil. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1992(4): 193-206. KARATAJUTE-TALIMAA, V. 1973, Elegestolepis grossi gen. ct sp. nov., ein neuer typ der Placoidschuppe aus dem Oberen Silur der Tuwa, Palaeontographica A 143: 35-50. KLAPPER, G. & JOHNSON, J.G, 1980, Endemism and dispersal of Devonian conodonts. Journal of Paleontology 54; 400-455, LELIEVRE, H., JANVIER, P. & BLIECK, A. 1993. Silurian-Devonian vertebrate biostratigraphy of western Gondwana and related terranes (South America, Africa, Armorica-Bohemia, Middle East), Pp, 139-173, In Long, J,A. (ed.), ‘Palacozoic vertebrate biostratigraphy and biogeography”, (Belhaven Press; London) LISZKOWSKI, J. & RACKI, G. 1993, Ichthyoliths and deepening events in the Devonian carbonate plal- Torm of the Holy Cross Mountains, Acta Palacon- tologica Polonica 37: 407-426. LONG, J,A. 1991, The long history of Australian Fossil fishes. Pp. 337-428. Jn Vickers-Rich, P. 113 Monaghan, J,M., Baird, R.F. & Rich, T.H, (eds), ‘Vertebrate Palaeoniology of Australiesia'. (Monash University: Melbourne). MADER, H, 1986. Schuppen und Zahne von Acan- thodiem und Elasmobranchiem aus dens Unier- Devon Spaniens (Pisces), Göttinger Arbeiten zur Geologie und Paläontologie 28; 1-59, MAWSON, R. & TALENT, J.A, 1989, Late Emsiun- Givetian stratigraphy and conodont biofacies - carbonate slope and offshore shoal to sheltered lagoon and nearshore carbonate ramp - Broken River, North Queensland, Australia, Courier Forschungs-Institut Senckenberg 117: 205-259. OBRUCHEV, D. & KARATAJUTE-TALIMAA, V. 1967. Vertebrate faunas and correlauon of the Ludlovian-Lower Devonian in castern Europe. Journal of the Linnean Society (Zoology) 47(311); 5-14, OELOFSEN, B.W. 1981. The fossil record of the Class Chondrichthyes in Southern Africa, Palaeon- lologica Africana 24: 11-13. ORVIG, T. 1969. Vertebrates from the Wood Bay Group and the position of the Emsian-Eifelian boundary in the Devonian of Vestspitsbergen. Lethaia 2: 273-328. PICKETT, J., TURNER, S$. & MYERS, B. 1985. The age of marine sediments near Tumblong, souti- west of Gundagai. Geological Survey of NSW. Quarterly Notes. 58: 12-15. REIF, W-E. 1978. Types of morphogenesis of the der- mal skeleton in fossil sharks, Palsiontologischen Zeisehnift 52: 110-128, SCHULTZE, H-P. 1968. Palaconiscoidea-Schuppen aus dem Unterdevon Australiens und Kanadas und aus dem Mitteldevon Spitsbergens. Bulletin of the British Museum of Natural History (Geol- ogy) 16: 343-367. TURNER, S. 1982. Middle Palaeozoic clasmobranch remains from Australia, Journal of Vertebrate Paleomology 2: 117-131. 1986, Veriebrate fauna of the Silverband Formation, Grampians, Western Victoria, Proceedings of the Royal Soctety of Victoria 98: 53-62, 1991, Palaeozoic vertebrate mierofossils in Australasia, Pp. 429-464. In Vickers-Rich, P., Monaghan, J.M., Baird, R.F. & Rich, T.H. (eds), ‘Vertebrate Palaeontology of Australasia’. (Pioneer Design Studio; Melbourne), 1993, Palaeozoic microvertebrate biostraugraphy of eastern Gondwana. Pp. 174-207, In Long, J.A (ed.), 'Palaeozoic vertebrate biostratigraphy and biogeography”. {Belhaven Press: London). TURNER, S, & MURPHY, M.A. 1988, Early Devonian vertebrate microfossils [ram the Simpson Park Range, Eureka County, Nevada, Journal of Paleontology 62; 959-94. TURNER, S. & YOUNG, G.C. 1987. Shark teeth from the Early-Middle Devonian Cravens Peak Beds, Seong Basin, Queensland, Alcheringa 11; 233- 114 VIETH, J. 1980. Thelodontier-, Acanthodier- und Elas- mobranchier-Schuppen aus dem Unter-Devon der Kanadischen Arktis (Agnatha, Pisces). Gôttinger Arbeiten Geologie und Paläontologie 23: 1-69. VIETH-SCHREINER, J. 1983. Fisch-Schuppen und -Zähne aus der Eifeler Kalkmulden-Zone (Em- sium, Eifelium). Senckenbergiana Lethaea 64 : 129-177. WANG, N.-Z. 1984. Thelodont, acanthodian and chondrichthyan fossils from the Lower Devonian of southwest China. Proceedings of the Linnean Society of New South Wales 107: 419-441, WELLS, J.W. 1944. Fish remains from the Middle Devonian Bone Beds of the Cincinnati Arch Region. Palaeontographica Americana 3: 101- 161. YOUNG, G.C. 1982. Devonian sharks from south-east- em Australia and Antarctica. Palaeontology 25: 817-843. 1989. The Aztec fish fauna (Devonian) of Southern Victoria Land: Evolutionary and biogeographic significance. Pp. 43-62. In Crame, J.A. (ed.), ‘Origins and evolution of the Antarctic biota’, Geological Survey of London Special Publica- tion 47. MEMOIRS OF THE QUEENSLAND MUSEUM 1990, New antiarchs (Devonian placoderm fishes) from Queensland, with comments on placoderm phylogeny and biogeography. Memoirs of the Queensland Museum 28: 35-50, 1991. Fossil fishes from Antarctica. Pp. 538-567. In Tingey, R.J. (ed.), ‘The Geology of Antarctica’. (Oxford University Press: London). 1993. Middle Palaeozoic macrovertebrate biostratigraphy of eastern Gondwana. Pp. 208- 251. In Long, J.A. (ed.), ‘Palaeozoic vertebrate biostratigraphy and biogeography'. (Belhaven Press: London). YOUNG, G.C., TURNER, S., OWEN, M., NICOLL, R.S., LAURIE, J.R. & GORTER, J.D. 1987. A new Devonian fish fauna, and revision of post-Or- dovician stratigraphy in the Ross River Syncline, Amadeus Basin, central Australia. Bureau of Mineral Resources Journal of Australasian Geol- ogy and Geophysics 10: 233-242. ZANGERL, R. 1981. Paleozoic Chondrichthyes. In Schultze, H.-P. (ed.), 'Handbook of Palaeoichthyology', Vol 3a. (Gustav Fischer Ver- lag: Stuttgart). THE USE OF SPIDER SILK IN THE INITIATION OF NEST-BUILDING BY WEAVER ANTS (FORMICIDAE: FORMICINAE: POLYRHACHIS) PETER D. DWYER AND DANIEL PATRICK EBERT Dwyer, P.D. & Ebert, D.P. 1994 1201: The use of spider silk in the initiation of nest-huilding by weaver ants (Formicidae: Formicinae: Polyrhachis). Memoirs of the Queensland Museum 37(1): 115-119. Brisbane. ISSN 0079-8835. Field and laboratory observations of Polyrhachis doddi and P, pilosa in southeastem Queensland, Australia, showed that spider silk is used commonly in nest-building, is used often instead of larval silk when workers initiate nests or extend pre-existing nests and, aL least in the laboratory when larvae are unavailable, may be used to build entire nesls- Founding queens of both species also used silk from spiders when initiating nests. In nearly all cases, silk came from shelters of spiders such as salticids and clubionids. Silk fron) spider webs was rarely used, The functional and evolutionary significance of the use of spider silk are discussed. |] Polyrhachis, weaver ants, nest-building, spider silk, founding queens, Peter D, Dwyer & Daniel Patrick Ebert, Department of Zoology, The University of Queensland, Brisbane, Queensland 4072, Australia; 27 August 1994. Many species of the Old World formicine genus Polyrhachis use silk that is produced by their own larvae in the construction of nests (Hôlldobler & Wilson, 1983, 1990). This be- haviour is particularly evident in arboreal species of several subgenera (e.g. Cyrfomyrma, Myr- matopa, Myrmhopla) which build pocket nests of silk on tree trunks and the undersides of leaves or which use silk to join the leaves of various Irees and shrubs (Dorow & Maschwitz. 1990; Dorow et al., 1990; Kohout & Taylor, 1990; Ofer, 1970; Yamauchi etal., 1987). In nests of the latter kind, ws made by Australian Cyrtomyrma species for example, interior walls are covered by a thin, tough layer of silk and exterior portions, joining leaves, may be strongly reinforced hy debris. At completed nests, neither the larval silk nor the ants inside are visible. Nesi-weaving, as described in Polyrhachis species and in other weaver ants, poses two re- lated problems which have not received attention. First, in the earliest phase of colony formation, queens do not have larvae and, hence, lack access to larval silk with which to initiate nests. Second- ly, initiation of new nests by workers requires transport of larvae from sheltered sites to open situations where they may be exposed to both potential predators and exigencies of weather, One of us, for example, has observed substantial predation by the skink Lampropholis delicata upon Polyrhachis larvae during a mass move- ment from one arborcal nest to another. The skinks attempted to retrieve larvae from Polyr- hachis workers while avoiding direct contact with the workers themselves. Chemicals released by the ants are a powerful deterrent to these lizards (cf. Bellas & Hôlldobler, 1985). In the green tree ant, Oecophylla, founding queens do nol make nests; these ants are crypti- cally coloured and they rest, and commence rear- ing their first broods, in exposed positions on the undersides of leaves (Dorow et al., 1990; Peeters & Anderson, 1989; Wheeler, 1915). Again, when Oecophylla workers initiate nests, the security of exposed larvae is ensured firstly by the recruit- ment of many workers that may defend those larvae and, secondly, by the rapidity with which nests may be built (Hôlldobler & Wilson, 1990). Neither of these options is available io species of Polyrhachis (Cyrtomyrma) which are black and conspicuous and do not recruit large numbers of workers to new nest sites. In this paper we report that, in two Australian Cyrtomyrma, a common solution to potential problems associated with initiating nests entails the use of silk produced by spiders as a substitute for that produced by ant larvae. The use of alrernative silks by these weaver ants has not previously been recognised. Indeed, Dorow et al, (1990: 184) wrote thal, in the laboratory, the Southcast Asian P. muelleri did not accept silk from spider webs. Our obser- vations have significance for both functional and evolutionary interpretations of weaving be- haviour in Polyrhachis, STUDY AREA AND METHODS In southeastern Queensland, P. doddi and P. pilosa are abundant species of arboreal weaver ant. Both make nests in many different species of plants of diverse leaf form, For example, 60 nests 116 of these species censused in May 1993 at ihe Mt Coot-tha Botanic Gardens, Brisbane. were lo- cated in 30 plants of 23 species in 10 families (17 nests in Myrtaceae, 16 in Arecaceae and |2 in Theaceae; C. Hunter, pers. comm). Nests of these species are also found, among other places, in hollows of dead Plumeria branches, leaf bases of bromeliads, petioles of climbing Philodendron, rolls of Eucalyptus bark and a variety of artificial sites (e.g. plastic buckets and covered plant pots suspended in trees), From January 1993 to February 1994 we made opportunislic observations of nest-building by P, doddi and P. pilosa in suburban gardens, and at the St, Lucia campus of the University of Queensland, Brisbane. From September 1993 to February 1994 we also made observations of nest-building by captive P. doddi. Groups of 25 to 55 workers were established in 28 x 21 Xx 7cm plastic trays that were isolated by moats, (In the field P, doddi and P. pilosa initiate nests using groups of Jess than 20 workers). The trays were weighted with sand, sticks were provided as walkways and plant debris (very small leaves and broken seed capsules of Melaleuca and Lep- tasperrnum) was supplied. Each tray was provided with a small artificial tree, less Than or equal to 30cm high, consisting of two or three branches of * silk’ (rayon) Ficus, Artificial trees were made in sucha way that parts ofsome leaves of different branches were 0.5-1.0cem apart but not in contact. Leaves were aligned more or Jess vertically, Fibres from the * silk’ plants were hot used im construction of nests. Sampled nests were from Buckinghamia celsissima and Casuarina linoralis, Ants were fed on a mixture of egg and honey according to the recipe in Hélldobler & Wilson (1990: 632). All observations were at lemperalures between about 15 and 30°C. RESULTS FIELD OBSERVATIONS Spider silk is often visible in the walls of new F. pilosa and P, doddi nests before these walls are covered with debris. Spider silk is sometimes included with plant fragments, insect frass and fragments of chitin as part of the outer, protective cover of nests. Workers of both species have been seen harvesting silk from shelters built by spiders and transporting it to nest sites. In the field, we have not seen larvae being used in open siluations during the earliest phase of nest-building. P. pilosa have been observed to initiate nests without the ard of larvae on at least six occasions. MEMOIRS OF THE QUEENSLAND MUSEUM These nests were made in ôr between leaves of Syzygium luehmannii, Backhousia citriodora, Howea forsteriana, Mangifera indica, Caryota mitis and Brachychiton acerifolium. In the first three cases, 10 to 20 ants used silk from shelters of unidentified spiders; they worked for two to four days to provide partially enclosed spaces before carrying larvae to the nest sites. The fourth nest was initiated, but not completed, in a dead, furled leaf of Mangifera indica caught in the abandoned web of a Nephilu (Araneae), Ants manipulated the web to increase leaf curyalure and attached debris to strands of web that joined the edges of the leaf. After three days, when a partial shelter had been formed, one larva was carried to the new nest. The fifth nest, built by a founding queen, was in a rolled leaflet of Caryota mitis, this leaflet had originally housed a spider shelter and silk from the shelter, together with a little attached debris, formed the exposed walls of the nest, The sixth nest was initiated by P, pilosa workers that used waxy filaments secreted by a cocci (Homoptera) to join the leaves, Sites chosen for building nests in two large stands of Phoenix roebelenii, onc stand occupied by each species of Polyrhachis, contained shel- ters of salticid and clubionid spiders, including those of Opisthoncus and Clubiona. These spider shelters were spun between adjoining leaflets of fronds and, particularly with the tightly woven shelters of salticids, had the outcome that the leaflets were pulled together and overlapped for part of their length. Pelyrhachis workers with larvae cannot achieve this rearrangement of the thick, stiff leaflets (personal observations). Be- cause the orientation of leaflets at nearly all Polyrhachis nests in these palms matched the onentation produced by spiders we think that most observed nests (mn = 12) were initiated at spider shelters. At one P. doddi nest the 16cm long walls that joined two leaflets were made entirely of silk from the shelters of spiders. Workers were seen harvesting silk from a nearby salticid shelter and carrying it to the nest, Two large, galleried nests of P. doddi in Casuarina cristata and one small nest in Syzygium luehmannii were extended by workers that carried silk from spider shelters to the sites: larvae were not used until the extensions were Well underway and provided covered areas within which workers could effectively hide the larvae from view. Four nest sites used by founding queens of P, doddi were established at spider shelters in furled leaves of Tupidanthus calyptratis and Schefflera actinophylla. The USE OF SPIDER SILK BY POLYRHACHIS queens had teased apart the spider shellers and attached some debris to make open nests that, in two cases, were used for more than a month by up to eight queens. Founding queens of P. doddi have been often observed sheltering under or inside palm leaflets which had been either folded by spiders making silk shelters or rolled by the larvae of palmdarts (Cephrenes). In the latter case, abandoned Cephrenes shelters that had sub- sequently been used by spiders were preferred by the queen ants. When using temporary shelters during the daytime, P. doddi queens are often in contact with the silk of spider shelters, either between leaves or jn leaves that have been curled by the spiders. LABORATORY OBSERVATIONS In mid-September 1993, three groups of workers, each with access to an artificial tree, were provided with silk from old, abandoned salticid and clubionid shelters. Three additional groups were provided, respectively, with spider web and egg cases (the latter from Zosis sp.[Uloboridae: Araneae}), cotton wool and five large larvae, All groups, except that with cotton wool, initiated nest-building in late September and continued intermittently through Octoberand November. In late November the group with lar- vae and two of the groups with salticid and clubionid shelters had completed building nests between leaves of the artificial trees, After several aborted attempts, the group with web and Zosis egg cases used silk from the egg cases to complete a small nest beneath a stick on the floor of the tray. On December 24, 1993, three groups of workers in artificial trees were provided with silk from recently built salticed and clubjonid shelters. On February 8, 1994, these groups of ants had completed two-thirds, one-quarter and one half, respectively, of the areas of wall needed to enclose each nest. The trial was then terminated because, in two groups, larvae produced by workers were approaching the size at which they might produce silk, Donor colonies of P. doddi, housed in the laboratory, also built nests using silk from salticid and clubionid shelters. These large groups, with queens and larvae, initiated nests with spider silk more often than with larval silk. One group took apart, and reused, the entirety of an old nest; in another goup, with about 200 workers, ants began harvesting spider silk 10 minutes after the silk was provided, 117 In the laboratory, P. doddi workers harvested silk from spider shelters by tearing al the shelter and pulling and biting portions free, They did not cooperate in this activity. Often, however, when the load had been carried to the nest site it was attached jo an artificial leaf at places which had becn moistened by fluid from the mouth of another worker, Sometimes the ant that carned the spider silk waited as the other ant moistened the leaf and, on a few occasions, the latter ant retneved the silk from the former and attached it to the leaf, This behaviour differs from that of Polyrhachis muelleri which, when using old lar- val silk or cotton wool, glued the material into place with a droplet from the tip of the gaster (Dorow et al., 1990) P. doddi seldom isolated single fibres from the masses of spider silk that were transported to nest sites. They teased each load to produce a flocculent mass of fibres. These were either built out from one leaf surface in layers or stretched between two leaves by manipulating with man- dibles and antenor legs. Plant debris was inserted into the mesh of fibres rather than being stuck onto it as occurs when larval silk is used. In two cases, when a vertical wall was built from spider silk the weight of attached debris contributed to pulling silk fibres from the upper to the lower leaf edge. In the laboratory, all P. doddi nests built between vertically aligned leaves were com- menced at the bottom, proceeded through arch- ing, vertical walls and were completed by covering the top of the chamber and making an entrance. Nests made in analogous situations in the field were also built in this sequence. Nests made entirely from spider silk were not as strong as those that incorporated larval silk. The latter material provides a firm base to which debris can be attached with some precision. Walls constructed from spider silk lack rigidity and cannol support a complete cover of plant debris without collapsing. Nor can workers construct the characteristic well-formed circular entrance tò the nest without access to larvac. The quality of laboratory built nests, made from spider silk alone, is such that they would fall apart rapidly if exposed to even moderate rain or wind, DISCUSSION Field and laboratory observations of P. doddi and P. pilosa show that silk produced by spiders is used very commonly in nest-building, is used often instead of larval silk when workers initiate nests or extend pre-existing nests and, at least in 118 laboratory settings, may be used to build entire nests, In all these cases, Polyrhachis workers actually manipulated or transported spider silk in the process of nest-building, We are not dealing merely with the use of * preformed shelters of spiders and caterpillars’ which Maschwitz ct al. (1991: 308) reported for an arboreal dolichoderine ant (Technomyrmex sp.); they commented that sites of these kinds might be easily ' misidentified as nests produced by the ants themselves’. Founding queens of hoth P. doddi and P. pilosa have been also observed to use silk from spiders when initiating nests. In- deed, there appears to be a close association be- tween P. doddi queens and abandoned spider shelters. Our observations suggest that silk from spider shelters is much more likely to be used by P, doddi and P. pilosa than silk from spider webs. Nests of these species are, however, often built where leaves are tangled in web and we have sometimes seen P. pilosa tugging àl and manipulating silk in webs that were occupied by spiders. Dorow et al. (1990) reported that * silk from spider webs’ was not accepted by nest- building P. muelleri; it is possible, however, that iiis: species would accept other forms of spider sal Arboreal Polyrhachis species do not display the highly cooperative nest-building behaviour seen in Oecophylla where large numbers of ants, recruited to favourable sites, form lines or hang- ing chains and bend or manoeuvre leaves such that larval silk can be used to hold those Jeaves in their new positions (Hélldobler & Wilson, 1977, 1990). Rearrangement of leaves observed at many arboreal nests of both P, doddi and P. pilosa, especially in suff-leafed plants (e.g. some palms), could not be achieved by the ants them- selves, either with or without assistance from larvae. Most of these nests will have been in- itiated at places where leaves had been already repositioned through the actions of other animals, usually by spiders spinning silk shelters. By con- trast, in various Myrtaceae species, Polyrhachis workers preferentially build nesisin young leaves which are easily bent into unconventional pasi- tions; even in these plants, however, our observa- tions were only of nests imbated or extended using spider silk and not larval silk. Dorow & Maschwitz (1990) described rapid recruitment of P. hodgsoni workers and larvae to potential nest sites on broad-leafed bamboo (Gigantochloa and Schizostachyum), anis opportunistically ex- ploited occasions when changes in turgor caused MEMOIRS OF THE QUEENSLAND MUSEUM edges of a leaf to move towards each other, These authors also reported that nests of P. hodgsoni were also built in leaves that had been rolled by spiders, orthopterans, caterpillars and climbing plants. At exposed sites, where larvae would be easily visible, P. doddi and P. pilosa usually commence nesi-building without recruiting lar- vac to the site. With specific reference to Oecophylla, Hôlldobler & Wilson (1990: 618) argued that weaving behaviour freed ants from ` the spatial limitations imposed on species that must live in beetle's burrows, leaf axils ... and other preformed vegetation cavities” (see also Dorow etal., 1990, on Polyrhachis and Maschwitz et al., 1991, on Dolichoderus). Dorow etal, (1990: 188) commented further that in arboreal Polyrhachis, as in Oecophylla, weaving behaviour facilitates polydomy which, itself, may be beneficial in reducing distances to food sources, providing allernative nest siles after catastrophes and, per- haps, facilitating defence of foraging areas. Ob- servalions reported here both reinforce and extend earlier conclusions. Use of spider silk as an alternative to larval silk will increase the availability of nest-building sites to P- doddi and P, pilosa and allow assessment of the quality of potential nest and foraging sites by workers prior to incurring the costs of transporting larvae. Potential costs to larvae that would arise from silk production and risks associated with exposure will be also reduced when silk from spiders is used to initiate nests. And, further, our observa- tions show how founding queens, which lack access to larval silk, may initiate arboreal nests. Hilldobler & Wilson (1990) considered that the exploitation of larval silk by workers released weaver ants from their ancestral dependence upon a variety of preformed vegetation cavities. This opinion is difficult to accept if, as Dorow er al, (1990) suggested and our observations imply, the failure of Cynomynna species to make cocoons is independent of the amount of silk contributed to nest-building. The behaviour of both workers and founding queens of P. doddi and P. pilosa suggests that use of spider silk could have been a precursor to use of larval silk in the urigin of nest-building by at least some groups of arboreal Polyrhachis, Answers to questions regarding the origins of these behaviours will require comparative knowledge of silk produc- tion in the genus as a Whole and knowledge of the extent io which silk production is under worker or larva control. USE OF SPIDER SILK BY POLYRHACHIS ACKNOWLEDGEMENTS We thank Rudy Kohout, Geoff Monteith, Val Davies and Yvonne Ross for help with the iden- tification of Polyrhachis, other insects, spiders and plants, Chadden Hunter for providing data on nest composition and Ross McKinnon for authorizing censuses at the Mt. Coot-tha Botanic Gardens. Special thanks to Jeffrey Willmer for frequent access to his garden, plant identifica- tions and a keen ability to locate ants in hard-to- see places. Discussion with Simon Robson rekindled an interest in Polyrhachis and Monica Minnegal commented on the manuscript. LITERATURE CITED BELLAS, T. & HOLLDOBLER, B. 1985. Constituents of mandibular and Dufour’s glands of an Australian Polyrhachis weaver ant. Journal of Chemical Ecology 11; 525-538. DOROW, W.H.O. & MASCHWITZ, U. 1990. The Arachne-group of Polyrhachis (Formicidae, For- micinae): Weaver ants cultivating Homoptera on bamboo, Insectes Sociaux 37: 73-89, DOROW, W.H.O., MASCHWITZ, U. & RAPP, S. 1990. The natural history of Polyrhachis (Myrmhopla) muelleri Forel 1893 (Formicidae Formicinae), a weaver ant with mimetic larvae and an unusual nesting behaviour. Tropical Zoology 3: 181-190. 119 HOLLDOBLER, B. & WILSON, E.O. 1977, Weaver ants. Scientific American 237: 146-154. 1983. The evolution of communal nest-weaving in ants. American Scientist 71: 490-499, 1990. ‘The ants’, (The Belknap Press of Harvard University Press: Cambridge). KOHOUT, R.J. & TAYLOR, R.W. 1990, Notes on Australian ants of the genus Polyrhachis Fr.Smith, with a synoptic list of the species (Hymenoptera: Formicidae: Formicinae). Memoirs of the Queensland Museum 28: 509-522. MASCHWITZ, U., DUMPERT, K., BOTZ, T. & ROHE, W. 1991. A silk-weaving Dolichoderine ant in a Malayan rain forest. Insectes Sociaux 38: 307-316. OFER, J. 1970. Polyrhachis simplex the weaver ant of Israel. Insectes Sociaux 17: 49-81. PEETERS, C. & ANDERSEN, A.N. 1989. Cooperation between dealate queens during colony foundation in the green tree ant, Oecophylla smaragdina. Psyche 96; 39-44, WHEELER, W.M. 1915. On the presence and absence of cocoons among ants, the nest-spinning habits of the larvae and the significance of the black cocoons among certain Australian species. Annals of the Entomological Society of America 8: 323- 342. YAMAUCHI, K., ITO, Y., KINOMURA, K. & TAKAMINE, H. 1987. Polycalcic colonies of the weaver ant Polyrhachis dives, Kontyu 55: 410- 420. 120 NEW CONSERVATION STATUS FOR ANOMALOPUS PLUTO (SCINCIDAE). Memoirs of the Queensland Museum 37(1):120. 1994 :- Recently (17 May 1994) the authors col- lected three specimens of Anomalopus pluto on the Glennie Tableland (13°27'S 142758'E), FNQ. Two were found at 260m, under sandstone slabs on a narrow, flat sandstone ridge, in dark sandy loam and humus. This is an area of dense shrubland with low, windswept trees. The third (collected 18 May, 1994) was found under a rotting log in a patch of monsoon vine forest in the same soil, at the same altitude. These specimens were photographed and released. Copies of the photographs have been lodged in the Queensland Museum photographic index (NP 79-83). These specimens extend the range of this species almost 200km south. Couper (1992) reviewed the range and status of A. pluto following its collection in the Heathlands area (1 1745” 142°35’). He treated itas a '2K' species under the conservation status definitions used by McDonald et al,, 1991, That is, as a species '... with a very restricted distribution ... and with a maximum geographic distribution of less than 100km ...”; poorly known ... accurate field distribution information + inadequate’. NEW DATA ON THE DISTRIBUTION OF PYGOPUS LEPIDOPODUS (PYGOPODIDAE). Memoirs of the Queensland Museum 37(1):120. 1994:- That Pygopus lepidopodus occurred north of the Yeppoon area, MEQ (23°08'S 150°44'E) was unknown until 1991. Ingram & Raven (1991) noted its occurrence at Yungaburra (QMJ3899, 17°16’S 145°35'E) and Shipton's Flat (QMJ47145, 15°48'S 145*15'E), NEO. Two more specimens have been recorded recently, on northern Cape York Peninsula; OMJ57778, Heathlands, 11°44’8 142°32'E, FNEO, B. Lyon, M. Cohen; OMJ58986, llkm NE Heathlands Ranger Station, at 150m, 11°41'S, 142°39’E, FNEO, P.J. Lethbridge, T.A. Hawkes. This is a northern extension of the range of this species by 600km. Both Heathlands specimens were found in low heath with sclerophyll emergents on sand. MEMOIRS OF THE OUEENSLAND MUSEUM Discovery of A. pluto on the Glennie Tableland changes its conservation status to “3K': '... with a range greater than 100km ... but occurring in small populations which are mainly restricted to highly specific habitats; poorly known ...”. Acknowledgements Patrick Couper and Jeanette Covacevich assisted us to prepare this note. Literature Cited Couper, P.J. 1992. Anomalopus pluto Ingram, a poorly known skink from Cape York Peninsula. Memoirs of the Queensland Museum 32(1): 54. McDonald, K.R., Covacevich, J.A., Ingram, G.J. & Couper, P.J. 1992. The status of frogs and reptiles. Pp.338-345. In Ingram, G.J. & Raven, R.J. (eds), 'An atlas of Oueensland's frogs, reptiles, birds and mammals'. (Board of Trustees, Oueensland Museum: Brisbane). T.A. Hawkes, P.J. Lethbridge, M, Anthony & M. McGregor, Cape York Herpetological Society, PO Box 114, Palm Cove (Cairns), Queensland 4879, Australia; 30 September 1994. Acknowledgements Patrick Couper and Jeanette Covacevich provided data for this note, Literature Cited Ingram, G.J. and Raven, R.J. 1991. *An atlas of Oueensland's frogs, reptiles, birds and mammals.’ (Board of Trustees of the Oueensland Museum: Brisbane). P.J. Lethbridge & T.A. Hawkes, Cape York Herpetological Society, PO Box 114, Palm Cove (Cairns), Queensland 4879, Australia; 30 September 1994. COURTS AND SEASONAL ACTIVITIES AT THEM BY MALE TOOTH-BILLED BOWERBIRDS, SCENOPOEETES DENTIROSTRIS (PTILONORHYNCHIDAE) CLIFFORD B. FRITH AND DAWN W. FRITH Frith, C.B. & Frith, D.W. 1994 12 01: Courts and seasonal activities at them by male Tooth-billed Bowerbirds, Scenopoeetes dentirostris (Ptilonorhynchidne), Memoirs of the Queensland Museum 37 (1):121-145, Brisbane, ISSN 0079-8835, Court seasonality and maintenance of male Tooth-billed Bowerbirds Scenopoeetes den- Hrostris were studied over the display seasons of 1978, 79 and 80 in a SOha upland rainforest study area on the Paluma Range, north Queensland. Seasonality of court establishment was also recorded during nine subsequent seasons. Initial court establishment varied from season to season, the abundance of fruit in the sy area having a greater influence than oe yi and/or rainfall, Lack of rain and associated higher temperatures and fruit crop size influenced court decoration and maintenance levels at established courts during the display season, The onset of the wet season usually terminated court activity but significant dry spells thereafter caused a brief return to courts and their maintenance. Excessively dry, seasonally abnormal conditions caused a cessation of court activity altogther, The size of the cleared court area of more established courts of older males varied greatly, the largest being 2.75 X 1.95m and the smallest 1.0 X 0.8m, Partly established, temporary, courts of presumed younger males were smaller. Mean number of Jeaves on courts during peak display activity was 69. Of newly placed fresh leaves 79% were removed from courts within a week, During peak display activity a mean of 6.3 leaves werc placed on courts/day and a mean of 6,9 removed, The turnover of leaves as court decorations was highest during hot dry spells. Leaves of Polyscias australiana were conspicuously favoured as court decoration by older males and the vast majority of leaves stolen from courts were of this plant, That leaves of a given tree species favoured by males as court decorations varied between age classes of males and geographical areas suggests the influence of local tradition based on imitive learning, as has been recorded in males of other polygynous bower-building bowerbirds. Results of leaf-marking experi- ments lend some support to the hypothesis that a male social hierarchy is established and maintained within groups of more densely dispersed and possibly older males, by leaf stealing. Leaf stealing was most frequent between immediate neighbours, more so between those with smallest nearest neighbour distance Attendance levels and behaviours of male Tooth-billed Bowerbirds at their courts were monitored throughout display seasons 1979, 80 and 89. Males spent an average 64% (range 52-79%) of daylight period 0600-1800 at their courts during peak display activity. Duration of a court visit averaged 23mins, most of which (96%) was spent singing from favoured court perches, the remaining time being spent decorating/maintaining the court (1,290 of total time) or displaying (<1% of total time). Males adopted a static posture when an unidentified intra- or inter-specific bird visited their court. Males performed Joud and prolonged advertisement song followed by quietand brief display vocalisations at courts, Advertisement song included a large proportion of avian mimicry the repertoire of which was added to opportunistically, Loud and complex vocalisations appear to substitute for epigamic plumage and a sophisti- cated bower in this cryptically-plumaged sexually monomorphic, simplistic court-clearing polygynous bowerbird. This represents an atypical sct of evolutionary adaptations to polygyny in the bowerbirds thatis, however, paralleled in several similarly atypical members of the evolutionary convergent but non-bower-building polygynous Cotingas (Passerifor- mes, Cotingidae) of thé neotropics. [_] Court seasunality, leaf decoration, leaf turnover rates, leaf theft, male attendance, court behaviour, court vocalisalians, Queensland, Australia, Toorh-billed Bowerbird, Scenopoeetes dentirosiris. Clifford B. Frith & Dawn. W. Frith, Honorary Research Fellows af the Queensland Museum, 'Prionodura', P.O. Bax 581, Malanda, Queensland 4885, Australia; 2] February 1994. The Tooth-billed Bowerbird Scenopoeetes (19°30'S, 146°57'E) just south of Townsville dentirostris lives in upland rainforests at 600- (Frith & Frith, 1985a; Nix & Switzer, 1991). The 1400m above mean sea level, from Mt Amos Tooth-bill is an atypical bowerbird in that al- (15742'S, 145*18'E) southward to Mi Elliot though males are promiscuous and females uniparental, it is cryptically plumaged, sexnally monomorphic (Gilliard 1969, Donaghey et al., 1985; Frith & Frith, 1985a, 1989, 1990a,h, 1993, 1994) and males form exploded leks (Frith, 1993), Unlike males of other polygynous bowerbird species, male Tooth-bills do not construct stick howers but instead clear leaf litter from an area on the forest floor (Borgia, 1986). This 'court” encompasses the trunk of at least one “display tree’. The males place leaves, with their paler sides uppermost, onto the cleared area and per- form courtship displays upon it (Frith & Frith, 1993), The court location is advertised by loud male vocalisations that deter nvals and attract potential mates, and the arrival of a female Tooth- bill stimulates the male to drop onto his court and display (Marshall, 1954; Warham, 1962; Frith & Frith, 1993), It is assumed that females assess male quality on the basis of vocalisations, dis- plays, courts, decorations and perhaps male atten- dance (time investment) before soliciting and mating. Outside the display season Tooth-bills are mostly silent and secretive inhabitants of the forest canopy. That male bowerbirds steal decorations from the bowers of nval males, with a preference for articular colours and items, has long been wn (Marshall 1951, 1954 & references there- in), but has been only briefly alluded to with the Tooth-bill (Frith & Frith, 1993). In recent years buwer decoration theft has been studied inten- sively and it has been demonstrated that males steal predominantly from immediate neighbours (Borgia, 1985a), preferentially steal items rare in the birds’ environment (Borgia & Gore, 1986; Frith & Frith, 1990c) and that the greater numbers of more favoured or stolen decorations on a bower positively influenced relative male mating success (Borgia, 1985a,b; Pruett-Jones & Pruett- Jones, 1993). Froth & Frith (in press) found that 71% of individually-marked (n = 24) male Tooth-bills attended only one court site, that mean annual survival rate averaged 88% and mean expectancy of further hfe after banding was 8.1 years. Courts of Tooth-bills have been known to persist at one site for 16 years (Frith & Frith, in press), those of the Satin Bowerbird Prilonarhynchus violaceus for up to 30 years (Vellenga, 1980), and those of the Great Bowerbird Chlamydera nuchalis and Golden-Bowerbird Prionodura newtoniana lor more than 20 years (Frith & Frith, unpubl. data). Temporal and spatial animal distributions are considered indicative of environmental pressures MEMOIRS OF THE QUEENSLAND MUSEUM and the presence/absence of conspecifics (Brown & Orians, 1970). Whilst studies of bowers and/or bird activities at bowers ure numerous, no study has fully documented male bower maintainance seasonality over one or more seasons and at- tempted to relate the so defined 'bower” or “display” season to environmental factors such as climate and food resources, Marshall (1951) noted that the Tooth-bill display season varied from year to year. Vellenga (1970) observed that male Satin Bowerbirds in New South Wales had a display season of early August to Jate October, and that year to vear variations possibly related to the irregular Flowering of banksias and eucalypts and summer bushfires ’ which affected food supplies’. No studies have attempted to re- late such seasonality to the temporal availability of females which, in the case of the sexually monomorphic and secretive Tooth-bill, would be extremely difficult. The display season of the male Touth-bill starts with calling and court clearance. On the Paluma Range the season started in September or early October (Frith & Frith, 1993). The display season continues through to January or early February. Adult males perch above or adjacent to their court where they are conspicuously vocal, incorporat- ing fine mimicry of other bird calls, and other sounds, into their repertoire (Frith & Frith, in press), All bowerbirds regularly attending and vocalizing at bowers have proved to be male (Marshall, 1954; Vellenga, 1980; Gilliard, 1969; Cooper & Forshaw, 1977; Frith & Frith, 1993). As there is no evidence that female bowerbirds attend and vocalize at bowers for any length of time we assume that all Tooth-bills doing so were male, The results of our long-term observations of Tooth-bills support this assumption (Frith & Frith, in press). Male Tooth-bills spend most time at their courts performing loud and prolonged court-ad- vertisement yocalisauons, including avian mimicry, from several favoured perches immedi- ately aboye or adjacent to the court (Marshall, 1954; Cooper & Forshaw, 1977; Warham, 1962). Remaining time at courts is spent maintaining andor decorating the court or displaying. Most displays are performed on the court, are started at the base of a display trec, and are accompanied by a Complex 'subsong' of avian mimicry (Frith & Frith, 1993), As interspecific mimicking by Tooth-bills has long been noted or denied in the literature (Frith & Frith, 1993) we provide both qualitative and quantitative data of it. TOOTH-BILLED BOWERBIRDS Males of other polygynous passerines that at- tend and maintain a traditional courting perch(es) of area, 10 which they attract females by vocalisa- tions, have been found to be predomiantly frugivorous or nectanvormous and, as a conse- guence, have been able to devote a high propor- tion of daylight hours to attending and advertising their traditional location (Snow, 1962, 1976, 1982; Snow & Snow, 1988). For some predominantly insectivorous polygynous pas- serines, however, see Bechler (1989). In this paper we describe the characteristics of Tooth-bill courts with reference to leaf decora- trons, leaf turnover rates and inter-court leaf theft rates and pattems. Court seasonality and main- tenance over three display seasons (1978-80) arc discussed in relation to fruit crop, rainfall and temperature. Seasonality of court establishment was also recorded during nine subsequent seasons. Male attendance levels, behaviours and vocalisations were monitored over three display seasons (1979, 80 and 89). METHODS STUDY AREA, CLIMATE AND PHENOLOGY The study was performed in 50ha of upland tropical rainforest, altitude c, 875m, on the Paluma Range (19°00'S, 146°10°E), NE Queensland, 7km from Paluma Township and 80km north of Townsville, The rainforest, clas- sified as simple notophyll vine forest by Tracey (1982), occurs at c.600-1000m altitude. Because the Oueensland Forestry Department selectively logged the study area c.1938, the canopy was uneven and disturbed, varying from c.17.5-25m high, with taller emergenis. The undersiorey was dominated by lawyer palms, Calamus spp., ter- restrial ferns including Cyathea spp., and sa- plings. Medium-sized woody lianes and climbing pandans, Freycinetia spp., were common. The 1 x 0.5km forested study ares (Fig. 1) was permanently gridded using compass, measuring tape and metal stakes, 25ha into 25 x 25m squares and the remainder into 25 * 5Ôm rec- tangles. Topography to the north of the road was a flattish ndge c.30-50m wide and 600m long with a discrete knoll (= Tooth-bill Hill). Other terrain to thc north was steep and dissected by gullies. Of the area to the south of the road, that to the NE was flat with less dense forest inter- spersed with disturbed patches of Calamus- dominated undergrowth. The central southern area sloped gently from the road into a system of creeks, with a few low ridges. The west end was 123 another rise, less discrete than Tooth-bill Hill, contiguous with a flattish c.75m wide ridge slop- mg down foward the southern comer of the area, Annual rainfall and temperature show marked seasonality, The dry season extends from April- November with June-August being the driest and coldest months. The hotter wet season is from December-March. Rainfall figures were provided by the Paluma meteorological substa- tion for the years 1978-1990, Mean annual ràin- fall was 2379mm «during the study period. Maximuni-minimum temperatures were recorded in a standard screen located 20m from the edge of the rainforest from 1 August 197%- February 1981. To obtain fruiting phenology data we cxamined 602 trees from September 1978- Apnl 1979, and thereafter c,500 of these trees at six-weekly (July 1979-August 1980) or eight- weekly (November 1980-February 1981) inter- vals, Only trees with a girth ut breast height of >32cm were examined in Wo sample plots, each measunng 500 X 5m, along ridges inhabited by court-owning Tooth-bills. DEFINITIONS We refer to a display season by the year in which it started, By ‘court’ we mean a cleared terrestrial area and ‘site’ describes the location of a court or its immediately adjacent replacement court(s). A “traditional” court site was one at- tended in the same location for at Jeast two years. A ‘traditional court owner’ was an individually marked court-attending bird known to attend that court in a previous season(s). A ‘court tree” is any irce or sapling rooted within the cleared court area anda ‘display tree’ is an individual tree or sapling trunk rooted within a court area that is used by a male in initial display activity (Frith & Frith, 1993). Attendance refers to known, individually marked, males perching, calling, displaying or maintaining their own court. Thus # male visiting the court of another to steal a leaf was not attend- ing it. As we could see only the male during most displays we refer to them only as ‘displays’ as distinct from ‘courtship displays” COURT SEASONALITY AND MAINTENANCE Intensive fieldwork was performed from | August 1978-28 February 1981, apart from | May-I8 July 1979 when we were absent. This included the three display seasons 1978, 79 and 80 and non-display months March-April 1979 and March-August 1980, We divided the display seasons 78, 79 and 8O inte three peniods based on the degree and kind of activity at courts. *Pre- 124 Gully Toolh-oill Hill 800m ate #1 +37 Eaa * as’) 4 FIG, 1. The fifty hectare study area on the Paluma Range. Note; = location and reference number of court sites examined for leaf tumover rates, inter-court theft and/or male attendance at courts. See Frith & Frith (in press) for locations of all court sites within the study area. Solid black line = road. peak’ was the period of courtestablishment, com- mencing with the first sign of litter clearance and decoration (courts partly established) and con- cluded when they were cleared and decorated (well-established). We examined courts as they were established and recorded the extent of clearance and number of leaf decorations. “Peak' activity was the penod of regular court main- tenance and attendance, commencing when courts were well established and continuing until courts began to deteriorate. "Post-peak' activity was the period of declining court maintenance and attendance, when decoration leaves rotted and courts became covered by fallen leaves. Court maintenance levels were assessed by counting leaves on five adjacent courts, initally chosen at random, every seven to 12 days throughout the peak and post-peak periods of 79 and 80. Court attendance levels by males were assessed in two ways. Faeces were collected at weekly intervals during peak and post-peak periods from fine black mesh traps suspended beneath favoured perches that were above or immediately adjacent to ten courts. In addition, court attendance levels were monitored by direct observations, by use of a tape-recorder, or both. COURT CHARACTERISTICS AND DECORATIONS All courts were measured during 1989, from the cleared court edge, directly behind the display tree, to the opposite court perimeter and at right angles to this across the widest part of the court. Torank court sizes we multiplied these two meas- urements. The relative position of display trees and their diameter at breast height and the Ioca- MEMOIRS OF THE QUEENSLAND MUSEUM tion of court trees and lawyer canes, Calamus spp., within the court were noted. Numbers of leaves in use as decorations were counted over several con- secutive days during the earlier part of peak activity of seasons 79, 80 and 81 and during the late peak activity in 86 and 89. Leaves were identified to species when possible, LEAF TURNOVER RATES AND THEFT Decoration leaves were counted on five adjacent courts (sites 6, 32, 33, 35, 37) on Tooth-bill Hill and adjacent slopes (Fig. 1). Counts were made during peak and post-peak activity at seven day intervals during 79 and at 7-12 days (mean = 9) during 80 (see Fig. 4 for dates), Each court was attended throughout this period by only one individually marked bird. Unmarked court leaves were marked with the court number and date on their natural upper surface, using an oil-based felt-tipped pen. Pre- viously marked leaves, including those moved from one court to another, were re-dated and re-numbered. Numbers of leaves newly brought to the five courts (= decoration rate) were compared with numbers removed from them (= removal rate); the data are expressed as mean number of leaves per day. Leaf thefts were also monitored at two additional courts (sites 23, 34) during 79 and four additional courts (sites 34, 38, 47, 55) during 80. Leaf movement between a pair of courts was counted as one stealing event, distances between the courts being noted. MALE ATTENDANCE LEVELS, BEHAVIOUR AND VOCALISATIONS AT COURTS Males were mist-netted at or near courts banded with a meta! band and a unique two Colour band combination (= marked) and released im- mediately. Observations of marked males did not start until at least a full week after their release. We established cryptic canvas hides six metres from each court two weeks before starting obser- vations each season. Each observation lasted three hours, during 0600-0900, 0900-1200, 1200- 1500 or 1500-1800h, over peak activity. Male TOOTH-BILLED BOWERBIRDS attendance levels at courts were monitored during 7-28 November of peak activity and 8 December of post-peak activity during season 1979, and between 1 October-28 December of peak activity and 29 January-14 February of post-peak activity during season 80. The same eight court sites were monitored during both seasons, six of the eight males attending the same court throughout. The other two males were unmarked during 79 but were possibly the same birds we individually marked at their courts in 80, During seasons 79 and 80 male attendance was monitored simal- taneously al two courts, al one by direct observa- tion by DWF from a hide while tape-recording sound, and at another hy tape recording sound only. The microphone was placed at the perimeter of the court and camotflaged with leaf litter, DWF started each three hour direct observation ten minutes after being put in the hide by CBF, but earlier if the male retumed to his court and behaved normally. When a twelve hour cycle (i.e. (600-1800h) was completed at a pair of courts we repeated it, but then making the direct observa- tion at the court previously only tape recorded. Thus both calling and silent behaviours of all males were observed, as a control in order to enable us to imerpret audio-recordings made in our absence. A total of 96 and 220h of sound were audio-recorded at eight courts during peak ac- tivity of seasons 79 and 80 respectively, of which 48 and 110h respectively were accompanied by direct observations. A total of 6 and 18h of audio- recordings were made during post-peak activity of seasons 79 (during 1200-1] 500h) and 80 (0900- 1200h) respectively. During peak activity (17 October-22 Decem- ber) of season 89 four males were monitored at their courts by direct observation by DWF during three hour watches for a total of 119 hours. One site (35) was attended by the same individual that had attended it during 79 and 80, a different male attended site 33, and the other two siles were nol previously examined. Each male was radio- tagged with a 4 Biotrack single stage radio- trans- mitter (SS-1), using a tail mount, and was tadio-tracked by G. Moore using a Telonics (U.S.A.) Tr-2 receiver with a Yagi antenna (Cus- tom Electronics, U.S,A.). This enabled us to determine whether 2 silent male, unseen by the observer in the hide, was in fact present above his court, Male activities at courts were classed into ad- vertisement song, court maintenance, display, and silence, each being monitored, The number of court perches used for advertisement song by 125 the same four marked males (at sites 18, 28, 33 & 35; Table 3) during 79 and 80 were noted and time spent on each recorded during direct observation. Male vocalisations, including mimicry, per- formed at courts during 79 and 80 were analysed to determine whether they differed between in- dividuals, seasons or years. RESULTS COURT SEASONALITY AND MAINTENANCE The display season was started by males calling above uncleared or slightly cleared and decorated court sites, Initially only 1-5 leaves were placed on a site with little or no ground clearance. Place- ment of leaves and ground clearance sub- sequently increased simultaneously through pre-peak activity, Courts were well established by the third to fourth weeks of September during seasons 80, 83, 85, 87 and 88, and by the third to fourth week of October during seasons 78, 79, 81, §2, 84 and $9. Detailed results from display seasons 79, 80 and 81 are summansed in Figure 2. Differences between those seasons may be related to differen: ces in climate and phenology (Fig. 3) and these are disctissed below. Season 78, Males started calling on 27 Septem- ber and pre-peak activity started between 29 Sep- tember-5 October and lasted three weeks (Fig. 2). This period was dry (9mm of rain over five days of rain) and mean weekly temperatures were 20.9-21.8°C. Peak activity started 20-26 October with all courts well cleared and decorated with many leaves (mean = 73.1, SD =34.7) and faecal samples accumulating. This phase lasted ten weeks, until 28 December, during which 441mm of rain fell over 43 rain days and mean weekly temperatures were L8.0-27.8°C (mean of means = 22.97C). Post-peak activity lasted seven weeks, Courts deteriorated during heavy carly January rains but some faeces indicated males were sull attending, but were not maintaining, courts. During 19-25 January, the rain eased and courts were re-cleared, decorated and attended but this activity was brief and declined after a few sub- sequent days of increased rain. By 15 February all courts were derelict (Fig, 2). Season 79. On 12 September some males were calling above 10 known but uncleared traditional sites, Court establishment began during 15-21 Scptember and pre-peak activity lasted five weeks (Fig, 2), This period was dry (37mm of rain over nine rain days) and mean Weekly tempera- tures were 19,1-20,9°C for three weeks, rising tò 126 Post-peab Stean number aenm Stan D bama 4 (pT 4 > » Misna >|9Je ne ' uu reyes Seanon 79 Ween number Yn artendwnce | 9152215 AMT mt ARS SON i 9n E U U z + iwy 8 “a arendame Mean number m mn FNISENGIUDDIDIMWITHIUNDIZIHSO?OIDAI'**D 5 i] N b J ' FIG. 2. Court maintenance and attendance by male Tooth- billed Bowerbirds at courts during pre-peak, peak and post-peak activity of display seasons 78, 79 and 80.Note: Solid column = mean number of court leaves (pre- peak n = 5-25 courts, peak & post-peak n = 5 courts); Clear column — mean number of defaecaled seeds (pre-peak n = 5-10 courts,, peak & post-peak n = 10 courts); Line = % of time males attended courts (n = 8 males). 24.5°C, Peak activity started 20-26 October and lasted six weeks, until the end of November, whilst remaining extremely dry and hot (51mm of rain over 13 rain days) with mean weekly temperatures of 21.5-24.8°C (mean of means = 23,97C). Leaf numbers peaked (mean =75, SD = 31.0) between 3-9 November but had declined by the end of November. Male attendance levels remained high until 23 November. This suggests that while males were attending courts they were not maintaining them during these extremely dry conditions, There were significantly fewer leaves on courts when less rainfall preceded counts (rs = 0.97, P<0.05) and mean weekly temperatures were higher (rs = 0.92, P<0.05). Post-peak ac- MEMOIRS OF THE QUEENSLAND MUSEUM tivity lasted cight weeks. During 1-21 December it remained extremely dry (0.2mm of rain) and hot with mean weekly temperatures of 24.6- 28.3°C (mean of means = 26.2*C). Activity at courts rapidly declined and ceased by 21 Decem- ber. Rain started on 25 December and on 8 January, when rain eased, there was a bref penod of renewed court activity for two weeks, but courts subsequently became derelict with no fur- ther activity or calling during the extremely wet late January and February (Figs 2, 3B). Season 80, Pre-peak activity was carly, starting on 27 August when a Tooth-bill was calling above an uncleared traditional site. No traditional sites (n = 10) were cleared on 1 September but between 8-14 September a few were partly estab- lished and within two weeks, during which no rain fell and mean weckly temperatures were 21.3-24.4°C, courts were well established (Fig. 2). Peak activity started 22-28 September and lasted 14 weeks, to 28 December, during which 298 mm of rain fell over 20 rain days and mean weekly temperatures were 21.2-25.2°C (mean of means = 23.0°C). Leaf numbers reached a maxi- mum (mean = 119.4, SD = 50.1; Fig. 2) between 13-19 October. During the next three weeks the numbers of leaves on courts declined, although court attendance levels remained high, Leaf num- bers increased again during mid-December. There were more leaves on courts when male attendance levels were relatively lower and this correlation was significant (rs = 0.32, P<0,1). There were significantly fewer leaves when less rainfall preceded counts (rs = 0.72, P<0.01) and when temperatures were higher (rs = 0.54, P<0.05). Courts were still maintained and at- tended at the end of December, but all activity stopped when torrential January rains started (Fig. 3B). Post-peak activity lasted nine weeks. Courts became waterlogged, leaves rotted in situ, and no attendance or maintenance took place. Between 26 January-l February courts were re- cleared and decorated as rain cased (Figs 2, 3B). During February, when it again rained heavily (Fig. 3B), courts deteriorated, few leaves were added, and whilst some faeces were collected, court attendance levels were low. This indicated that males were seldom maintaining, or perching at, their courts, Non-display seasons, We heard Tooth-bills calling above traditional court sites during March and/or April of seasons 78, 79, 80,81 and 83, after wet Season rains eased. During March 1980 we found 15 traditional court sites to be uncleared, or slightly cleared, but decorated (<12 leaves). TOOTH-BILLED BOWERBIRDS Males then called above court sites with much mimicry. We identified one male as the marked traditional site owner but two birds, at sites usual- ly occupied by traditional marked males, were unbanded ‘intruders’, On 11 June, we heard a bird calling above an uncleared, undecorated, tradi- tional court site. Between 19-23 June we ex- amined ten traditional sites to find six uncleared or slightly cleared and decorated (< nine leaves) and males calling infrequently above them. All courts were derelict by 3 July. During five other winter seasons no similar activity was observed. The observed winter activity may have related to an abundant fruit crop at that time (Fig. 3C). COURT CHARACTERISTICS AND DECORATIONS Size und form. Of 59 courts attended in peak activity of season 89, 54 were well established and five partly so (see below). Of the 54 well established courts, three were almost circular and $1 were oval. The largest oval court measured 2.75 xX 1.95m (we have seen one 3.65 X 2.1m elsewhere) and the smallest 1.0 x 0.8m, Mean length of the longer oval court axis was 1.96m (SD =0.35, n= 51), and of the shorter 1.31m (5D = 0.34, n — 51). One extremely elongate court measured 2.64 x 0.75m. Large courts tended to have more leaves but there was no significant correlation between court size/leaf numbers (rs = 0.03, P>0.05) during peak activity of season 89, Of the 54 well established courts 50 had a tree rooted just within their cleared area. The ground 1o at least one side of this display tree was clean of floor litter and leaf decorations. The remaining four courts had two display trees, the ground around part of both being cleared. The display tree on most (90%) oval-shaped courts was lo- caled near one end of the longer axis. Display trees varied in size from a large sapling to a substantial tree (mean diameter at breast height = 113.6mm (SD = 1.4, range 30-340mm, n = 53). Twenty one of the 54 courts were flat, the rest being on sloping ground. The display tree on sloping courts was usually (9196) at the higher end. Twenty of the 54 courts had lawyer canes, Calamus spp., growing on them. A few to 10 saplings of <30mm diameter were prowing on 54% of courts. One or two heaps of germinating seeds beneath commonly-used singing perches were found on many courts. The five partly established courts were all small. One was a 1.4 x 0.75m oval and the others were circular, of e * Pa = p" 55 s a = -a . = A of a ia . oe 5 s . *. . ys Å .. y^ Ed . g 3 Minime Š iw ow SHONDPIEUAMI AAS ON DS EMIEMS TAS ANH ome, ie 1000 B dey eateitelt tet e E E rf = = x ASONDIJFMAMITASONDI FMAMJIASOND I 178-1981 an c Es & 3 io = ~ es = A ASONDIPMAMSIASONDIFMAMFIASOND) & 1978 - 1901 FIG, 3, Mean monthly iemperature (a), mean monthly rainfall (1978-1990) (b), and monthly percentage of sampled trees in fruit (c) on the Paluma Range, north Queensland. Leaf numbers and types. The mean number of leaves on courts during peak activity of seasons 79, 80, 81, 86 and 89 varied from 60-78 (mean of means = 69, Table 1). There were more leaves on courts during 79, 80 and 81 than during 86 and 89 but differences were not significant (x° = 4.45, P>0.30), Variations between seasons were pos- sibly duc to the dates counts were made during peak activity (see Table 1). The most abundant decoration leaf species (mean = 92%, Table 1) was Polyseius australiana, Araliaceae (= Polyseias in the fol- lowing). Other leaves often used as decorauon included Sloanea langii (Elaeocarpaceae), Cryp- tocarya hypospodia, C. mackinnoniana, C. putida, Neolitsea dealbata, Litsea bindoniana (Lauraceae), Symplocos cochinchinensis (Symplocaceae), Alphironia whitei (Rham- naceae), Commersonia bariramea (Ster- 128 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1. Mean number of leaves and percentage of them thal were Polyscias on male Tooth-billed Bowerbird courts during peak activity of five display seasons. (Months are indicated by their first capital letter). Seasons 78 40 81 B6 89 Courts and leaves " (200-9N) _(228-260) (270-9N) _ (15-210) (47-30N)_ Number of well established courts 21 36 17 46 63 Mean number of leaves a court 70.0 75.2 78.2 59.5 594 Range 20-120 198-179 44-150 19-108 20-120 so 28.3 34.2 25.9 22.3 228 % of total leaves that were Polyscias 97 98 80 91 B3 Number of courts with > 8096 20 34 12 34 21 70-90% 1 2 4 7 20 < 70% Polyscias 0 o 1 5 2 Number of new court sites 3 6 1 5 7 Number of partly established cours 1 2 2 2 5 Mean number of leaves a court 16.5 245 31.0 19.8 “yp of total leaves that were Polyscias a 73 53 52 Number of new court sites 1 2 2 2 g Court sites (n = B) with one owner Mean number of leaves a court 78.6 92.8 792 61.0 455 % of total leaves that were Polyscias 98 97 88 B4 73 Courts sites with 2 (n = 3) or 3 (n = 4) Owners Mean number of leaves a court 75.0 69.4 81.0 59.4 659 Polyscias 99 98 ‘98 98.0 92.0 % of total leaves that were culiaceae), Smilax australis, 8. glyciphylla (Smilacaceae) and Rubus moluccanus (Rosaceae). Leaves of Asplenium and Alpinia spp. were used seldom. The percentage of Polyscias leaves on well established courts during 79 and 80 was higher than during other seasons, but these differences were not significant (x? = 1.62, P>0.80). A lower proportion (83%) of Polyscias were used as decoration during 89, the remaining proportion consisting mostly (12%) of Cryptocarya spp.. During seasons 78 and 79, 95% of courts were decorated with >90% Polyscias. During seasons 81, 86 and 89, 85% of courts were decorated with >70% Polyscias. Seventeen courts over these five seasons had <70% Polyscias (range = 30-65%), and another had only 10%. Of 21 newly estab- lished sites over these five seasons, five were decorated with 70-90% and three had <70% Polyscias (Table 1). The numbers of leaves on traditional courts attended by the same male during the five seasons averaged 71.6 (SD = 18.8, n = 6) and was similar (mean = 70.1, SD = 8.3, n = 7) to the numbers on traditional courts attended by two or three males over the period (Table 1). These courts were all mostly decorated with Polyscias, Twelve courts were only ever partly estab- lished, during the peak activity of the five seasons. These were decorated with fewer leaves and a smaller proportion of Polyscias (Table 1). Ten of the 12 were at new court sites and the other two (season 89) were on traditional sites of nine and 11 seasons standing. The latter site had been attended by the same individual male during the previous 11 seasons, but during 89 was attended by an unmarked male who cleared three small separate areas < 1m apart and decorated each with a few leaves. LEAF TURNOVER RATES Season 79. The total number of leaves on ad- jacent courts (sites 6, 32, 33, 35 and 37; sec Fig. 1 and 5) were counted four times over 21 days (mean sampling interval = 7 days) during peak activity and 871 newly-placed leaves were marked on them. Of these leaves, 79% were removed from courts within 7 days, 18% (1% of which had been moved from one court to another) between 7-13 days and 3% between 14-20 days. The mean number of days an initially fresh leaf remained on a court was 6.1. There was an average of 59 leaves on each court during peak activity, courts 35 and 37 having most (Table 2). Mean numbers of leaves brought TOOTH-BILLED BOWERBIRDS 129 TABLE 2. Court decoration and leaf turnover rates by the same five male Tooth-billed Bowerbirds during peak activity of display seasons 79 and 80. — Gourt reference number Season Decoration and removal rates * 6 32 33 35 _37__ Mean of means 79 Mean number of total leaves 135 685 46.0 87.8 79.5 59.4 Mean number of new leaves a day 1.1 68 3.8 99 a 59 Mean number of leaves removed a day 24 7.0 62 112 S5 7.3 B0 Mean number of total leaves 63.3 62.1 66.8 103.6 10682 84.5 Mean number of new leaves a day 63 67 5.6 7.8 TT 6.8 Mean number of leaves removed a day 64 58 52 74 77 64 % of Polyscias leaves 91 100 ga 99 85 95 *= Mean numbers are for over 21 days in 78 and 97 days in 60, lo the five courts per day during peak activity varied considerably (range 1.1-9.9, SE=1.57), as did the daily removal of old leaves (range 2.1- 11.2, SE = 1.58). Daily decoration and removal tates were significantly correlated during peak activity (r = 1.33, P<0.001). Decoration and removal rates on the five courts declined as peak activity progressed, mean num- bers of leaves placed on courts declining from 51 to 24 and leaves removed from 66 to 30 (Fig. 4). Peak activity ended at the end of November. During the first two weeks of December (= post- peak activity) few new leaves were placed on courts and none were removed. Decline in decoration placement and removal rates this season were associated with extremely dry con- ditions. Only one of the five sample courts was, slightly, re-cleared and decorated during a brief period of renewed post-peak activity following January rains. Season 80, The total number of leaves on the five adjacent courts used in season 79 was counted twelve times over 97 days (mean sam- pling interval = 9 days) during peak activity and 3219 newly-placed leaves were marked on them. Of these 53% were removed from courts within 9 days, 42.9% (2.5% of which were moved from one court to another) were removed between 9-17 days, 4% (0.3% of which were moved from one court to another) between 18-26 days and 0.1% between 27-34 days. The mean number of days an initially fresh leaf remained on a court was 5.9. There was an average of 85 leaves on each court during peak activity, courts 35 and 37 having the most (Table 2). Mean numbers of leaves brought to a court per day throughout peak activity by each of the five males were similar (range 5.6-7.7, SE = 0.42), as were the daily removal of old leaves (range 5.2-7.7, SE = 0.48). Daily decora- tion and removal rates were significantly corre- lated during peak activity (r = 0.96, P<0,01). Decoration and removal rates on the five courts varied considerably during peak activity. The mean number of new leaves on courts at each Inspection ranged from 49-85 and of leaves removed from 24-91, Decoration and removal rates were highest during the early peak activity. Males brought more leaves to courts per day during the first seven weeks of peak activity (mean = 7.8, SE = 0.57) than during the later seven weeks (mean = 5.9, SE = 0.4), and the difference was significant (Mann Whitney U=1, - = 3 Peak | Post-yreak az 2 + = a Remnwel ruter 8 au < 2 Decaratinn raies -y ` Z a . = Season 79 . E = SN 12N 1N 26N 3p wo Dates of counts Ei = a S © í a S wv = 22S 293 60 150 240 310 WN 1TN 25N 6D 16D 28D Dates of count war I1F FIG. 4. Mean number of leaf decorations brought to (= decoration rate) or removed from (=removal rate) courts by the same five male Tooth-billed Bowerbirds during display seasons 79 and 80. Months are indicated by their capital letter. Season 79 Season 80 inom FIG, 5. Inter-court leaf theft by six (season 79) and eight (season 80) adjacent male Tooth-billed Bowerbirds on Tooth-bill Hill and slopes. Note: Large-type number by court site (+) = site number; Arrows indicate direc- tion of leaf theft with total number of leaves involved indicated by smaller-type number beside the line. p=0.008). Similarly, more leaves were removed per day during the first half (mean = 7.4, SE = 0.67) than during the latter half (mean = 5.4, SE = 0.38) of peak activity, the difference being significant (Mann Whitney U = 1, p = 0.008), Immediately after January rains eased males removed all old leaves from courts and placed new ones on them (Fig. 4). More Polyscias leaves (mean = 98%, range 93-100%) were used on these courts during the first half of peak activity than during the latter half of it (mean = 90%, range 72-93%). Fewer (mean of means = 61%) Polyscias leaves were used as decoration during post-peak activity, LEAF THEFT Season 79, Of 871 leaves marked on six ad- jacent courts during peak activity, ten (19b) were subsequently found on neighbouring courts, All were stolen within 7 days of being marked. Three of the six male court owners were thieves, males MEMOIRS OF THE QUEENSLAND MUSEUM at courts 33 and 37 cach stealing one leaf, and the male at court 35 stealing a total of eight leaves from three neighbouring courts (Fig. 5), Daily theft rate was 0,38 leaves/day/male. Season 80. Of 3219 leaves marked on nine adjacent courts during peak activity, 93 (3%) were subsequently found on neighbouring courts, Of stolen leaves 88% were taken within a mean of 9 days of being marked and the remainder were moved >9 days after they had been marked on the victim's court. More leaves were stolen during the first half (63% of thefts) than the latter half of peak activity. Three of the nine male court owners were thieves, these being the same individuals found to be thieves in season 79 (Fig. 5). Daily theft rate was 0,88 leaves/day/male. Most (97%) leaves stolen were Polyscias. The male at court 33 stole six leaves from one neighbour, and the male at court 37 stole four leaves from three neighbours (Fig. 5). The male al court 35 was never stolen from but stole a total of 85 lcaves from seven neighbouring courts (mean inter-court distance = 60.3, SD = 21.5m, range 36-95m). There was a significant correla- tion between number of thefts by the court 35 male and the distance he travelled to a victim's court (rs = 0.97, P<0.01). He stole more leaves from closer neighbours (Fig. 5) but did not steal from another neighbour, at court 38, 105m dis- tant. Court 34, from which a single leaf was stolen, was excluded from this analysis as it was not established until the last two weeks of peak activity. Neighbouring males usually stole leaves during a court owner's absence. On four oc- casions, however, we watched the bird from site 35 remove a leaf from site 33 whilst the owner continued singing above it. MALE ATTENDANCE LEVELS AT COURTS Seasonal variation, During peak activity of seasons 79, 80 and 89 males spent an average of 52, 79 and 61% respectively of time attending courts. The proportion of total time individual males spent at courts during peak activity (Table 3) varied significantly during season 79 (x? = 26.5, P<0.001) but not during 80 (X? = 2.25, P>0. 90) or 89 YN = 2.81, P>0,30). Males spent an average of 25 (range 18-31, SE = 1.58), 22 (range 17-27, SE=1.18) and 2] (range 20-25, SE = 1,03) mins at the court per presence during seasons 79, 80 and 89 respectively (Table 3). Thus, despite lower attendance levels during season 79 and, to lesser extent, during 89, once males were at their courts the duration of each visit was similar. Mean duration per absence from TOOTH-BILLED BOWERBIRDS 131 TABLE 3. Court attendance levels by individual male Tooth-billed Bowerbirds during peak activity of display seasons 79, 80 and 89. HOURS ABSENCE PRESENCE Season Site Audio- Observations Mean mins % of total Mean mins % of total number recordings _(2 absence _ time absent @ presence time present 79 18 * 12 12 23.5 55.5 18.9 44.5 28 * 12 12 15.0 35.4 25.8 64.6 33 * 12 12 324 67.6 1B.0 32.4 35 * 12 12 23.5 45.6 28.0 54.4 6* 12 37.4 624 24.6 37.6 17 12 16.0 377 28.0 623 21 12 13.0 27.2 30.9 72.8 32* 12 24,0 53.1 24.1 47.0 Total / Mean /% 96 48 225 48.0 24.9 52.0 ao 18* 27 12 8.2 22.6 25.6 77.4 28 * 26 17 5.1 16.0 23.8 64.0 cr Ad 30 18 76 25.7 20.6 74.3 35 * 27 21 53 21A 20.0 78.9 6^ 27 6 7.9 29.6 17.0 70.4 17 27 15 4.9 17.9 21.8 82.1 21 26 9 53 15.1 27.1 64.9 32 * 30 12 47 17.1 19.9 82.9 Total / Mean / % 220 110 6.1 20.7 21.6 79.3 B9 33 23 23 16.9 44.1 20.3 54.3 (1.8) ** 35 * 24 24 17.7 39.2 24.7 58.4 (2.4) 23 36 36 13.6 37.9 20.4 59.4 (2.7) 37 36 36 8.4 27.0 21.6 71.9 (1.1) Total / Mean / % 119 119 13.1 36.0 21.5 62.0 (2.0) * = same male owner each season ** in parenthesis = % total time males were present but detected only by radio-tracking. courts during season 80 was much briefer and far less variable between individual males (range 5-8mins, SE = 0.53) than during 79 (range 13- 37mins, SE = 3.02) or 89 (range 8-18mins, SE = 2.11). During season 89 transmitters on the four radio- tagged males enabled us to learn that they spent 1.1-2.7% (mean = 2) of time silently in the forest canopy above their court, this figure being similar during different months of peak activity, and times of day. Thus, figures for absence of other males, lacking radio-tags, during seasons 79-80 (Tables 3-5) could include an error of up to 3% and figures for time present could be higher by up to 3%. This potential error was not taken into account in time-budgeted court activity data presented for season 89 (Tables 6-8). Monthly variation. Peak activity lasted six weeks (20 October-30 November) during season 79, 14 weeks (22 September-28 December) during 80 (Fig. 2), and 11 weeks (13 October-28 December) during 89. Season 79 was excessively dry and, although December rainfall was near average, it did not start raining until 25 Decem- 741 MW Kt mm Monthly rainfall mm 2 FIG, 6. Monthly rainfall during display seasons 79, 80 and 89 and mean monthly (= W) for 1978-1990 on the Paluma Range, north Queensland. 132 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 4. Monthly court attendance levels by male Tooth-billed Bowerbirds part of the day (Table 5). during peak (seasons 79, 80 & 89) and post-peak (79 and 80) activity.* During season 80 court at- tendance levels remained ABSENCE PRESENCE A high throughout the day. Season Months Hoursef Mean mins %oftotal Mean mins % oftotal Mean duration per presence audio- absence timeabsent @ presence time present at courts did not v ary great- _____ Fecordihgs 3 : i - ly at differing times of the 79 day overeach season (Table Paak November s6 225 48.0 24.9" 52.0 5) Post-peak December 6 100.0 8 i MALE BEHAVIOUR AT 80 Peak October 78 64 23.0 193 770 Gas i November al 49 16.8 222 83.4 vertisement song. December 61 75 23.2 245 76.8 Male Tooth-bills spent Post-peak February 18 274 66.0 153 34.0 most (mean of means = rT) — 95.7%) of their time at Peak October 47 13.4 344 236 638(23)** courts giving loud adver- November 35 97 286 228 69.7 (17) tisement song from December 26 16.3 460 178 52.0 (1.9) *= court sites and sample sizes as in Table 3 ** in parenthesis = % total time males were present but detected only by radio-tracking ber. Little or no rain fell during September 80 or 89, and October rainfall of those seasons was below average, November 79 was dry, and November 89 wet (Fig. 6). Court attendance levels were higher during each month of season 80 than during each cor- responding month of 79 and 89. Attendance levels reached a peak in November of seasons 80 and 89. Court attendance levels were lowest during November 79 and December 89. Mean durauon per presence at courts varied little between the respective months of favoured perches. This was similar for all males (range 90-98%) during each season, each month of peak activity and different umes of day (Tables 6-8). Mean duration of advertisment song periods was similar during seasons 79 (range 19-30mins, SE = 1.46), 80 (range 17-27mins, SE = 1.18) and 89 (range 20-27mins, SE = 1.31, Table 6). Mean duration of song periods was similar each month of peak activity except during December 89 when seasonal activity declined. It was also much lower during post-peak activity of season 80 (Table 7). TABLE 5. Variation in daytime court attendance levels by male Tooth-billed Bowerbirds dunng peak activily of display seasons 79, 80 and 89. * each season, Mean duration ABSENCE PRESENCE per AU iin courts waj Season Time Hours Mean mins 9& of total Mean mins — 9b of total notably low during a — ru _ @ absence time absent @ presence time present months of the 80 season. During the extemely dy 7» peemeo 3s geo m early December 79 court at- 1200-1500 24 24.8 455 271 545 tendance stopped. During 1500-1800 24 39.2 626 28.4 374 season 80 there was a brief A remt To postpéak ACTIVITY s0 06000900 48 a4 182 20,6 338 when torrential January 0900-1200 60 7.0 232 216 768 rains stopped but atten- 1200-1500 50 6T 227 20.6 713 dance levels were low, 1500-1800 52 61 19,7 23,9 80.3 males spending little time at - e- courts each visit (Table 4). s9 0600-0900 29 82 274 19.5 70718) z neii 0900-1200 30 16.1 40.1 21.8 58.7 (1.8) Daytime variation. Court 2200-1806 30 sh Fry 20g 55.5 (2.8) attendance levels were al- 1500-1800 30 14.1 34.5 25.6 63.9 (1.6) ways highest during the period 0600-0900h. During season 79 attendance declined during the latter *= court sites and sample sizes as in Table 3 * |n parenthesis = % total time males were present but detected only by radio-tracking. TOOTH-BILLED BOWERBIRDS TABLE 6. Time-budgeted activities performed by individial male Tooth-billed Bowerbirds at courts during peak activity of display seasons 79, 80 and 89. * ADVERTISEMENT SONG MAINTENANCE DISPLAY SILENCE TOTAL TIME PRESENT Season Court site Mean mins Sboftine Meanmine € "of time Meanmms %of time Mean mins 9b of time (mins) number song present maintenance present @ display present @ silence present period singing penod maintaining penod displaying period spent silent T3 t+ 184 969 1.3 18 14 os 10 3 324 2a 253 984 0,8 0.8 19 04 10 04 465 nu 18.0 89.6 20 17 18 39 37 4a 233 35 25.5 547 23 12 oo 0.0 20 41 392 6, 4 4 4.0 04 11 1.8 13 18 274 7 292 97.6 28 13 1.6 07 os U4 449 21 304 eB4 14 05 16 og 0.5 0.2 525 32° 23.8 96.4 14 ae 09 00 os 08 334 Total / Mean / % 25.2 6.7 13 1o 15 [AJ “7 14 2993 50 15" 251 882 17 oT 4 0.3 De 08 1253 28° 24.1 974 13 oB 14 10 D.B og 1311 33° 237 odo 15 21 a7 1.8 43 20 +338 “ue 12.8 828 20 27 18 07 12 28 1278 6* 16.8 84 1 15 30 *3 12 D8 17 4141 17 231 95.6 17 24 15 04 1.0 T.6 1330 2 BT 287 0.8 D4 15 D2 0.3 D7 1325 iss 351 24 24 20 os 10 10 1495 Total / Mean / % ao 95.5 1.6 48 16 08 41 18 10469 a Š. o “ “ - 3 204 926 10 12 20 08 V6 54 750 35 * w6 948 os 0.6 22 05 22 40 640 2a ms 939 10 08 1.6 02 1.8 51 1282 7 238 954 12 11 16 04 13 31 1552 Total / Mean / % 23.2 944 12 oa 1.8 04 16 43 4424 "= game mele owner each season = = silent. periods of > 30 seconds duration (see Results), but excluding maintenance time, Mean duration of song periods did not vary great- ly during different times of the day over each season (Table 8). Almost all (>99%) advertisement song was given from perches immediately above or about courts, males only rarely (<1%) singing loudly on the ground at the base of the display tree, prior to the distinctly different display subsong. Males usually sang from more than one perch during a court visit (mean = 1.9), spending an average of 13mins on each (Table 9). Song perches were high above the court (high perches), adjacent to it (adjacent perches) or just above the court (court perches). High perches were >5m above the ground in the understorey or lower canopy, and up to 8m horizontally from the court location. Most males had two high perches (mean = 1.9). Adjacent court perches were mostly horizontal or sloping sapling or small tree branches, 3-5m above ground and between 1-5m from the court perimeter. Males used 1-4 adjacent perches (mean = 1.8). Court perches were horizontal branches of saplings and, less com- monly, vines or fallen branches lying across the court at 0.3-3.0m (mean = 1,4) above the cleared ground or within a metre radius of its perimeter. More court perches were used during season 80 (mean = 8.3, range 5-12) than during 79 (mean = 4.8, range 4-7). Males spent an average of 21, 1] and 68% of total song time singing on high, adjacent and court perches respectively, spending more time singing on high and adjacent perches during season 79 than 80 (Table 9). High perch singing decreased as season 80 progressed, from 22% of October song time to 13% of November and 4% of December song time. High perches were most- ly used (90% of occasions frequented) for singing when males firstreturned to their court area, birds subseguently flying down to their adjacent and/or court perches. Returning males someumes flew directly to an adjacent perch and sang there before moving to a court perch or returned direct- ly to a court perch (Table 9). Of returns to a court perch 37% were, however, via the court to place a newly-aguired leaf there. During 0600-0900, 0900-1200, 1200-1500 and 1500-1800h males sang from high perches for 12, 19, 19 and 8% of total song time respectively. Thus, males spent more time singing on high perches during the hotter, brighter, part of the day. Of 31 perches used by four males at court sites 18, 28, 33 and 35 during season 79, 84% were still 134 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 7, Monthly time-budgeted activities performed by male Tooth-billed Bowerbirds at courts during peak (seasons 79, 80 and 89) and post peak (80) activity. * ADVERTISEMENT SONG MAINTENANCE DISPLAY SILENCE™ | TOTAL TIME PRESENT Season Months Mean mins % of tima Mean mins @ % of time Mean mins of lime Mean means % of time {mins) @ song present mainteranee prasant @ display present @ silence present pernod _ singing period maintaining period displaying penod spent silent 79 Peak November Bz m7 3 15 1.5 ps iT 14 z993 80 Peak Octobar mz M4 15 22 ts 02 i4 at 554 November 220 62 12 13 rs 11 12 4 4055 December ms M2 14 2c Tè 12 10 2e 2811 Posi-paak February 138 318 oa 16 ts ac 2e Ki 89 Peak October 25.7 34 13 15 MU oe 3 36 Th November 2&7 356 40 06 17 04 20 34 1808 December 17A 323 44 D5 23 1 37 65 Ha * court sites and sample sizes as ìn Table 3 = = silent periods af > 30 seconds duration (sêe Results), bul excluding maintenance time- în use by the same males during 80. Of six perches used by the site 35 male during 79, all were in use during 80 and he was still using three of them during 89. Maintenance. Male Tooth-bills spent a mean of only 1.2% of total time actually on courts and maintaining them. This figure was similar for all males each season, during different months of peak activity and different times of day (Tables 6-8). Mean length of court maintenance periods varied between males but was never >2.8mins (Table 6). Of 104 observed court maintenance periods, 95 immediately followed a male's arrival with a leaf and nine occurred during or after singing or dis- play and involved court clearing only. After plac- ing a new leaf and/or performing court maintenence males hopped up to a perch to sing (72% of arrivals) or simply flew away from the court. Number of new leaf decorations brought to the court in each three hour period was higher during season 80 (mean = 1.3) than 79 (mean = 0.8) or 89 (mean — 0,7). More leaves (60%) were brought to courts during 0600-1200hrs than during the rest of the day. Once a male had placed a new leaf on his court he usually tidied or re-arranged his fresher leaves and removed old or fallen ones to beyond the court perimeter. Males were usually silent when maintaining courts, only occasionally giving a single ‘chuck’ note. Displays. Upon firstnoticing a conspecific near his court 4 male would drop to the base of his display tree to give quieter, more intense, adver- tisement song or, more often, would immediately commence display 'subsong” vocalisations. On a few occasions a male adopted a static 'frozen' sleeked posture prior to subsong and display (see below), Display vocalisations and posturing are described elsewhere (Frith & Frith, 1993). Most times (83%) males performed only one display, but on a few occasions they performed a sequence of two to four immediately repeated ones. During peak activity males spent <1% of their tolal time al courts displaying, the mean duration of a display (including subsong) ranging from 1,1-2,2mins (mean = 1.6, Table 6). Number of displays per three hour period was greater during the peak activity of seasons 80 (mean = (0.8) than during 79 (mean = 0.5) and 89 (mean = 0.3). The total ime males spent displaying was greatest during the peak activity month of November (1.5 displays/display period) and December (0.9 dis- plays per three hours, Table 7). Most displays were performed during 0600-0900h of seasons 79 and 80 and least during 1500-1800h (Table 8). Males continued to display during post-peak (court maintenance) activity (Table 7), in fact spending relatively more time displaying then. We could not sex birds seen to be displayed to by their appearance or behaviour. No mating was seen. Stlence. The proportion of total court atten- dance time males spent silently (= >30secs without calling), other than during court main- tenance periods, was greater during season 89 (4.3%) than 79 (1.4%) or 80 (1.6%, Table 6). These higher 89 figures were reflected during each month of peak activity (Table 7) and dif- ferent times of day (Table 8), notwithstanding up TOOTH-BILLED BOWERBIRDS 135 TABLE 8. Variation in daytime-budgeted court activities by male Tooth-billed Bowerbirds during peak activity of display seasons 79, 80 and 80. * ADVERTISEMENT SONG MAINTENANCE. DISPLAY SILENCE ™ TOTAL TIME PRESENT Season Time Mean mins- %of time Meanmins@ "eof time Meanmins %of time Mean mins % of time (mins) @ song present maintenance present @ display present @ silence present period singing period maintaining period displaying penod spent silant 73 0600-0900 254 97.4 10 14 16 14 14 os S26 0500-1200 21.5 96.9 15 20 20 10 1.2 a7 741 1200-1500 27.9 85.9 1.0 04 1.5 10 24 30 786 1500-1800 27.6 973 18 10 T1 02 16 15 539 80 0600-0800 203 95.5 13 18 14 12 12 14 2414 0900-1200 228 96.3 14 14 11 10 11 13 7785 1200-1500 21.2 850 18 19 17 70 1.0 21 2785 1500-1800 24.1 %6,1 19 21 1.8 03 4.0 45 2506 89 0500-0800 24.7 25.4 1.0 11 14 o4 11 34 1229 0900-1200 26.4 MB 12 oe oo oo 16 46 1045 1200-1500 206 $46 15 10 1.8 08 20 34 B88 1500-1800 261 $2.9 13 10 1.8 0a 20 53 1150 * court sites and sample sizes as in Table 3 = = silent periods of > 30 seconds duration (see Results), but excluding maintenance time. to an additional 3% silence of potential, but un- detected, presence (see Table 3). The possibility that the movements of two people radio-tracking in the area might have caused relatively longer silent periods during season 80 is considered un- likely but cannot be dismissed. There were more silent periods over 1200-1500h during 79 and 80 than at other times of day (Table 8). Most (92%) silent periods occurred immedi- ately before, or during, advertisement song when birds preened, changed perches, turned on their perch to face the opposite direction, or when they stopped singing to apparently listen to neighbours' calls. Having apparently noted a neighbouring male was not calling, a court owner flew immediately in the direction of the presumably unattended court to steal a leaf. The other 8% of silent periods occurred when males adopted a sleeked ‘frozen’ posture. We recorded thc frozen posture 34 times (mean dura- tion = 1.1mins), 28 of which were performed in response to a conspecific flying toward the court. On nine of these latter occasions the male imme- diately followed the frozen posture with display. On the 19 other occasions he chased the con- specific, which we assume to have been a neigh- bouring male (sometimes confirmed by band colours), Court owners sometimes chased off potential thieves without first adopting a frozen posture, the latter usually being performed by birds apparently initially unaware of the identity or sex of the visitor, Four frozen postures oc- curred when we heard, but could not see, a bird land high above the Tooth-bill at its court and two were given when a Grey-headed Robin Poecilodryas albispecularis, flew directly to a court. This behaviour appears typical of male Tooth-bills surprised by the appearance or sound TABLE 9. Perch type used for advertisement song by the same four male Tooth-billed Bowerbirds at courts during peak activity of display seasons 79 and 80, “ HIGH PERCHES (> 5 m} ADJACENT PERCHES (3-5 m) 'COURTPERCHES(<3m) GROUND TOTAL SONG TIME* Season Mean % of total % of times Mean % of total 96 of times Mean %oftotal %oftimes % of total (mins) mins @ song used on mins @ song used on mins @ song used on song __ perch time arrival perch timè arrival perch time arrival __time 79 112 26 s0 21.5 14 56 147 59.8 49 01 1349 20 8.9 16 Bg 114 B 50 10.5 75.4 37 06 2878 * = direct observation watches only at court sites 1B,28,33,&35, see Table 2. MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 10. Analysis of advertisement song given by male Tooth-billed Bowerbirds at eight courts during peak activity of display seasons 79 and 80.* Court sites as in Table 3. SQUEAL CHUCK CHUCK+ BABBLE CHUCK + MIMICRY EABBLE BABBLE + MIMICRY MIMICRY TOTAL SONG TIME Semn Mean % of total Maan % of total Mear % cf tobe Monn % of total Mest 3h nf total Mean 4 of total Mean % of total (mins) mine @ song ne @! song mine @ tonga = min» Pb wong tmins 0 song mins @ tong mins @ song time phase fire phase time phate____ldlme___ phass tna phase ima __ phase tma o n 23 17 26 5a 22 i? 1 at 9 234 "n7 555 51 74 2788 (al 4 7 13 ag 20 33 22 29 TT 357 BD 40 42 a9 = of an incoming bird as we often saw Grey-headed Robins, Chowchillas Orthonyx spaldingii, East- em Whipbirds Psophodes olivaceus, Yellow- throated Scrub-wrens Sericornis citreogularis, Brush-turkeys Alectura lathami and Spotted Cat- birds Ailuroedus melanotis foraging toward a court without concerning the Tooth-bill owner. We did not witness a single aggressive en- counter between Tooth-bills at a court other than an owning bird chasing off another. On a few occasions we did note two birds confronting each other in a food (fruiting) tree by facing each other with sleeked plumage and agitated vocalisations. MALE VOCALISATIONS AT COURTS Advertisement song content was similar in seasons 79 and 80. We could divide song into seven component calls: ‘squeal’, ‘chuck’, “chuck + babble’, ‘chuck + mimicry’, ‘babble’, ‘babble + mimicry’, and exclusively pure ‘mimicry’, changes from one to another usually being quite obvious. Calls were given in phases, a male per- forming a phase of ‘chuck’ calls, following this with a phase of “chuck + mimicry’ and so on. Each song period (= a court visit) consisted of several to many phases. Each phase lasted an average of 8 and 5mins during seasons 78 and 80 respectively, The relative time spent giving each call component and mean length of cach phase was similar each season (Table 10), 'Sgueals' were sgueal-like screctchy notes. Phases of ‘squeals’ represented 1,7% of total song time and were brief (Table 10). “Squeals' were mostly (64%) given first by returning birds or as second phase calls (19%) that usually fol- lowed ‘chucks’ (Table 11). Occasionally ‘squeals’ were again given, briefly, when a male changed perches. *Chucks’ consisted of a repeated ‘chuck’ note, represented 6% of total song time, and were brief (Table10). ‘Chucks’ were mostly (72%) given first by returning birds (Table 11), but also fol- lowed ‘squeals’. Males sometimes briefly reverted to ‘chucks’ after changing perches. “Chuck + babble’ and ‘chuck + mimicry’ were dominated by ‘chuck’ notes interspersed with occasional ‘babble or mimicry’, These calls rep- resented < 4% of total song time and phase dura- tion was brief (Table 10). Males sometimes gave these calls having just retumed to their court instead of a ‘chuck’ call, but mostly they were given as second phase calls (Tablel 1) following ‘squeals’ and/or ‘chucks’, ‘Babble’ and “babble + mimicry’, often includ- ing occasional ‘chucks’, were the most common calls and represented 30 and 49% of tolal song time respectively with long phase duration (see Table 10). Males sometimes first gave these calls upon returning to the court, but mostly they were given as second, third or fourth phase calls (Table 11) following ‘chuck’, ‘chuck + babble’ and/or ‘chuck + mimicry’. Some males continuously alternated between ‘babble, babble + mimicry’ and/or ‘mimicry’. Males gave more intense and quieter advertisement song, of ‘babble’ or ‘bab- ble + mimicry’. immediately prior to display of a mean duration of 20.9mins (range 0-65). Exclusive mimicry was performed by all males, represented a mean of 8% of total song time, and each phase lasted an average of 4.7 mins (Table 10), A far larger proportion (60%) of song time involved mimicry, however, when ‘mimicry’, ‘chuck + mimicry” and ‘babble + mimicry’ were combined. Mimicry usually fol- lowed ‘chuck + mimicry” or “babble + mimicry’. Cicadas. crickets, frogs. dripping water and 27 bird species were identified as models for mimicry. We audio-recorded 1074mins of exclusive mimicry during advertisement song over seasons 79 and 80, Of total *mimicry' time, 43% was of Bower's Shnke-thrush Colluricincla boweri and 30% of medleys of various bird calls mostly dominated by that species. Mimicry of Bower's Shnke-thrush also dominated ‘chuck + mimicry’ and ‘babble + mimicry’. King Parrots Alisterus scapularis were also commonly mimicked (11% of *mimicry'). Other bird species mimicked for a combined total of TOOTH-BILLED BOWERBIRDS 137 TABLE I 1, Phase sequence of each call component òf advertisement song given by male Tooth-billed Bowerbirds al eight courts during peak activity of display seasons 79 and 80, * Data are expressed as % of total number of phases of each call component.** Phase sequence Call component i — 2 4 5 8 7 8 9 10 11-18 Squeal 64.0 194 56 5.6 og *.B 0.8 0.8 oe 0.0 0.0 Chuck 724 71 67 43 2e 26 1.0 1.5 0.2 05 1.0 Chuck + babble 17.1 442 122 o4 55 33 1.1 0.6 37 0.6 23 Chuck + mimicry 12.4 34 147 7.6 53 40 24 3.0 0.6 12 24 Babble 13 217 69 173 125 6&7 4.8 3.5 1.0 12 31 Babble + mimicry 93 183 216 17.3 107 7.9 58 2.6 +3 18 28 Mimicry 77 198 212 184 10,5 65 4.1 2.8 2.8 12 20 Display subsong 83 33 150 234 150 83 &3 16.0 5:0 0.0 34 —— AO <íY'N— — —u———b>»b———— *= court sites as in Table 3 ** = data for seasons 79 & 80 combined. See Table 10 for total song time 13% of “mimicry” time were: Fan-tailed Cuckoo Cuculus flabelliformis, Yellow-eyed Cuckoo- shrike Coracina lineata, Grey-headed Robin, Golden Whistler Pachycephala pectoralis, Chowchilla, Eastern Whipbird, Yellow-throated Scrub-wren, Little Treecreeper Climacteris minor, Bridled Honeyeater Lichenostomus frenatus and Golden Bowerbird Prionodura new- toniana. Bird species mimicked less frequently (combined total = 3% of 'mimiery') included; Pale-yellow Robin 7regellasia capito, Black- faced Monarch Monarcha melanopsis., Spec- tacled Monarch M, trivirgatus, Rufous Fantail Rhipidura rufifrons, Grey Fantail R. fuliginosa, Australian Fernwren Crateroscelis guiterlats, Large-billed Scrub-wren S, magnirostris, Brown Gerygone Gerygone mouki and Mountain Thornbill Acanthiza katherina. 'Tooth-bills mimicked several distinct calls of a given bird species. For example, they commonly mimicked both the whistled single note song and the ‘chee-chee’ greeting calls of Grey-headed Robins, both whip-crack song and the ‘chip- chop’ calls of Eastem Whipbirds and both squeals and rattle calls of Golden Bowerbirds. Mimicry was sometimes given opportunistically, in imme- diate response to calling birds, such as those of a passing flock ef Crimson Rosellas Platycercus elegans, lorikeeis Trichoglossus spp. or, less fre- quently, Sulphur-crested Cockatoos Cacatua gelerita or Red-tailed Black-Cockatoos Calyp- torhynchus magnificus. Tooth-bill$ sometimes “duetted” with other species including King Parrot, Grey-headed Robin, Golden Whistler, Bower's Shrike-thrush, Eastern Whiphird, Yellow-throated Scrub-wren, Little Treecreeper and Bridled Honeyeater. Neighbounng Tooth-bills also appeared to in- stantaneously copy each others ‘mimicry’, ‘chuck’ or ‘babble* calls. Tooth-bill mimicry models changed seasonal- ly. For example, during season 79 Fan-tailed Cuckoos were extremely vocal in the study area and their calls were mimicked by all eight males (12% of ‘mimicry’). During season 80 we heard fewer Fan-tailed Cuckoos and less (1%) 'mimicry' of them. The proportion of 'mimicry' incorporated into song did not differ greatly over the threc peak activity months of season 80 (Oc- tober = 7%, November = 9%, December = 10% of total song time), although increasing towards peak mating time and falling to an average of 6% in post-peak activity. Moreover, it did not differ much throughout the day. During the periods 0600-0900, 0900-1200, 1200-1500, and 1500- 1800h males performed ‘mimicry’ for6, 10, R and 10% of total song time respectively over seasons 79 and 80. Bird species that regularly called in the study area during Tooth-bill display seasons but were not, or only rarely, mimicked included the Noisy Pitta Pitra versicolor, Victoria's Riflebird Piiloris victoriae and the Spotted Catbird. The whistled bower advertisement call of male Satin Bowerbirds and the sound of dripping water was mimicked by Tooth-bills in display subsong but not during advertisement singing. DISCUSSION COURT SEASONALITY AND MAINTENANCE Male Tooth-bills started court clearing in Sep- tember-October, approximately six to eighi weeks before first known egg-laying (Marshall, 1951; Poth & Frith, 1985a, 1993) and increasing- ly attended courts from October-December (Fig. 138 2). Food resources are abundant at this titie of year (Frith 1984; Frith & Frith, 1990, 1985b), Although Marshall (1951, 1954) suggested in- creasing daylight tîme, rising temperatures and decreasing humidity might instigate the seasonal sexual resurgence of male Tooth-bills our data do not reveal clear-cut relationships between Weather patterns and variation in the initiation of pre-peak activity across years. Pre-peak activity lasted three, five and two weeks during seasons 78, 79 and 80 respectively; interesting differences because whilst «empera- tires Were seasonally increasing slightly, relative rainfall differed between seasons (Fig. 3a,b). Relative fruit crop, however, was poor dunng 79 and rich during 78 and 80 (Fig, 3c), and it is possible that this influenced these differing lengths of pre-peak activity. Peak court activity lasted for ten, six and 14 wecks during seasons 78, 79 and 80 respectively. A prolific fruit crop became available in Novern- ber 78, the 79 fruit crop failed and in season 80 peak activily was preceeded and accompanied by much fruit (Fig. 3c). Late peak court activity, and nesting coincided (November-February) with the months of relatively far greater beetle abundance at Paluma (Frith & Frith, 1985a,h), Coleoptera apparently being important in the nestling diet (Jackson, 1909; Marshall, 1951). Lapses in court decoration rates during the long period of peak court attendance in scason 80 (Fig. 2) coincided with increasing temperatures during drier periods. At such times court decoration leaves dry and curl quickly and extremes in these conditions, such as during season 79, may cause birds to desert courts, The start of post-peak activity is strongly as- sociated with persistent heavy rains of wel season months January-February but during season 79 it preceeded this due to excessively dry conditions (Fig. 3b). Significant dry spells within the wet season rains stimulated a brief return to renewed court activity, Warham (1962) correctly thought the onset of the wet season caused cessation of court attendance and the start of moult. During March-April of several seasons we ob- served some males calling and poorly clearing and decorating courts. This would appear to be a post-moult activity performed mostly by sub- adult males, as occurs m other bowerbird species (Frith & Frith, unpubl. data). Similar behaviour was also observed in June-July 1980, which may have related to an abundant winter fruit crop (Fig, 3c) thal supported a subsequent carly start to the 80 display season. MEMOIRS OF THE QUEENSLAND MUSEUM COURT CHARACTERISTICS AND DECORATION Marshall (1954) found the average number of leaves on a Tooth-bil] court to be c.40 and the maximum 103 whereas Schodde & Tidemann (1988) give an crroneousty low &-15 Jcaves/court. Grant & Laurance (1991) found an average of 23-95 leaves a court for their 27 September-17 December study and an average of 52 leaves to 15 October and only 37 thereafter. We recorded an average of between 60-78 per season and a largest number of 108-179 leaves per coun over five seasons (Table 1), Chaffer (1984) gained the impression that fewer leaves were used if they were of larger size but this could not be tested in our study area where birds preferred plant species with smaller leaves, Male Tooth-bills m this study decorated courts predominantly with Polyscias leaves (Table 1) and only infrequently with leaves of 14 other tree species. Jackson (1910) listed the 14 most com- monly used leaves he found on Atherton Tableland courts but does not include Polyscias. Marshall (1954) lists identifications of 181 Icaves from five Atherton Tableland courts and does not include Polyscias which are, however, used by Atherton Tableland birds (Frith & Frith, unpubl. data). Grant & Laurance (1991) found thal decorations on 13 Atherton Tableland courts con- sisted of >90% Schlefflera actinophylla leaves, These facts suggest geographic variation in favoured court decorations, a situation found in bower decorations of the two Satin Bowerbird subspecies (Chaffer, 1984; Frith & Frith, unpubl. data) and between populations of the Vogelkop Bowerbird Amblyernis. inomaus. Diamond (1986, 1987) considered this tn be indicative of local tradition based on imitive learning, Whilst traditional court-owning Tooth-bills at Paluma very predominantly used Polyscias leaves we noted that courts established at new sites or traditional sites that were only partly re-established during a season had fewer Polys- cias leaves, This suggests to us that more Polycias was indicative of relative male experience as we consider males attempting to establish new siles or replacing traditional owners at their traditional sites to be younger birds (Frith & Frith, in press). In some cases where an unknown, unmarked, male took over the traditional court site of a traditional marked male al the start of a display season. we observed the presumed relatively in- experienced male to clear an inferior court and decorate it poorly. Such newly-aquired courts and their decoration improve over subsequent seasons, Thus, long established, older, coun- TOOTH-BILLED BOWERBIRDS owning males used mostly Polysciay leaves whereas younger and/or peripheral males used less, Older males also selectively stole Pelyscias leaves from courts of peripheral, probably sub- dominant, neighbouring males. These con- clusions are supported by the findings of Borgia (19&5a, 1986) who found the ability of male Satin Bowerbirds to build better quality bowers and retain greater numbers of decorations to be a function of age. The more significant bower decorations of some other bowerbird species are items rare in the birds’ environment (Borgia & Gore, 1986; Frith, 1989; Frith & Frith, 1990c) and it appears that an abundance of such decorations on bowers im- proves the owners mating success (Borgia, 1986). Rare decorations might, thus, indicate something significant to nval males and to females about fitness, and possibly level of dominance, of the males acquiring and retaining them. Itis doubtful, however, that Polyscias leaves are rare in the Tooth-bill*s present environment for in consider- ing this possibility the modification or degrada- tion of Tooth-bill habitat must be kept in mind, Pioneering Polyscias in our study area were char- acteristic of disturbed forest edge and gaps. Prior Io any upland rainforest disturbance (i.e, roads, snig tracks, logging ete.) this plant would certain- ly have been less abundant and more patchily distnbuted than it is today. This aside, however, it is possible that Polyscias presents a leaf that has a structure and moisture content that gives a long 'court-life” and provides a strikingly contrasting pale underside. We were unable to study male mating success relative to court position, size, or number of leaf decorations, We observed that a display tree, at or within the court periphery, is fundamentally important 1o enable males ro initiate courtship by hiding from the female behind (Frith & Frith, 1993), Whilst all long-term traditional courts had at least one display tree a number of small and temporary courts, apparently made by younger males early or late in thc display season, lacked such a tree. As the initial clearing of the court of all litter and debris doubtless involves consider- able effort by, and cost to, the male and, given the observed variation in the size of area cleared it is possible that court size may be of some sig- nificance to females in mate selection. It should be noted that Grant & Laurance (1991) did not discuss court-size but unfortunately used the term ‘court size’ to represent the number of leaves on 4 court. 139 Of 54 courts examined for the presence of lawyer cane plants growing within them 37% proved to have them, indicating that this is not a Prerequisite for a court location as suggested (Jackson, 1910). All fruit remains found on courts, often in discrete piles beneath favoured perches, are excreted by birds and not ‘cast or vomited' as suggested by Jackson (1910). LEAF TURNOVER RATES AND THEFT Courts examined for leaf numbers, species and evidence of leaf stealing were those of part of the dense population of males on Tooth-bill Hill and adjacent slopes. These were traditional court- owners, which we concluded consisted of older and more expenenced individuals. In most cases, colour-banding proved this to be true (Frith & Frith, in press). Most leaves remained on courts for two weeks or less but some did so for up to three (dry season 79) or four (season 80) weeks, leaf turnover rates being faster with drier conditions. These results agree with Grant & Laurance (1991) who found most leaves were removed at 13 courts after Wo wecks or less. We found Ieaf turnover rates to be significantly higher during hotter drier periods of peak activity and, given thal leaves are presumab- ly selected for contrasting paler undersides, large size, [Tat structure and moisture retention (anti- curling properties), this is predictable as dehydra- tion renders leaves unsuitable decorations more quickly, We performed no experiments on rela- tive dehydration and deterioration rates of various Jeaf species, but observed that leaves of other species dried, curled and discoloured quick- er than did Polyscias. We found a peak activity court decoration rate of 5.9 and 6.8 and a removal rate of 7.3 and 6.4 leaves/day in seasons 79 and 80 respectively. These rates are higher than the 4.05 and 3.96 leaves added and removed per day respectively observed by Grant & Laurance í 1991). Differen- ces in leaf decoration and removal rates between individual males may reflect relative differences in experience, and/or dominance within the population of adjacent males. Grant & Laurances' (1991) period of study, however, in- cluded pre-peak activity, which would have lowered the overall rate of observed leaf replace- ment. There were more leaves on courts during early peak uctivity in season 80 with a greater propor- tion of them Palyseias. Relatively fewer Polys- cias leaves were present during later peak activity (and fewer still during post-peak activity), which 140 may sugges! competition for favolired decora- tions is More intense initially each season, when males are establishing or re-establishing a social heirachy, More leaf thefts occurred during early peak activity. indicating greater inter-male còm- petition, and these mostly involved Potyscias. Movement of stolen leaves among several ad- jacent well-established marked males’ courts in- dicated the possibility of a social hierarchy. That one Of nine adjacent individuals proved to steal more leaves from more adjacent cours than did al) eight other birds combined, and apparently inhibited al! other males from stealing from him (Fig, 5), is indicative of a system similar to that operating in some other polygynous bowerbird species (Borgia, 1985a,b; Borgia & Mueller, 1992). It is possible that more acquisitively suc- cessful court-owning males might obtain most successful matings within the localised dense tion of court-owning males, or exploded lek (Frith & Frith, in press), Borgia (1985a) demonstrajed that male Satin Bowerbirds restrict their bower decoration theft raids to immediate neighbours, their most likely sexual competitors, as we do in the Tooth-bill (Fig. 5). He found, moreover, a highly significant correlation between male social dominance of individuals at feeding sites, and the number of bower destructions performed by the individuals. thereby providing evidence in support of bower interference being an aggressive act by which more dominant males can detrimentally affect the quality of bowers of rival males. We obtained no empincal data to support our hypothesis bul we think it possible that a relatively large number of fresh and appropriate leaves on the court of a male, within an exploded lek. reflects that male's expenence among the local male population and perhaps also his relative dominance, MALE ATTENDANCE LEVELS AT COURTS Over the peak court activity of season 79, 80 and 89 males spent an average of 64% of time attending courts, By radio-tagging four males, we G. Moore showed that a bird was potentially present but silent high above his court where he might have otherwise been overlooked by an observer within a hide for up to 3% of total time. Differing court attendance levels by males during seasons 79, 80 and 89 clearly reflect seasonal environmental differences, being lowest during 79 (52%) and highest during 80 (79%). This pattem was the same during each month of peak activity and at different times of day. MEMOIRS OF THE QUEENSLAND MUSEUM The time that individual male Tooth-bills spent at courts during season 79 was lower and more variable than during $0 and £9. Males took longer absences from courts both during different months of peak activity and times of day in season 79, than 80 and 89. During 79, peak activity lasted only six weeks and it would appear that the dry conditions and poor fruit crop prevailing during this season were detrimental. The exceptionally dry early December of season 79 combined with a poor fruit crop caused birds to quit courts (Figs 3 & 6). Coun attendance levels by all males was high during season RO and mean duration of absences from their courts was 3,7 and 2,2 times briefer than during seasons 79 and 89 respectively (Table 3) which would seem attributable to relatively greater availability of fruit (Fig. 3c), While the period of early peak activity of season 80 was relatively dry, fruit was prolific throughout the prolonged peak activity of 14 weeks. The onset of wel season rains terminated court attendance and, while significantly dry spells thereafter brought birds briefly back to courts, court atten- dance was low during post-peak activity (Table 4), Peak activity lasted ten weeks in season 89 with attendance levels being intermediate be- tween those of 79 and 80 (Table 3). That the carly 89 peak activity period was relatively dry, the November wet (Fig. 6), and the fruit crop rela- tively poor (Moore, 1991) possibly accounts for this. Most polysynously-breeding passerines are in- habitants of tropical rainforests that eat predominantly, if not exclusively, fruits which are abundant in space and time in that habitat and require, therefore, a small proportion of the day to harvest, Jt has been argued that these ecological factors favour the evolution of polygyny because seasonally abundant fruit resources enable males to devote most of their time to courtship and females are able to provision their offspring un- aided by the promiscuous males (Snow, 1976, 1982: Snow & Snow, 1988; Beehler , 1989). Male Tooth-bills are almost exclusively frugivorous during their courtship and mating season (Frith & Frith, unpubl, data) and while nesting females are known to also feed insects to their young, the diet of the offspring is probably predominantly fruit (Frith & Frith, 1985a). Thus, this bowerbird fits well the ecological scenario common to most polygynnus passennes. In the case of some neotropical polygynous passerines, males have been observed to spend as much as 87-90% of daylight hours attending display TOOTH-RILLED BOWERBIRDS courts or perches (Snow & Snow, 1988), It is most noteworthy, therefore, that male Tooth-bills appear far more frigivorous than males of other bowerhird species and spend sigmficantly longer al their courts than do males of other bowerbirds. This was emphasised because during this study, male Tooth-bills spent significantly longer at their courts during the season of highest fruit abundance. Male Satin Bowerbirds, a species with a 67% fruit component of annual diet, spent an average of 73% of daytime within 50m of their bowers (Donaghey, 1981), Males of the predominantly frugivorous Macgregor's Bowerbird Ambiyornis macgregoriae spent 54% of daylight at, or within 20m of, their bowers (Pruett-Jones & Pruett- Jones, 1982, 1985). A male Great Bowerbird, a species considered Io be predominantly frugivorous (Diamond, 1986; Schodde & Tidemann, 1988) spent 47% of daylight hours at or near his bower (Veselovsky, 1978). The Regent Bowerbird's annual diet includes 81% fruit and yet males were found to spend a mere 3% of daytime at or near their bowers. The ex- planation for this exception is that regent bower- birds Sericulus species represent an early stage in the evolution of bower-building, in which bowers have no: yet replaced elaborate male nuptial plumage, Male Regent Bowerbirds S. chrysocephalus imitiate courtship in the forest canopy before accompanying the female to the bower where a prolonged courtship display primarily presents colourful puptia) plumage, not the bower, to the female (Lenz, 1993). MALE BEHAVIOUR AT COURTS The length of time males spent at courts at each visiL was similar for individual males during each season, different months of activity and at different times of the day (Tables 3-5), Once males started Io attend courts the proportion of time they spent giving advertisement song, per- forming court mainienance or displaying was similar each season (Tables 6-8). Of their time at courts males invested 95,7% in performing loud advertisement song from favoured perches, 2.4% in silence, 1.2% in court maintainence and 0.7% in display, Most (99%) advertisement song was performed by male Tooth-bills from favoured high, adjacent or court perches, males rarely singing on the ground prior to display subsong. Birds spent more time singing on high and adjacent perches during the drier and hotter 79 season than in 80 and spent more time on higher perches during the brighter 141 and hotter time of day, During season 80, males spent more time on court perches as the season progressed, Most birds frequented only one or two, favoured, adjacent court perches, resulting in large accumulations of excreted fruit remains beneath them. Male Tooth-bills spent a mean of 1.4mins maintaining courts. Given that males were present immediately above their courts for 64% of total time, the 1,2% of time spent in court maintenance is remarkably brief compared with mean proportion of lime spent in bower main- tenance by Macgregor's Bowerbirds (12%; PruetiJones & Pruett-Jones, 1982), Sann Bower- birds (8%; Donaghey, 1981), Great Bowerbirds (Veselovsky, 1978), Western Bowerbirds C. gut- fate (Bradley, 1987) Spotted Bowerbirds C. maculata, Fawn-breasted Bowerbirds C. cer- viniventris and Lauterbach's Bowerbirds C. lauterbachi (Frith & Frith, 1989, unpubl. data). It is similar, however, to that invested by the Regent Bowerbird which builds the most rudimentary and sparsely decorated of howers (Lenz, 1993), Male Tooth-bills do not spend much time on the court ‘searching for insects and snails’ as reported by Forshaw (in Cooper & Forshaw, 1977). The limited time they did spend on the cleared court area presumably merely reflects the low main- tenance required once it is initially cleared and decorated, The cryptic morphology of males, their habit of remaining all but motionless when perched, of being silent when maintaining the court, the ventriloquistic quality of their calls, the exireme brevity of their courtship display (Frith & Frith, 1993), the adoption of a sleeked frozen posture when alarmed and Ihe fact that they are not infrequently preyed on at their courts by goshawks Accipiter spp. (Frith & Frith, unpubl, data) all suggest that predation might have sig- nificantly influenced some or all of this be- haviour. The fact that we witnessed not a single aggressive encounter between birds at a court other than owning birds chasing off another sug- gests such events are extremely rare. MALE VOCALISATIONS AT COURTS Bower-attending bowerbirds studied to date advertise the bower location with specific calls given relatively infrequently (Gilliard, 1969; Cooper & Forshaw, 1977; Veselosky, 1978; Donaghey, 1981; Frith & Frith, unpubl, data) and use a distinctly different, usually quieter and more complex, song in courtship display that in the case of the Satin Bowerbird (Loffredo & Borgia. 1986), the Regent Bowerbird (Lenz, 142 1993) and gardener bowerbirds Amthlyernis spp. (Frith & Frith, 1993, Frith & Frith, unpubl. data) often includes avian mimicry. Tooth-bill, Satin, Macgregor's, Streaked Bowerbird Amblyomis subalaris and Archbold’s Bowerbird Archboldia papuensis calls lack directional cues or have ventriloguistic gualities (Frith & Frith, unpubl. data). It has been suggested that this is a result of the enhanced risk of predation upon birds calling from long-term traditional locations (Robinson, 1974). Advertisement song of Tooth-bills consisted of seven component calls. Phases of several to all of these calls were performed dunng each visit and usually followed a definite sequence. Thus males returning to courts invariably first gave a ‘chuck’, as noted by Chaffer (1984), or less frequently a squeal. “Chuck” and ‘squeal’ calls were followed by ‘chuck + babble’ or “chuck + mimucry'. These were followed by ‘babble’, ‘babble + mimicry’ and/or ‘mimicry’. Display vocalisatrons (Frith & Frith, 1993) usually followed the latterthree calls. Sometimes, bul not often, the sequence of calls was broken when males changed position on a, or moved to another, perch, In these instances they often gave ‘chuck’ culls again. The ‘chuck’ ap- peared to us the loudest, most far-carrying, and most locatable call, This may explain why “chuck' is used to recommence calling, as it clear- ly and quickly establishes the bird's presence and location. Itis possible the ‘chock’, which we here treat as a natural Tooth-bill call, ts in fact mimicry of a similar note of Bower's Shrike Thrush, par- ticularly as calls of this species dominated Tooth- bill mimicry. Of the lotal lime males spent advertisement singing at their courts, at least 61% of yocalisations included elements of mimicry (Table 10). Calling male Tooth-bills at locations remote from Paluma sound similar in general terms but incorporate some different avian model species into their mimicry and/or give different bias to different species (C. & D. Frith, unpubl. data) as is found tn Satin Bowerbirds (Loffredo & Borgia, 1986) and Jyrebirds Menure spp. (Smith, L988) at different areas. This is indicative of the kind of “culturally” transmitted behavior seen expressed in the bower architecture and decorations of bowers of geographically distinct populations of bowerbitd species (Diamond, 1986, 1987). While the loud song of the Tooth-bill presumably functions to indicate presence to rival males (Marshall, 1950) we think the conspicuous mimicry content may function to advertise court location and bird status to females (see below). MEMOIRS OF THE QUEENSLAND MUSEUM Once a female shows serious interest in the court the male Tooth-bill performs a far softer 'subsong” exclusively of mimicry (Fnth & Frith, 1993), as do male Satin Bowerbirds and Noisy Scrub-birds Atrichornis clamosus (Loffredo & Borgia, 1986; Robinson, 1974), We think it likely that this subsong is in fact more informative to females than advertisement song as it has been demonstrated that female Satin Bowerbirds use the quality of more intimate male mimicry to assess the relative merits of prospective mates (Loffredo & Borgia, 1986), Contrary to Robinson (1974), but consistent with Dobkin's (1979) in- terpretation that male bowerbird mimicry is given in precopulatory display (and in nest defence by females), the male Tooth-hill does specifically perform mimicry prior to and during courtship displays, as male Satin Bawerbirds appear to do (Loffredo & Borgia, 1986). Loffredo & Borgia (1986) showed that among competing male Satin Bowerbirds, older males produced longer bouts of higher-quality avian vocal mimicry than younger males and as a con- sequence gained higher mating success. These authors also found that the male Satin Bowerbirds in their study area performed mimicry of only two bird species during courtship and that when both were given lhey were always performed in the same order. During subsong mimicry male Tooth-bills showed a strong tendency to perform their repertoire of approximately ten model species and other sounds in a particular order and within a minute (Frith & Frith, 1993), The nature of male Tooth-bill vocalisauons would suggest that females may make some general assessment of male experience from their loud court advertisement song prior Io selecting individual males and/or courts for visitations. This initial loud ‘broadcasting’ of mimicry may reflect pressures of greater male competitor den- sity. During the subsequent court visit females can assess the quantity and quality of the more stereotyped sub-song exclusively of mimicry (Frith & Frith, 1993), Thorpe (1985) stated that there is some evidence that variety in male bird song is attractive to females, and suggested that mimicry may simply be a way of increasing repertoire size. The repertoire of singing male court owners was found to change, by other bird calls being opportunistically mimicked, thus incorporating novel characters into the sang. Bourke & Austin (1947) also noted that males would stop calling to listen to other bird species and then immedi- ately mimic the bird listened to. Goodwin (1986) TOOTH-BILLED BOWERBIRDS reported several wild Jays Garralus gladurius mimicking the call of the Grey Heron Ardea cinereus immediately upon seeing this species, Smith (1988) observed the same behaviour by mimicking, promiscuous male, lyrebirds. The significance of such individualistic call com- nts is unknown. The idea that females may in fact select for novel secondary sexual charac- teristics has, however, recently been presented (Ten Cate & Bateson, 1988; Christidis & Schodde, 1993), lt is possible this theory may have significance to the question of avian mimicry in addition to the finding of Loffredo & Borgia (1986) that male Satin Bowerbirds per- forming more and better mimicry were older individuals obtaining higher mating success, Hoglund (1989) observed that in lekking birds all species that display on the ground have larger males than females, with the exception of the Great Snipe Gallinago media. He noted that the male Great Snipe uses vocalisations rather than nuptial plumage to enhance reproductive success. It is noteworthy, therefore, that the terrestrially- displaying Tooth-bill is cryptically monomorphic in plumage, nearly so in size, and that males use prolonged complex vocalisations but a markedly bricf display (Frith & Frith, 1993) in courtship. Five adult male Macgregor's Bowerhirds, decorated only with an extensive orange crest, Spent an average of 71% of court attendance time perched silently and 14% vocalising from tradi- tional perches (Pructt-Jones & Pructt-Jones, 1982), Brilliantly-plumuged adult male Regent Bowerbirds spent a mere 1.01% of their total, brief, court attendance time vocalising (Lenz. 1993), It is possible, therefore, that continuous Ioud complex vocalisations by male Tooth-hills, at least in part, compensates for the lack of visual secondary sexual characters by containing infor- mation that rival males and potential mates can use to assess male ‘fitness’, It has been noted that in the evolutionarily convergent polygynous Cotingas (Passeriformes, Cotingidae) of the Neotropics, in which most species are con- spicuously sexually dimorphic, the lekking males of the few species with much-reduced or no sexual dimorphism use vocal behaviour to attract and court females far more than their relatives. (Snow 1982). Male Tooth-bill fitness. could be indicated by the amount of calling (time invested at the court) and/or its qualities and proportion of avian mimicry (reflecting male experience), Lof- fredo & Borgia (1986) found that female Satin Bowerbirds use male yocalisations, particularly 143 the quality of mimicked bird calls, as an indica- tion of relative male age. They found that older males perform better mimicry and that females selected for the ability in males tò learn compli- cated songs. Thus, while female Tooth-bills cannot simul- taneously compare members of an ‘exploded! aggregation (lek) of males displaying their vigor and plumage (as females in true leks can) they may be able to simultaneously assess court atten- dance levels as indicated by vocalization time and the quality of song content. By this means females could initially assess male age (cf. Lof- fredo & Borgia, 1986) from perches above or adajcent to areas of grealer court densities that in effect form exploded leks (Frith & Frith, in press), prior to selecting a male(s) to visit ACKNOWLEDGEMENTS The initial three years of this study were in part supported by National Geographic Society Grants 1709 and 1870 tò Alan Lill, Ecology and Evolutionary Biology Department, Monash University. We thank Alan Lill for his support, encouragement and advice and the then Zoolozy Department, Monash University for help in many ways. The Wet Tropics Management Agency and Authority kindly provided funds to assist with preparation and production of this contribution. For thoughtful and constructive criticism of and discussion on a draft(s) of this paper we sincerely thank Jared Diamond, Peter Dwyer, Glen Ingram, Norbert Lenz, Alan Lall and Stephen Pruett-Jones for their time and trouble. Andrée Griffin provided valued company and assistance in the field. Bernie Hyland and Jeff Tracey kindly iden- tified leaf decorations, We thank Geoff Moore of the Department of Zoology, James Cook Univer- sity of North Queensland for use of radio-tracking equipment and for radio-tracking marked birds with us at and away from their courts during our 1989 study, this being part of his PhD study on seed dispersal by male Tonth-billed Bowerbirds. Dedicated to Andrée Griffin. in appreciation. LITERATURE CITED BEEHLER, B.M. 1989. The birds of paradise. Scientific American 261:117-123, BORGIA, G. 1985a. Bower quality, number of dewi- tions and mating success of male satin bowerbirds (Ptilonorhynchay violaceus). an experimental analysis. Animal Behaviour 33:266-271. 1985b. Bower destruction and sexual competition Mi the satin bowerbird (Ptilonorhynchus violaceus}. Behavioural Ecology and Sociobiology 18: 91- 100 1986, Sexual selection in bowerbirds. Scientific American 254:70-79. BORGIA, G. & GORE, M.A, 1986, Feather stealing in the satin bowerbird (Pnilenorhynehus violaceus): male competition and the quality of display, Animal Behaviour 34:727.738. BORGIA, G. & MUELLER, U. [992, Bower destruc- bon, decoration, stealing and female choice in the Spotted Bowerbird Chlumydera mdculata, En 92:11-18 BOURKE, P.A. & AUSTIN, A.F. 1947. The Atherton Tablelands and its avifauna. Emu 47: 87-116. BRADLEY, J. M. 1987. Vocal behaviour and annual cycle of the Western Bowerbird Chlamydera gwt- tata. Australian Bird Watcher 12:83-90. BROWN, J. L. & ORIANS, G. H, 1970, Spacing pat- tems in mobile animals. Annual Review of Ecol- ogy and Systematics 1;239-262. CHAFFER, N. 1984, ‘In quest of bower birds’, (Rigby; Adelaide). CHRISTIDIS, L. & SCHODDE, R, 1993, Sexual selec- tion for novel partners: a mechanism for ac- celerated morphological evolution in the birds-of-paradise (Paradisaeidae). Bulletin of the British Omithologists’ Club, 113: 169-172, COOPER, W.T. & FORSHAW, J.M. 1977. “The birds of paradise and bower birds’. (Collins:Sydney). DIAMOND, J. 1986. Biology of the birds of paradise: and bowerbirds, Annual Review of Ecology and Systematics 17;17-37, 1987. Bower building and decoration by the bower- bird Ambiyornis inornatus. Ethology 74:177- 204 DOBKIN, D.S. 1979. Functional and evolutionary relationships of vocal copying phenomena. in birds, Zeitschnifl Tierpsychologie 50:348-363. DONAGHEY, R. H. 1981. “The ecology and evolution of bowerbird mating systems’, (Unpublished PhD Thesis; Monash University, Clayton), DONAGHEY, R.H.. FRITH, C.B. & LILL, A. 1985. 'Bowerbind', Pp 60-62, In Campbell, B. & Lack, E. (eds), ‘A dictionary of birds”. (Poyser: Calton). FRITH, C 1989, A construction warker in the rain- forest. Birds Intemational 1:29-39. 1993, The Tooth-billed Bowerbird (Scengpoeeter dentirostris). North Queensland Naturalist 195:7-14. FRITH, C,B. & FRITH, D.W, 1985a. Parental care and investment in the Tooth-billed Bowerbird Scenopeeetes. dentirosiris (Pulonochynchudae), Australian Bird Watcher 113103-113, 1985b, Seasonality of insect abundance in an Austrafain upland tropical rainforest. Australian Journal of Ecology 10:237-248. 1989. Miscellaneous notes on the bowerbirds Chlamydera cerviniventris and C, laurerbachi (Pulonorhynchidae) in Papua New Guinea, Australian Bird Watcher 13: 6-19, MEMOIRS OF THE QUEENSLAND MUSEUM 19%. Notes on the nesting money of the Great Bowerbird Chlamydera ruchalis (Pulonoc- hynchidae). Australian Bird Watcher 13:137- 148 1990b, Noles on the nesting biology af the Spotted Bowerbird Chlamydera macilata (Ptilonor- hynchidae). Australtan Bird Watcher 13:218- 225 1990c. Archbold's Bowerbird Archboldia papuen- sis (Ptilonorhynchidae) uses plumes from King of Saxony Bird of Paradise Pteridophora alberti (Paradisaeidae) as bower decoration, Emu 90136-137. 1993. Courtship display of the Tooth-billed Bower- bird Scenopoeetes dentirostris; its behavioural and systematic significance, Emu 93:129-136, 1994, The nesting biology of Archbold's Bowerbird Archbaldia papuensis and a review of that of wher bowerbirds (Pulonorhynchidue). Ibis 136: 153-160. IN PRESS. Court site constancy, dispersion, male survival und court ownership in the male Tooth- billed Bowerbird, Scenopoeeres dentirostris (Pulonorhynchidae). Emu 95. FRITH, D. 1984. Foraging ecology of birds in an upland tropical rainforestin north Queensland. Australian Wildlife Research 1 1:325-347. FRITH, D. & FRITH, C. 1990. Seasonality of litter invertebrate populations in an Australian upland tropical rain forest. Biotropica 22:181-190. GILLIARD, E.T. 1969. “Birds of paradise and bower birds’, (Weidenfeld & Nicolson:London). GOODWIN, D, 1986. "Crows of the world’. (British Museum of Natural History:London), GRANT, J.D. & LAURANCE, W.F. 1991. Court size and maintenance in the Tooth-billed Catbird. Sun- bird 21;90-92. HOGLUND, J. 1989, Size and plumage dimorphism in lek-breeding birds: a comparative analysis, The American Naturalist 134;72-87. JACKSON, S.W. 1909. In the Barron River Valley, North Queensland. Emu 8:233-285. 1910, Additional notes on the Tooth-billed Bower- bird (Scensopoeeles dentirastris) of North Queensland. Emu 10;81-88, LENZ, N.H.G. 1993. Behavioural and reproductive biology of the Regent Bowerbird, Sericulus chyr- secephalus (Lewin, 1808). (Unpublished PhD Thesis, Griffith University: Nathan). LOFFREDO, C.A. & BORGIA, G. 1986. Male courtship vocalisations as cues for mate choice in the Satin Bowerbird (Piilonerknychus vielaceus), Auk 103:189-195. MARSHALL, A. J. 1950. The function of vocal mimicry, Emu 50; 5-16. 1951. Leaf-display and the sexual cycle in the Tooth-billed “Bowerbird” (Scenopoeeres den- (irostris, Ramsay), Proceedings of the Zoological Society of London 120:749-758. 1954. ‘Bower-birds - their displays and breeding cycles’. (Oxford University Press: Oxford). TOOTH-BILLED BOWERBIRDS MOORE, G.J. 1991. Seed dispersal by male Tooth- billed Bowerbirds Scenopoeetes dentirostris (Ptilonorhynchidae), in north-east Queensland rainforest: Processes and consequences. (Un- published PhD Thesis: James Cook University of North Queensland: Townsville). NIX, H.A. & SWITZER, M.A. 1991. Rainforest animals - atlas of vertebrates endemic to Australia's Wet Tropics. Kowari 1:1-112, PRUETT-JONES, M.A. & PRUETT-JONES, S.G. 1982. Spacing and distribution of bowers in Macgregor's Bowerbird _ (Amblyomis macgregoriae). Behaviour, Ecology and Sociobiology 11:25-32. 1985. Food caching in the tropical frugivore Macgregor's Bowerbird (Amblyornis macgregoriae). Auk 102:334-341. 1993. Sexual competition and courtship disruption : why do male bowerbirds destroy each others bowers? Animal Behaviour. 47: 607-620. ROBINSON, F.N. 1974. The function of vocal mimicry in some avian displays. Emu 74:9-10. SCHODDE, R. & TIDEMANN, S. C. 1988. “Reader's digest complete book of Australian birds'. (Reader's Digest:Sydney). SMITH, L. H. 1988. 'The life of the lyrebird'. (Heinemann: Richmond). SNOW, D.W. 1962. A field study of the Black and White Manakin Manacus manacus in Trinidad. Zoologica 47, 65-104. 1976, ‘The web of adaptation’, (Collins: London). 1982. ‘The cotingas’. (British Museum of Natural History & Oxford University Press: London). SNOW, B.K, & SNOW, D.W. 1988. ‘Birds and berries’, (Poyser: Calton). TEN CATE, C. & BATESON, P. 1988. Sexual selec- tion: the evolution of conspicuous characteristics in birds by means of inprinting. Evolution 42:1355-1358. THORPE, W.H. 1985. Mimicry, vocal. Pp. 353-355. In Campbell, B. & Lack, E. (eds), *A dictionary of birds'. (Poyser: Calton). TRACEY, J.G. 1982. ‘The vegetation of the humid tropical region of North Queensland’, (CSIRO: Melbourne). VELLENGA, R. E. 1970. Behaviour of the male Satin Bowerbird at the bower. The Australian Bird Bander 8:3-11, 1980. Distribution of bowers of the Satin Bowerbird at Leura, NSW, with notes on parental care, development and independence of young. Emu 80: 97-102. VESELOVSKY, Z. 1978. Beobachtungen zu biologie und verhalten des groben kragenlaubensvogels (Chlamydera nuchalis). Journal fiir Ornithologie 119:74-90. WARHAM, J. 1962. Field notes on Australian bower- birds and cat-birds. Emu 62:1-30. 146 ANEW FRESHWATER BIVALVE FROM THE TRIAS- SIC OF SOUTHEASTERN QUEENSLAND. Memoirs of the Queensland Museum 37 (1):146. 1994:- Collections made from the Blackstone Formation, Ipswich Coal Measures, have yielded, Protovirgus clellandi sp. nov. in addition to bivalves reported by Etheridge (1892), McMichael (1957), and Hill et al. (1965). P. clellandi sp. nov. is also reported from the Tingalpa Formation. A Late Triassic (Carnian) age has been ascribed to the Blackstone Formation by de Jersey (1975), and the Tingalpa Formation (de Jersey & Hamilton, 1965). Class BIVALVIA Linne, 1758 Order UNIOIDA Stoliczka, 1871 Family UNIONIDAE Fleming, 1828 Protovirgus McMichael, 1956 Protovirgus clellandi sp. nov. Material. Holotype: QMF29473, from mullock heaps at Ebbw Vale, which are derived from the Blackstone Formation, Claypave Quarry, Dinmore. Paratypes: QMF29475, QMF32226, same data as holotype; QMF27523, from road cutting on Gateway Arterial road, Brisbane, Tingalpa Forma- tion. Diagnosis. Medium sized, equivalved, elongate unioid with inflated umbonal area and fine comarginal ornament. Hinge straight and long. Tapering strongly to the posterior. Ventral margin is sharp, Dentition unknown. Anterior adductor scars are raised on platforms near the umbones. Description. Maximum height ranges from 28-39mm, width 27-36mm and in length 75-123mm (Table 1). Height and width is approximately egual along the shell length. Hinge straight, the ligament is apparently wide and extends ap- proximately 2/3 of the shell length. Umbones anterior, placed at no more than 1/6 of shell length. Strongly tapering posteriorly, shell is apparently thin with ornament of fine growth lines. Rounded at posterior margin. Anterior adductor scar just ventral of umbo consists of slightly raised platform with 3 or 4 small slightly ovate pits. Posterior muscle scar unknown, Dentition unknown. FIG. 1. Protovirgus clellandi sp.nov. A-C, OMF29473, x0.3. A, left valve; B, right valve; C, dorsal view. D, paratype, QMF29475, right valve x0.5. MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1. Morphometric data for specimens of Protovirgus clellandi sp. nov. from the Blackstone and Tingalpa Forma- tions. Measurements in mm. Tar ri a ink ad a YY WG QMF29473 |39 123 36 Remarks. The tapering form, rounded posterior end, sharp ventral margin and strongly anterior umbones ally the present material with Protovirgus McMichael, 1956. The type species, P. dunstani (Etheridge Jr., 1888 described by Mc- Michael 1956: 232, fig. 8) from the Triassic of New South Wales is much smaller and less robust than P. clellandi. P. flemingi McMichael (1956: 232-233, figs 1-3) from Cretaceous of New Zealand is also smaller and the umbones are less anteriorly placed. P, jaenschi Ludbrook, 1961 from Triassic strata at Leigh Creek, South Australia has a greater height to length ratio, and is more gently tapering than P. clellandi. P. coatsi Ludbrook, 1961 is smaller and does not taper as strongly as P. clellandi, resulting in a far more ovate outline. Etymology. For John Clelland Hocknull. Acknowledgements Alex Cook and Natalie Camilleri are to be thanked for their encouragement Literature Cited DE JERSEY, N. J. & HAMILTON, M. 1965. Triassic microfloras of the Mooroka and Tingalpa Formations. Queensland Government Mining Journal 66: 315-318. DE JERSEY, N.J. 1975. Miospore zones in the Lower Mesozoic of southeastern Queensland. Pp. 159-163. In Campbell, K. S. W. (ed.), ‘Papers from the Third Gondwana Symposium’. (ANU Press: Canberra). ETHRIDGE JR., R. 1892. The organic remains of the Ipswich Formation. In Jack, R. L. & Etheridge, R.(eds), “The Geology and Palaeontology of Queensland and New Guinea’. Geological Survey of Queensland, Publica- tion 92. HAAS, F. 1969. Superfamily Unionacea. Pp. N41 1-468. In Moore, R. C. (ed.),* Treatise on Invertebrate Palaeon- tology Part N’. Vol. 2, Mollusca 6, Bivalvia. (Geologi- cal Society of America and University of Kansas: Lawrence, Kansas). HILL, D., PLAYFORD, G. & WOODS, J. T. 1965. ‘Triassic fossils of Queensland’. (Queensland Palaeontographi- cal Society: Brisbane). LUDBROOK, N. H. 1961. Mesozoic non-marine Mollusca (Pelecypoda: Unionidae) from the north of South Australia. Transactions of the Royal Society of South Australia 84: 139-148. McMICHAEL, D.F. 1956. A review of the fossil freshwater mussels (Mollusca, Pelecypoda) Of Australasia. Proceedings of the Linnean Society of New South Wales 81: 232 - 233. Scott A. Hocknull, Queensland Museum PO Box 3300, South Brisbane ,Queensland 4101, Australia; 10 August 1994. HOME RANGE AND EXTRA-COURT ACTIVITY IN THE MALE TOOTH-BILLED BOWERBIRD, SCENOPOEETES DENTIROSTRIS (PYILONORHYNCHIDAE). CLIFFORD B. FRITH, DAWN W, FRITH AND GEOFFREY J. MOORE Frith, € B., Frith, D. W. & Moore, G. J. 1994 12 01: Home range and extra-court activity in the male Tooth-billed Bowerbird yN dentirostris (Piilonorhynchidae). Memoirs of the Queensland Museum 37(1):147-154. Brisbane. ISSN 0079-8835 Home range, time utilisation and activities of four male Tooth-billed Bowerbirds away from their adjacent courts were monitored by radio-tracking throughout the 1989 display season in upland rainforest of north Queensland. Three of the males were known to be adults, having attended their courts for ten or more seasons. Males spent an average of 36% of their lime absent from courts during peak display season activity. The mean home range of males was 9.5ha, with a mean overlap of 50%. Median distances travelled by males from their courts for all activities ranged from 40-86m, with a maximum range of 311-391m. Activities away from courts included foraging, collection of leaf decorations directly trom trees or by theft from neighbouring courts, interactions with conspecifics, bathing and drinking. Most absen- ces (60%) were primarily for foraging which represented 83% of total ime absent with a mean duration of 1 8mins per absence. Some 18% of absences, representing 7% of total time absent, involved exclusive non-foraging activities, Almost all of these absences were brief with a mean duration of 5 mins. The remaining 22% of absences, representing 10% of total lime absent, from the court involved unknown activities, Absences from courts were shortest during November, and during the early part of the moming (0600-0900h). The implications of these data are discussed, [_] Tooth-billed Bowerbird home range size, time urilisarion and activities away from courts, Plilanorhynchidae, Australia. Clifford B. Frith & Dawn W, Frith, Honorary Research Fellows of the Queensland Museum, 'Prionodura', P.O. Box 581, Malanda , Queensland 4885, Australia; Geoffrey J. Moore, Zoology Department, James Cook University of North Queensland, Townsville, Queensland 4811, Australia; present address ; Australian Antarctic Division, Tasmania 7050, Australia; 24 June 1994, Several families of passerine birds include species that exhibit polygynous mating systems in which promiscuous, competitive and often or- nate males take no part in nest duties. In most of these species males spend much of the day at either solitary or communal (lek) display perches; females are attracted to these sites by the calls of the males and are courted by a variety of displays (Gilliard, 1969; Snow, 1982; Beehler & Pructt- Jones, 1983, Beehler 1989). The Tooth-billed Bowerbird, Scenopoeetes dentirostris is one such species (Frith & Frith, 1985, 1993), Males of this species establish cleared courts on the forest floorin areas of suitable topography; mean nearest neighbour distances in these areas are 50m (Frith & Frith, in press). Frith and Frith (1994) found that during peak activity male Tooth-bills spent an average of 64% of the day (0600-1800h) at courts; court visits averaged 23mins and court absences averaged 14mins. The birds spent more time at their courts during seasons of greater fruit abundance. The behaviour of male Tooth-bills at courts is increasingly well understood (Frith & Frith, 1993, 1994; m press). However, nothing is known about home range size, time utilisation and ac- tivities of male Tooth-bills away from their courts, This paper reports observations on the behaviour and movement of four court-ownin male Tooth-bills using radio-tracking and simul- taneous observation from hides near courts. The aims of this work were threefold: to identify the role of male Tooth-bills as seed dispersers within tropical rainforest (Moore, 1991); to determine whether male Tooth-bills spent much time present, but silent, in the canopy above or within the vicinity of their court (Frith & Frith 1994); and to ascertain home ranges and activities of males away from the courts. The third aim is the subject of the present paper, STUDY AREA AND METHODS The study area comprised 50ha of upland trepi- cal rainforest, altitude c.875m, on the Paluma Range (19°00'S, 146°10°E), north-eastern Queensland, 7km from Paluma Township, 80km north of Townsville. Descriptions of the Paluma Range study area and its climate appear in Frith and Frith (1994), Half of the site was divided intà permanently marked 25 X 25m grid squares amd 148 lhe remainder was divided into 25 xX SOm rec- tangles; this enabled accurate plotting of loca- lions at which the birds werc observed. Radio-tracking was undertaken during the 1989 display season (17-27 October, 16-26 November and 12-22 December). Four males (from courts 23, 33, 35, 37; Frith & Frith, 1994: fig. I) were caught in mist nets near their courts and were banded with a metal band and a unique two colour-band combination, Males at courts 23 and 37 had been the traditional owners at those sites for ten previous display seasons and the male al court 35 for 11 previous seasons. The male at site 33 was unbanded at the start of the 89 display season and may have been the unmarked owner of this site for the lwo previous seasons. Each male was monitored for four periods (0600-0900, 0900-1200, 1200-1500, 1500- 1800h) during October, but only three males were monitored during November (at sites 23, 35, 37) and December (23, 33, 37). All males were radio- tracked for a total of 120 hours, The first hour of the first tracking period (at site 33) was excluded from the analysis due to possible disturbance (see below). Male 37 lost a transmitter during one tracking period and, although his court arrivals aid departures were recorded, his activities away from the court were unknown. Racio-tracking was also cerried out after 2000h, for two 30 minute periods at cach court, to ascertain whether the owner Toosted above his court. Each male was fitted with a 4g Biotrack single stage radio-transmitter (SS-1) using a tail mount (Kenward, 1978). Radio-tagged individuals were readily identified by the antenna extending beyond the tail. To assist rapid identification of individuals when they were in the canopy a small umount of contrasting-colaured paint was placed on the underside tips of the outer tail feathers. Radho-racking was iniiated a week afler capture to allow sufficient time for the birds to adjust to the transmiller, We established cryptic canvas hides six metres from each court two weeks before starting the study. The observer (DWF) remained within the hide during each tracking period and all bird behaviour, locations, departure and retum times and flight directions were noted, Each three hour observation period was started cither ten minutes after establishing DWF in the hide or earlier if the male returned to his court and behaved normally. The observer was in constant radio contact with the radio-tracking team and alerted them as soon a bird left nr arrived ar his court. MEMOIRS OF THE (QUEENSLAND MUSEUM The radio-tracking team remained at least 30m distant from the courts of the radio-tagged birds to minimize disturbance to the court owner or to other Tooth-bills in the area. The tracking team consisted of GJM, with the radio antenna and receiver, who followed the bird’s signal and marked destinations, and CBF who attempted to maintain visual contact with the bird and in- formed GJM of its location and behaviour in the canopy and of the compass bearing of signals, Males were tracked using a Telonics (U.S.A.) Tr-2 receiver. To improve reception at close range and direction finding ability, an enhanced manual gain control was fitted to the reciever, A three element, folding hand held Yagi antenna (Custom Electronics, U.S.A.) was used for direc- tion finding. Use of a small single earphone as- sisted communication between observer and tracker, this was necessary because many forays resulted in a dash of up to 400m to determine the bird's destination. Once a male had been followed to a destination, the tree was tagged and, ata later date, the posi- tions of all destination trees were mapped in relation to the pre-established grid. On a few necasions males flew beyond the limits of the study site; in these cases the locations of destina- tion trees were plotted by tape and compass. Signal strength and consistency were used as a guide to bird activity and if the bird could not be sighted, its position was determined by triangula- tion, If the signal was strong and consistent, this usually meant the bird was stationary. A bearing was then taken from the nearest grid reference point and a second bearing taken at the next gnd reference point to obtain a fix. To test the ac- curacy of this method, calibration transmitters were placed in trees and the resulting fixes were found to be within 5m of the actual transmitter position, A highly variable signal, in both strength and consistency, indicated the bird was moving. In these cases the signal was followed until a strong and consistent signal was obtained and the process wis then repeated, Tf foraging, a strong variahle signal was obtained as the bird would change its orientation whilc moving between branches, The sound of falling debris or fruit was often a good indicator of approximate bird loca- tion within the forest canopy. If interacting with other individuals, calls and chase flights were ofien heard. Establishing a final destination using radio-as- sisted surveillance usually resulted in tracker and observer reaching a destination some lime after ACTIVITIES OF THE MALE TOOTH-BILLED BOWERBIRD 149 FIG. 1. Minimum Convex Polygon home range areas (n = 636 fixes*) for four male Tooth-billed Bowerbirds, based upon their adjacent courts, during the 1989 display season. Numbered stars = male’s court location and number. Solid dots = sightings of males in tree canopy.* male 23 = 202 fixes; 33 = 148 fixes; 35 = 122 fixes; 37 = 164. the bird. This problem increased the further the male travelled from his court. In addition, male Tooth-bills are well camouflaged and rely on cryptic habits to avoid predation (Frith & Frith, 1993, 1994). For these reasons it was often dif- ficult and sometimes impossible to establish either the final destination or the activities of a bird, leaving the reason for many forays by a bird as unknown, and destinations as approximate areas only. In this paper absences from courts are classed broadly as foraging, non-foraging and unknown activities. Foraging absences were those primari- ly concerned with visits to food trees where fruits were eaten but sometimes brief non-foraging ac- tivities were observed before or after feeding. Non-foraging absences did not include visits to fruiting trees or, the the best of our knowledge, feeding. Foraging and non-foraging activities were either observed directly or inferred from characteristics of the destination of the bird, the radio signal or sounds from the canopy. Direct return flights from a destination tree to a court were determined by the rapid loss of a signal and the subsequent recording of the male at the court by DWF in the hide. Many home range estimators are available for home range analysis (Mohr, 1947; Southwood, 1978; Anderson, 1982; Worton, 1989). Follow- ing animals using focal sampling results in autocorrelation of locations (i.e. non-independeni fixes). This may create problems in the analysis of radio-tracking data. However, several recent studies have found that home range estimates based on independent versus non-independent locations are statistically indistinguishable (Anderson & Rongstad, 1989; Gese et al., 1990). As most fixes obtained during this study were MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1. Three-way fixed factor Analysis of Variance * examining the Variance using arcstn square root effect of month, time period and individual male on home range es- transformed proportions: F=6.36, temates for four male Tooth-billed Bowerbirds during peak activity of gf — 3, P = 0,0164). The greatest the 1989 display season. home range overlap was observed betwecn males whose courts were Treatment af F Month g 142 Time period 3 1.26 Individual male 3 2.31 Month x Time period 6 0.22 Time period x Month E] 0.36 Month w Time period x Bird 16 Total 33 * = values obtained using Minimum Conyax Polygon method non-random the Minimum Convex Polygon (Mohr, 1947; Southwood, 1978) estimator was used to estimate home range size. This method is sufficiently robust to deal with non-independence of locations. Home range and home range overlap were estimated using the commercial package Ranges IV. Most frequency distributions of time and space use about a court by the male owner were highly skewed towards the court; therefore the median was chosen as the best indicator of the central place tendency of the data, rather than mean (Zar, 1979), RESULTS HOME RANGE The mean home range area for four male Tooth- bills over the 1989 display season was 9.5ha (range 6.9-12.5) as determined from a total of 636 fixes obtained away from courts (Fig, 1), In a three way fixed factor analysis of variance examining the effect of month, daytime period and individual on the home range estimate, there were no significant differences (Table 1), Mean overlap of the four home ranges was 50% with a range from 23-80% (Fig. 1, Table 2). There was a significant difference in the amount of overlap observed between individual males and their nearest neighbours (one way Analysis of TABLE 2. Percent overlap of the home ranges of four radio-tagged male Tooth-billed Bowerbirds away from their courts during peak activity of the 1989 display season, Male 5 a3 35 37 2 - 46.0 za 454 x 30.7 aa 547 36 226 Tes ‘ 417 ET ME 786 52 p Closest together (Fig. 1, Table 2). The median distance males travelled from their courts for all 0.263 an 0320 activities was 59m, with a mini- bess mum of 8m and maximum of os8 391m (Table 3). Minimum distan- ces occurred when males stole leaf decorations from a neighbour's court or chased off conspecifics u short distance from their own court. Maximum distances oc- curred when males flew to a creek to bathe, usually in the middle of the day. The nearest significant body of water was located in a gully about 300m from the courts (Fig. 1), TIME UTILISATION Males spent an average of 64% (range 56-73%) of total time (0600-1800h) attending courts during peak activity of the 1989 display season. Up to 3% of this time present males were detected above their courts by radio signals, but remained unseen or unheard by the observer in the court hide (see Frith & Frith 1994: table 3). In a three way fixed factor analysis of variance examining the effects of month, daytime period and in- dividual male on the proportion of time spent at the court during 1989, there were no significant differences (Table 4). Court attendance levels and time-budgeted activities at courts by the four males during different months (October-Decem- ber) and during different times (0600-1800h) of the day during this season are reported elsewhere (Frith & Frith, 1994). The results of radio-track- ing at courts at night revealed that each male Tooth-bill roosted above his court. Males spent an average ot 36% (range 27-44%) of total time (0600-1800h) absent from their courts during peak activity of the 1989 display season, with a mean duration per absence of TABLE 3, Distances (m) of all movements away from courts by four male Tooth-billed Bowerbirds during peak activity of the 1989 display season. Male Number Median Minimum Marium ot fixes distanse distance distance 23 202 50.6 272 334.4 33 148 58.6 13.9 370.7 a5 122 Bba 43 ana 37 164 404 az Abie Total 836 59.4 a mn a ——— — — o ACTIVITIES OF THE MALE TOOTH-BILLED BOWERBIRD 151 TABLE 4. Three-way fixed factor Analysis of Variance * examining the ACTIVITIES AWAY FROM effect of month, time activity of the 1989 display season. period and individual male on proportion of time COURTS four male Tooth-billed Bowerbirds spent at their courts during peak Of 197 total absences from courts 119 were foraging absen- ces (60%), 35 non-foraging ab- Trusten W F: P sences (18%) and 43 absences 7 | with unknown activities (22%), a FU = as Seve, representing 82.8, 6.8 and 10.4% Individual male 3 1.06 osei8 Of total time absent respectively Tio veriod we Toh g ta DW (Table 5). A foraging absence Month x Time period X Bird 16 averaged 17,9mins (range 11 4- Toni “ 24), a non-foraging absence averaged 5mins (range 2.9-13) *< using arcsin square root transformed proportion data, and an absence with unknown ac- livities 6.3mins (range 5.1-6.9, 13.1mins (range 8.4-17.7). The proportion of Table 5). total time males were absent and the mean dura- tion of absences were lower during November than during October or December. Males spent less time at courts during the middle of the day between 0900-1500h (Table 5). During the first hour (0600-0700h) of the first October tracking period male 33 cleared a small subsidiary court some 40m away from his primary one. Because this unusual behaviour may have been caused by banding, transmitter attach- ment or both the records from this hour have been excluded from analyses. During subsequent Oc- tober tracking periods this male occasionally flew over, and perched briefly above, the rudimentary court (<3% of total time absent from his estab- lished court), Male 33 spent more time absent from his court than the other three males and Male 37 spent less time absent (Table 5), but differen- ces between the four males were not significant (x= 3.78, P > 0.02). During foraging absences the birds were seen in destination food trees for 67% of the time; the remaining time being spent en route to or from the court (30%) or performing non-foraging ac- tivities (3%). Sixty six of the 119 foraging absen- ces were to known food trees. The median foraging distances travelled by three males to these trees ranged from 19-42m, but male 37 flew much further to forage, with a median distance of 119m. During a foraging absence males usually flew directly from the court to a food tree. On eight occasions, however, a court-owning male chased a conspecific from his court before continuing on to a food tree (sec below). Males also occasional- ly flew directly towards a creek (before or afler feeding) where it was assumed they went to bathe or drink, but trackers were only able to visually confirm this twice, Most times (86% of foraging absences) the bird flew directly back to the court TABLE 5. Activities performed by four individual male Tooth-billed Bowerbirds during absences from their courts over peak activity of display season. ABSENCES; FORAGING NON-FORAGING UNKNOWN ACTIVITIES TOTAL Male Hours Number % of Mean Number % cf Maan Number Tê of Mean Number 3% ot Mean of radjo- at tme mins @ of tima mins @ of time mins @ of totel tims mins @ tracking absences absent absence absences absent absence absences absent absence absances sbeent sbaence Mele a 23 36 43 64.8 181 5 78 13.0 12 7.5 54 60 125 ye 33 23 26 90.8 212 5 a7 48 5 6.6 68 36 16.9 tat 35 24 Vi 89.8 ao 7 52 54 4 40 58 2 wi 382 7 36 29 66.3 134 18 Be 29 z 25.9 68 be 6a mb Month October 47 a9 60.8 199 15 69 “7 16 108 ae 72 13.4 341 November 35 35 784 139 12 89 42 16 124 48 a4 a7 266 December 38 45 87.5 183 4 a5 6s 1? BO 76 LU 16.5 480 Time 0600-0900 2 32 765 11.4 16 10.3 31 10 132 63 5e uz 27,4 0900-1200 30 28 65.1 220 4 34 £3 3 11.6 64 46 183 404 1200-1500 30 29 80.1 20,9 5 a3 105 15 11.3 5T 0 151 A9 1500-1800 80 ao 77.8 182 b 62 43 S &3 75 4 144 us Total/means 119 119 52.8 7s z ae so «a 04 43 187 134 me 152 but on 15 occasions (14% of foraging absences) a leaf decoration was collected during the return trip. It is possible that undetected activities oc- curred during flights between courts and food irees, Of the 35 non-foraging absences 12 entailed collection of a leaf decoration from a tree (3 leaves) or from another court (9 leaves), 12 were visits to the forest canopy above a neighbouring courts, and 11 involyed the male court-owner chasing a conspecific and then returning immedi- ately to his own court (see below), Theft involved a male flying directly to a neighbouring court. quickly and quietly stealing a leaf from it, and returning directly to his own court. Leaf theft represented 0.5% of total time absent from the cour, with a mean duration of 1.5mins per theft The 12 visits to a neighbouring court were brief and involved the court-owner flying directly to, and perching quietly above, his neighbour's court for a short period (mean duration = 3.3mins), possibly with the intention of stealing a leaf. This behaviour was performed by all four males. Males were seen to chase conspecifics away from the court on 19 occasions flying on toa food tree after eight such chases and returning directly back to the court after 11 of them. Three chases were in pursuit of a presumed female (having previously been displayed to). It is possible other chases were after a neighbouring male, On onc occasion a leaf was stolen by a marked, neigh- bouring male just as Male 37 returned to his court. The latter male immediately chased the thief to the thief’s court, about 70m away, where he stayed briefly before retuming to his own court withoul a leaf. He was, however, subsequently seen to steal from that particular neighbour's court Interactions between conspecifics were ob- served away from the court. On four occasions a male left his court and flew to where other Tooth- bills were heard displaying, either above or on another a court. On one occasion Male 35 was seen calling and displaying lo an assumed female near a creck that was about 300)m from his court. On two occasions two birds were seen facing each other in a food tree, face to face, with conspicous- ly sleeked plumage and agitated vocalisations. DISCUSSION Bechler & Pructt-Jones (1983) reviewed avail- able information concerning the spatial disper- sion of adult males of nine polygynous bird of paradise species (Paradisacsidac) from New MEMOIRS OF THE QUEENSLAND MUSEUM Guinea rainforests. These species were known or presumed to inyolve promiscuous males. Males of species with a predominantly arthropod diet were found to be territorial while males of predominant frugivorous species were non-ter- ntorial. Lack of territoriality in predominantly frugivorous passerine species is considered to result from the fact that tropical fruits are economically undefendable (Lill, 1976; Bechler, 1989; Beehler & Pruett-Jones, 1983). During the 1989 display season the mean home range size of male Tooth-bills was 9 5ha and the median distance travelled from courts for all ac- tivities was 59m, Comparable data on the breed- ing season home range of other tropical frugivorous birds which display at focal sites are scarce. Pruett-Joncs & Pructt-Jones (1988) reported that the home range of one adult male Blue Bird of Paradise, Paradisaea rudolphi, was 4.Tha over the season, about half the size ob- served in male Tooth-bills. Display trees of the former species are much more widely spaced than are the courts of Tooth-bills (220m vs 50m respectively). Blue Birds of Paradise are nol as exclusively frugivorous through the breeding season as are Tooth-bills (cf. Beehler & Pruett- Jones, 1983) and this, together with a possible greater abundance of food near display sites, may account for differences in home range sizes be- tween these similar sized species. It is noteworthy that the all-purpose territory of three male Superb Birds of Paradise, Lophorina superba, a species with only 24% fruit in its dict, averaged 1.5ha (range 1.2-1.7) (Beehler & Pruett-Jones, 1983). The availability of fruit in time and space may therefore have major effects on both male disper- sion and home range. Male Tooth-bills do not move far from their courts and, as would be expected, males at an exploded Ick exhibit considerable overlap (mean = 50%) of home ranges. Those with closer courts show the greatest overlap, and males from dif- ferent courts were observed to bathe in the same portions of creeks and 1o feed in the same trees. Although distances incurred in travelling to bath- ing and drinking locations, ur those associated with leaf thefls, are likely to remain constant across years this will not be truce of distances Iravelled to and from feeding locations. These latter distances will vary between years as dif- ferent individual trees ur species of tree fruit, Seasonal environmental differences, in particular fruit crop size, affect the length of Tooth-bill display seasons and the proportion of time that males attend courts (Moore, 1991; Frith & Frith, ACTIVITIES OF THE MALE TOOTH-BILLED BOWERBIRD 1994), Fruit crop was poor during the 1989 dis- play season (Moore, 1991), Consequently, males spent a grealer proportion (36%) of their time absent from courts, to feed, during the 1989 season than during the 1980 season (21%) when fruit was abundant (Frith & Frith, 1994), During the exceptionally dry 1979 season an extremely poor fruit crop resulted in males spending even more time (48%) away from courts (Frith & Frith, 1994), All the observed movements away from courts appeared to entail direct purposeful flights to food, water or leaf (court decoration) resources, Activities such as interactions with conspecifics or the collection of leaf decorations that were observed away from courts, during both foraging and non-foraging absences, were extremely bricf (Table 5). Most absences were for foraging, rep- resenting 8390 of total time absent with a mean duration of 18 mins (Table 5), Moore (1993) found that Tooth-bills tended to forage more in food trees closer to the court than those further away and as a result shorter foraging absences were more frequent than longer ones. Median distance travelled from courts exclusively for foraging at food trees was 32m (median range 27-119). Male 37 flew much further than the other three males to forage. often visiting fruiting trees some 380m to the east of his court that were not visited by the other males studied (Fig. 1). While males other than those of this study doubt- less fed in these trees the observation suggests the possibility that males on an exploded lek may travel greater distances to utilise rich food resour- ces where competition may be less. It is possible, however, that Male 37 also fed in this area duc to the local accessibility of the creek (see Fig.1). Donaghey (1981) presented the only com- parable data, He found that five rainforest-dwell- ing male Satin Bowerbirds, Prilonorhynchus violaceus, mostly foraged for food within 50m of their bowers (83% of 83 foraging records) during the breeding season of October-December. He obtained almost identical results for four wood- land-dwelling males (82% of 273 foraging records within 50m of bowers). His Satin Buwerbird's bowers did not form leks but were dispersed linearly along forest edges at a mean inter-bower distance of 312m (Donaghey, 1981). Male Tooth-bills, like male Satin and other bowerbirds, must remain close to their court if they are to successfully display and attract females to mate (Frith & Frith, 1993) as do males of other, mostly lekking, passerines (Snow, 1962; Lill 19744, bh), Thus, irrespective of the size of a 153 meal caten or the distance travelled to the food, males almost invanably return directly to their court in order to defend it from leaf stealing rivals and to call and display. In effect, male Tooth-bills attempt to maximise their time at their court (Moore, 1991), They can achieve this, however. only by utilising a food resource that is freely available and unable to be defended, namely, fleshy fruits. Snow (1976) considered a predominantly frugivorous diet one of the main pre-conditions to a mating system lacking pair bond and biparental brood care in which males spend much time attending and defending a tradi- tional courtship area. ACKNOWLEDGEMENTS CBF and DWF thank GJM and the Department of Zoology, James Cook University of North Queensland for use of radio-tracking equipment. GJM thanks Professor Rhondda Jones and Dr Annemane Watt lor reviewing initial drafts of his thesis manuscript and the Commonwealth Postgraduate Research Awards and the Australian Research Council for financial assis- tance for this project which formed part of his PhD study on seed dispersal by male Tooth-billed Bowerbirds. Glen Ingram and Peter Dwyer kind- ly provided constructive criticism of a draft that improved this contribution. LITERATURE CITED ANDERSON, D.J, 1982, The home ninge: 4 new non parametric estimation technique, Ecology 63:103- 12 ANDERSON, D.E. & RONGSTAD, (0.3. 1989, Home range estimates of red-tailed hawks based un rais- dom and systematic relocations, Journal of Wildlife Mangement 53:802-807 BEEHLER, B.M. 1989. The birds of paradise. Scientific Amenean 261:117-123. BEEHLER, B. M. & PRUETT-JONES, S. G.1983 Display dispersion and diet of birds of paradise: a comparison of nine species, Behavioural Ecol ogy and Sociobiology 13:229-238, DONAGHEY, R. H, 1981. “The ecology and evolution of bowerberd mating systems”. (Unpublished PhD Thesis; Monash University, Australia). FRITH, C.B. & FRITH, D.W. 1985. Parental care and investment in the Tooth-billed Bowerbird Scenopoeeles. dentirostris. (Pulonorhynchidae). Australian Bird Watcher 11:103-113. 1993. Courtship display of the Tooth-billed Bower- bird Scenopoeetes dentîrosiris,; its behavioural and systematic significance. Emu 93;129-136, 1994, Courts and seasonal activities at them by the male Tooth-billed Bowerbird, Scenapoeeres 154 dentirostris (Ptilonorhynchidae). Memoirs of the Queensland Museum 37(1):121-145. IN PRESS. Court site constancy, dispersion, male survival and court ownership in the male Tooth- billed Bowerbird, Scenopoeetes dentirostris (Ptilonorhynchidae). Emu 95. GESE, E.M., ANDERSON, D.E. & RONGSTAD, O.J. 1990. Determining home-range size of resident coyotes from point and seguential locations. Jour- nal of Wildlife Management 4:501-506. GILLIARD, E.T. 1969. 'Birds of paradise and bowerbirds'. (Widenfeld & Nicolson:London). KENWARD, R.E. 1978. Radio transmitters tail mounted on hawks. Ornis Scandinavia 9:220-223. LILL, A. 1974a. Social organization and space utiliza- tion in the lek-forming White-bearded Manakin, M. manacus trinitalis Hartert. Zeitschrift Tierpsychologie 36: 513-530. 1974b. Sexual behaviour in the lek-forming White- bearded Manakin (M. manacus _ trinitalis Hartert). Zeitschrift Tierpsychologie 36:1-36. 1976. Lek behaviour in the Golden-headed Manakin, Pipra erythrocephala in Trinidad (West Indies). Advances in Ethology 18:1-84. MOHR, C.O. 1947. Table of equivalent populations of North American small mammals. American Mid- land Naturalist 37:223-249. MEMOIRS OF THE OUEENSLAND MUSEUM MOORE, G.J. 1991, ‘Seed dispersal by male Tooth- billed Bowerbirds Scenopoeetes dentirostris (Ptilonorhynchidae), in north-east Oueensland rainforest: Processes and conseguences'. (Un- published Ph D.Thesis: James Cook University of North Oueensland, Australia). PRUETT-JONES, S.G. & PRUETT-JONES, M.A. 1988. A promiscuous mating system in the Blue Bird of Paradise Paradisaea rudolphi. Ibis 130:373-377. SNOW, D.W. 1962. A field study of the Black and White Manakin Manacus manacus in Trinidad. Zoologica 47; 65-104. 1976, ‘The web of adaptation’. (Collins:London). 1982. ‘The cotingas'. (British Museum of Natural History& Oxford University Press:London). SOUTHWOOD, T.R.E. 1978. 'Ecological methods: with particular refemce to the study of insect populations'. (Chapman & Hall:London). WORTON, B. J. 1989. Kernel methods for estimating the utilisation distributions in home-range studies. Ecology 70:164-168. ZAR, J. H. 1979. ‘Biostatistical analysis’. (Prentice- Hall:USA). REDISCOVERED PSEUDOSCORPION TYPE MATERIAL DESCRIBED BY BEIER FROM SOUTHEASTERN QUEENSLAND (ARACHNIDA: PSEUDOSCORPIONIDA) MARK S. HARVEY Harvey, M.S. 1994 12 01: Rediscovered pseudoscorpion type material described by Beier from southeastern Queensland (Arachnida: Pseudoscorpionida). Memoirs of the Queensland Museum 37(1):155-156. Brisbane. ISSN 0079-8835. All or part of the type series of f1ve pseudoscorpion species described by Beier (1969) from southeastern Oucensland have been considered lost for some time. They have been recently found in the Department of Entomology, University of Queensland and transferred 10 the Queensland Museum. The specimens are documented. [_] Arachnida, Pyeudoseorpionida, Tridenchthoniidae, Chthoniidae, type material. Mark S. Harvey, Western Australian Museum, Francis Street, Perth, Western Australia 6000, Australia; 15 December 1993. Beier (1969) recorded several species of pseu- doscorpions from southeastern Queensland, based upon material sent to him by Mr J.B. Wil- liams. Five were new and a further two species were recorded from southeastern Queensland for the first time. Beier claimed to have returned most of the material to Mr Williams, then at Monash University, Melbourne, while retaining several specimens of two species in the collections of the Naturhistorisches Museum, Vienna. Despite ef- forts to trace these specimens, the material ap- peared to be lost (Harvey, 1981, 1985, 1989). During a recent visit to the Department of Entomology, University of Queensland, Bris- bane, I located the missing vials, and now take this opportunity to comment on the condition and status of the types. All of the type material has been transferred to the Queensland Museum, Brisbane (QM). Family TRIDENCHTHONIIDAE Heterolophus australicus Beicr, 1969 Heterolophus australicus Beier, 1969: 171-172, fig. 1. MATERIAL EXAMINED HoLoTYPE: 9, Mt Tamborine, Joalah National Park, Queensland, 28°56'S, 153712'E, berlesate, rainforest Jeaf litter, 16 Aug 1966, B. Cantrell, QMS21333. PARATYPE: l trilonymph, same dala as holotype, QMS21334. REMARKS The specimens are in fair condition, and the right chela is detached from the holotype. A recurrent typographical error by Beier (1969) referred to Mt Tamborine as “Mt Jamborine” and Joalah National Park as “Toalah National Park”, Anaulacodithella australica Beier, 1969 Anaulacodithella australica Beier, 1969: 172-174, fig. 9 é. MATERIAL EXAMINED HOLOTYPE; d Lamington National Park, Queensland, 2712'S, 153°10'E, extracted from leaf Jitter, 6 Mar 1965, B. Cantrell, OMS21335. ALLOTYPE: 2, Mt Glorious, Queensland, 27°20°S, 152°46°E, 11 Jul 1965, berlesate, rainforest leaf litter B. Cantrell, OMS21336. PARATYPES: 1 4,1 9, Landsborough, Queensland, 26°48°S, 152°58°E, Australia. leaf mould, mixed forest, 16 Jul 1965, F.A. Perkins, OMS21337; 4 specimens, Mi Clunie, east spur, Queensland, 28/20'S, 152°31°E, leaf mould, rainforest, 5 Apr 1953, T.E. Woodward, OMS21338; 16 specimens, Mt Tam- borine, Joalah National Park, Queensland, 28°56'S, 152°12°E, berlesaté, 16 Aug 1966, B. Cantrell, OMS21339. REMARKS The holotype and allotype are in fair condition, but the remaining specimens are extremely bleached and fragmented. The left chela of ihe holotype has been derached; the right chela of the allotype has been detached and subsequently lost. Family CHTHONIIDAE Morikawia gueenslandica Beser, 1909 Morikawia queenslandica Beier, 1969: 174-175, fig. 3. MATERIAL EXAMINED SYNTYPES: 5 d,3 9, Mt Tamborine, Joalah National Park, Queensland, 28/56'S, 153'12'E, berlese, rain- forest leaf litter, 16 Aug 1966, B. Cantrell, OMS21341. 156 REMARKS This species was transferred to the genus Tyrannochthonius, based on examination of the remaining syntypes lodged in Naturhistorisches Museum, Vienna (Harvey, 1989). Morikawia semihorrida Beier, 1969 Morikawia semihorrida Beier, 1969: 176-177, fig, 4. MATERIAL EXAMINED HOLOTYPE: 2, Mt Nebo, Queensland, 27 23'S, 152°47°E, rainforest leaf litter, 28 Mar 1967, J.B. Wil- liams, OMS21340. REMARKS The recent transfer of this species to Tyran- nochthonius (Harvey, 1989) based solely on the original description is confirmed by examination of the holotype. The right chela has been detached from the pedipalp. Pseudotyrannochthonius queenslandicus Beier, 1969 Pseudotyrannachthonius queenslandicus Beier, 1969: 177-178, fig. 5. MATERIAL EXAMINED HOLOTYPE: â, Mt Tamborine, Joalah National Park, Queensland, 28°56’S, 153°12'E, 16 Aug 1966, B. Cantrell, berlesc, OMS21342. PARATYPE: 1 tritonymph?, same data as holotype, OMS21343, REMARKS The specimens are in fair condition. Austrochthonius australis Hoff, 195] REMARKS Bejer (1969) recorded this species from southeastern Queensland, based on one male from Mt Nebo, (berlesate, rainforest leaf litter, 28 Mar 1967, J.B. Williams), which is now lodged in the Department of Entomology, University of MEMOIRS OF THE QUEENSLAND MUSEUM Qeensland. This specimen conforms to recent descriptions of the species (Kennedy, 1989). Family OLPIIDAE Xenolpium longiventer (L. Koch, 1885) REMARKS Beier (1969) recorded this species from southeastem Queensland based on a specimen from Mt Nebo (rainforest leaf litter, 28 Mar 1967, J.B.Williams), which is now lodged in the Department of Entomology, University of Queensland. The specimen is clearly not con- generic with the type of X. longiventer (BMNH), and in fact represents an undescribed species of Protogarypinus. Members of this genus are abun- dant in southeastern Australia, and many new species await description. ACKNOWLEDGEMENTS Ms Margaret Schneider very kindly allowed access to the collections of the Department of Entomology, University of Queensland. LITERATURE CITED BEIER, M. 1969, Neve Pseudoskorpione aus Australien. Annalen des Naturhistorischen Museums in Wien 73: 171-187, HARVEY, M.S. 1981. A checklist of the Australian Pseudoscorpionida, Bulletin of the British Arach- nological Society 5: 237-252. 1985. Pseudoscorpionida, Pp,126-155. In Walton, D.W, (ed.), ‘Zoological catalogue of Australia’. Vol. 3. (Australian Government Publishing Ser- vice: Canberra), 1989. Two new cavernicolous chthoniids from Australia, with notes on the genene placement of the south-western Pacific species attributed to the genera Paraliochthonius Beier and Morikawia Chamberlin (Pseudoscorpionida; Chthoniidae), Bulletin of the British Arachnological Society 8; 21-29. KENNEDY, C.M.A, 1989. Redescription of Austroch- thonius australis Hoff (Chthoniidae: Pseudoscor- pionida). Proceedings of the Linnean Society of New South Wales 111; 233-240, SOME EARTHWORMS FROM THE WET TROPICS AND FROM BUNY A MOUN- TAINS, QUEENSLAND (MEGASCOLECIDAE: OLIGOCHAETA) B.G.M. JAMIESON Jamieson, B.G.M. 1994 12 01: Some earthworms from the Wet Tropics and from Bunya Mountains, Queensland (Megascolecidae: Oligochaeta), Memoirs of the Queensland Museum 37(1): 157-180. Brisbane. ISSN 0079-8835, A systematic study, supported by acladistic analysis, indicates that North Queensland species of Diporechaeta warrant separate generic status, as Terrisswalkerius gen. nov. Fletcherodrilus gen. nov. represents an apomorphic clade of this assemblage which ts, nevertheless, retained as a separate genus. The Bunya Mountains isolate of Diporochaeta merits separate generic status, as Hiatidriluy bunya, and Cryptodrilus semicinctus, from Grafton, on the Clarence River, New South Wales, is congeneric with it. The residue of Diporochaera ìs a paraphyletic and polyphyletic entity reguiring further resolution. The cladistic analysis also investigates wider relationships of the above genera with the remainder of the Perionychini. In the analysis the type-species of Diporochaeta, D. intermedia, Is separated by other genera from those Diporochaeta species here assigned to Terrisswalkerius and, indeed, from all other included Diporochaeta species. The analysis suggests that Perionychella, currently subsumed in Diporochaeta, is the sister-taxon of the Ter- risswalkerius-Fletcherodrilus assemblage. The phylogram is not inconsistent with the generic status of the remaining perionychin genera Graliophilus, Heteroporodrilus, Paraplutellus, Perionychella, Pinguidrilus, Plutellus, Pseudoperichaeta, Simsia, and Wood- wardiella but significant autapomorphies remain to be established for some of these, Within Terrisswalkerius the most plesiomorphic species appears to be T, blounti. T. atavius, is the most derived species and forms the sister-taxon of Fletcherodrilus, Pairs of sister-species are! T. grandis and T, terraereginae, T. millaamillaa and T, mcdonaldi sp. nov., T. ather- tonensis and T. oculatus, T. kuranda and T, canaliculatus, and T, nashi and T, liber sp. nov, A monophyletic, though weakly defined, species group contains T. athertonensis, T. oculatus, T. canaliculatus, T, kuranda, T, crateris, T. nashi and T, liber, The new species T, covacevichae and Fletcherodrilus menurus are also described,[_|Terrisswalkerius gen nova Hiatidrilus gen.nov,, Fletcherodrilus, Diporochaeta, Perionychini, cladistics. B.G.M. Jamieson, Zoology Department, University of Queensland, Brisbane , Queensland 4072, Australia; 28 August 1994, Diporochaeta was erected by Beddard (1890) with, as its Lype-species, the New Zealand species Diperochaeta (=Perichaeia) intermedia Bed- dard, 1889. Prior to the present work, Diporochaeta contained 78 Australian species, These species had been initially placed, on erec- tion, in some eight genera in addition to Diporochaeta. Of these, five (Cryptodrilus, Megascolex, Megascolides, Perionyx, Plutellus) are still recognised as dislinct genera, two (Perionychella and Vesiculodrilus) have been suhsumed in Diporochaeta (see Jamieson, 1976b) and one (Perichaeta) in Megasolex (see Michaelsen, 1900). Here evidence will be presented, in a cladistic analysis, in support of removal on “intuitive' grounds, of 16 species from North Queensland from Diporochaeta and their transfer to a new genus, Terrisswalkerius, and for transfer of two species to a further new genus, Hiatidrilus. The number of Australian species of Diporechaefa is thus reduced to 60. Three new species are added to Terrisswalkerius. The status and phylogenetic affinities of Fleicherodrilus will also be investigated. Jamieson (1976b) observed that little other than the unpaired genital pores warranted separation of Fletcherodrilus from Diporochaeta, that in- clusion in the latter genus as a subgenus would be sustainable, and that Fletcherodrilus may be regarded as the apomorph sister-group of Diporochaeia, A new species is added to Fletcherodrilus. Erection of a distinct genus for north Queensland species of Diporochaeta and the apomorphic status of Fletcherodrilus have been independently suggested by Dyne & Wal- lace (1994). The status of the predominantly Western Australian genus Graliophilus Jamieson, 19716, was rendered uncertain by addition of North Queensland species to Diporochaeta by Jamieson (1976b) which blurred the distinction between the two genera. Graliophilus was 158 retained by Jamieson (1974a) on the grounds, inter alia, that in Western Australian species the number of spermathecae was less (2 or 3 pairs) than the five pairs usual in Diporochaeta and that the avesiculate nephridia and totally Jumbricin condition of the setae were constant whereas the combination of these characters was known in only one species of Diporochaetg from eastern Australia (Tasmania). It was concluded, how- ever, that the low phenejic resemblance between Diperochaeta and Graliophilus demonstrated by Wallace (1972) and the very distinct morphology of some Graliophilus species suggested that this genus should be retained. The present study ex- amines cladistic relationships of Graliophilus to Diperochaeta s. lat. and includes in the analysis the type-species of all numed genera of the Perionychini, viz, the 13 genera Diporochuera, Fletcherodrilus, Graliophilus, Heteroporadrilus, Miatidrilus, Paraplutellus, Perionychella, Pin- guidrilus, Plutelius, Pseudoperichaeta, Simsia, Terrisswalkerius, and Woodwardiella. Retention of Perionychella, for forms lacking nephridial bladders (and possibly with racemose prostates, as in the type-species) as a subgenus of Diporochaeta deserves consideration but was not upheld (Jamieson, 1976b) as the distinction be- tween an avesiculate and a vesiculate group of species had become increasingly blurred, Thus Tasmanian Diporochaeta mortoni has avesicu- late and, typically, vesiculate individuals (Jamieson, 1974a), D. grandis has vesiculate and avesiculate populations, or even segments in the same individual and in D. terraereginae (both are Queensland species) nephnidial ducts are transi- tional between the two conditions. Perhaps more Significantly, in an unpublished taxonometric (numerical) study vesiculate and avesiculate spege formed mixed groups, especially where all nephridial characters are omitted, It is noteworthy that Michaelsen, who ignored the presence or absence of bladders, considered the avesiculate D. atavia to be merely a form of thc vesiculate D, athertonensis, a decision indicating the closeness of the entities with the antithetic conditions, notwithstanding the view (Jamieson, 1976b) that they are distinct species. Unless otherwise stated material studied is hotised in the Queensland Museum (OM). Terrisswalkerins gen. nov. DIAGNOSIS Setae numerous (>14) per segment. Genital markings other than porophores bearing the MEMOIRS OF THE QUEENSLAND MUSEUM female, male and spermathecal pores, absent. A pair of combined pores of vasa deferentia and tubular or tubuloracemose prostates on XVIII. Last hearts in XI or XIII. Gizzard in V or VI (or VII?) (well developed); intestinal caeca and typh- losole absent. Extramural calciferous glands ab- sent. Nephridia stomate exonephric holonephridia with or without bladders; their pores in straight or simuous lines but never with regular alternation. Spermathecae 1 to 5 pairs, rarely unpaired midventral, pre-testicular, diver- ticulate; diverticulum usually single, uniloculate; rarely double, rarely multiloculate, DESCRIPTION Terrestrial worms. Form circular in crass sec- tion or less commonly slightly depressed dor- soventrally; anus terminal. With or without pigmentation, Prostomium proepilobous to tanylobous. Body usually with a dorsal groove (canalicula) in part or the whole of its length. Dorsal pores present; in 3/4 - 6/7, usually 5/6. Setae perichaetine throughout. Nephridia in single lateral series throughout, or in irregularly sinuous series, varying from far dorsally to far ventrally but never showing regular alternation. Posterior limit of the clitellum shortly anterior to the male pores or including these. A pair of male pores on XVIII, each pore the combined opening of a prostate gland and the corresponding vasa deferentia. Female pores paired on XIV. Sper- mathecal pores | 10 5 pairs, rarely unpaired mid- ventral, between intersegmental furrows 4/5-8/9, rarely shortly behind the intersegments, Dorsal blood vessel single, continuous onto the pharynx; last hearts in XI or, rarely, XIII; hearts in X posteriorly latero-oesophageal, each receiv- ing a conneclive from the supra-oesophageal ves- sel and a lesser connective (sometimes absent?) from the dorsal vessel. Supra-oesophageal vessel restricted 1o the post-gizzard oesophagus, moderately to well developed. Subneural blood vessel absent, Gizzard in V or, more frequently, VI (or VII?) (well developed). Extramural cal- eifernus glands nor developed bur oesophagus vascularized and often internally rugose in some segments from VIII to XVI. Intestine commenc- ing in XVI to XIX, most frequently XVII; intes- linal caeca and typhlosole absent. Nephridia simple, exonephric holonephridia throughout; tufted nephridia absent; bladders usually present, with or without diverticula; exceptionally (D. grandis, part.) avesiculate in the forebody and vesiculate in the hindbody. Holandric (testes in X and XI); seminal vesicles two to four pairs, in IX NEW QUEENSLAND EARTHWORMS 159 to XII, exceptionally (T. terraereginae) in VIII (in addition to IX and XII), and often with pseu- dovesiclesin XIII. Metagynous (ovaries and fun- nels in XIII). Prostates tubular or tubuloracemose, never racemose. Penial setae ab- sent. Spermathecae diverticulate; diverticulum usually single, uniloculate; rarely double or mul- tiloculate, DISTRIBUTION The Wet Tropics of eastern North Queensland, from the Paluma Range, 19S., near Townsville, north to the Upper Endeavour River shortly below 15*S. TYPE SPECIES Terrisswalkerius (=Perichaeta) canaliculatus (Fletcher, 1887a); here selected because its anatomy is well known and it is the most widely distributed species of the genus and therefore the most likely to be available for molecular and other studies. ETYMOLOGY Named in memory of Terry Walker. Gender masculine. INCLUDED SPECIES T. atavius (Michaelsen, 1916); T. athertonensis (Michaelsen, 1916); T. barronensis (Fletcher, 1886b); T. blounti (Jamieson, 1976b); T, canaliculatus (Fletcher, 1887a); T. covacevichae sp. nov.; T. crateris (Jamieson, 1976b); T. erici (Michaelsen, 1916); T. grandis (Spencer, 1900); T. Kuranda (Jamieson, 1976b); T. liber sp. nov.; T. mcdonaldi sp. nov.; T. millaamillaa (Jamieson, 1976b); T. montislewisi (Jamieson, 1976b); T. nashi (Jamieson, 1976b); T. oculatus (Jamieson, 1976b); T. phalacrus (Michaelsen, 1916); T. raveni (Jamieson, 1976b); T. terrareginae (Fletcher, 1890). KEY TO SPECIES OF TERRISSWALKERIUS 1. Spermathecal pores 5 pairs, in or shortly behind intersegmental furrows 4/5-8/9. 22.2.0. | 2 Spermathecal pores 1 to 4 pairs (rarely unpaired, midventral), in or shortly behind some or all of intersegmental furrows 5/6-8/9. .....,... 4 Spermathecal pores 5 pairs 2. Spermathecal pores in a lines. Spermathecal duct very short. Last hearts in XIII, T, grandis Spermathecal pores median to a lines. Spermathe- cal duct Jong and slender though shorter than the ampulla. Last hearts in XII 3. Spermathecal pores of 8/9 c, 0.02 body circum- ference apart. Male pores c. 0.03 body circum- ference apart. Seminal vesicles in IX and XII. Nephridial bladders present. .. ... T. atavius Spermathecal pores of 8/9 e. 0.06 body circum- ference apart. Male pores c. 0.1 body circum- ference apart. Seminal vesicles in XI and XH (or absent). Nephridial bladders absent- . . _ , - T. phalacrus 4. Spermathecal pores in 4 intersegments, 5/6-8/9 or 4/5-7/8, paired orsingle. ... ..:. 5 Spermathecal pores 3 pairs or fewer: .. - . 9 Spermathecal pores in 4 intersegments 5. Spermathecal pores in 5/6-8/9, unpaired, mid- ventral. s we ee ee T. medonaldi sp. nav. Spermathecal pores 4 pairs, in 4/5-7/8 or 5/6-8/9 6. Spermathecal pores 4 pairs, in 4/5-7/8 «a .. Ag dicio df lg TW he Jo Je tet [ce T. terrareginae - Spermathecal pores 4 pairs, in 5/6-8/9.. ... 7 7. Male pores median to a lines, their papillae con- Uguous in the midline. Nephridial bladders ab- snt Jn. “y yd A hwi: T: millaamiliaa Male pores between setal lines a and d, their papillae well separated in the midline. Nephridial bladders present... 0. aaa 8 8. Prostomium tanylobous, with wide dorsal tongue. Nephropores in a straight series on each side . - Loved stem arate eu sae T. erici Prostomium epilobous or proepilobous; with a deep middorsal groove which continues to the hind margin of the peristomium. Nephropores in an irregularly sinuous series, varying from far dorsally to far ventrally, on each side... . . . 9. Spermathecal pores 3 pairs, in 4/5-6/7 or in or shortly behind 6/7-8/9 . RR 10 Spermathecal pores 1 or 2 pairs, in some of 5/6- Spermathecal pores 3 pairs 10. Spermathecal pores 3 pairs, in 4/S-6/7 .,., 11 Spermathecal pores 3 pairs, in or shortly behind E a ym Om dn dr dyd 13 11. Spermathecal diverticulum long and tortuous. Spermathecal pores in the vicinity of d to € fines de EE Codd b a A uh 12 Spermathecal diverticulum short and clavate. Spermathecal pores between a and b lines . - T. montislewisi (patt,) 160 12. Spermathecal diverticulum longer than ampulla plus duct. (Spermathecal pores ind lines) . . . fete fe Pe ee YF T. barronensis Spermathecal diverticulum shorter than ampulla plus duct. (Spermathecal pores in d-e lines) . nr WU tor fe ai bath NW T. raveni 13. Male pores very close together, almost con- tiguous midventrally, on a common papilla . , ydu Pa" koane usiye Sinn Ych T. athertonensis Male pores well separated, between setal lines b and f, ona pair of papillae ......... 14 14. Spermathecal diverticulum simple, shortly clavate i SR EE I SY py eb 15 Spermathecal diverticulum composite, consist- ing of as many as 4 parallel conjoined tubes with terminal, knoblike seminal chambers; ap- proximately as long as the spermathec . . . , T. oculat ont irbe