AO Ol a) DENIES of the LINNEAN SOCIETY of NEW SOUTH WALES VOLUME 125 NATURAL HISTORY IN ALL ITS BRANCHES THE LINNEAN SOCIETY OF NEW SOUTH WALES ISSN 0370-047X Founded 1874 Incorporated 1884 The Society exists to promote the cultivation and study of the science of natural history in all its branches. The Society awards research grants each year in the fields of Life Sciences (the Joyce Vickery fund) and Earth Sciences (the Betty Mayne fund), offers annually a Linnean Macleay Fellowship for research, contributes to the stipend of the Linnean Macleay Lecturer in Microbiology at the University of Sydney, and publishes the Proceedings. It holds field excursion and scientific meetings, including the biennial Sir William Macleay Memorial Lecture delivered by a person eminent in some branch of natural science. Membership enquiries should be addressed in the first instance to the Secretary. Candidates for elec- tion to the Society must be recommended by two members. The present annual subscription is $A53.00. The current subscription rate to the Proceedings is set at $480.00 per volume. In recent years a volume consists of a single annual issue. Back issues of all but a few volumes and parts of the Proceedings are available for purchase. Prices are listed on our home page and can also be obtained from the Secretary. OFFICERS AND COUNCIL 2003/2004 President: 1.G. Percival Vice-presidents: R.J. King, K.L. Wilson, A. Ritchie, J.P. Barkas Honorary Treasurer: M.L. Augee Secretary: J-C. Herremans Council: A.E.J. Andrews, M.L. Augee, J.P. Barkas, M.R. Gray, J-C. Herremans, M.A. Humphrey, D. Keith, R.J. King, H.A. Martin, P.M. Martin, J.R. Merrick, M.S. Moulds, D.R. Murray, P.J. Myerscough, I.G. Percival, A. Ritchie, S. Rose, and K.L. Wilson Honorary Editor: M.L. Augee Linnean Macleay Lecturer in Microbiology: P.R. Reeves Auditors: Phil Williams Carbonara The postal address of the Society is: P.O. Box 82, Kingsford NSW 2032, Australia Telephone: (International) 61 2 9662 6196; (Aust) 02 9662 6196 E-mail: linnsoc @acay.com.au Home page: www.acay.com.au/~linnsoc/welcome.html © Linnean Society of New South Wales Cover motif: diving platypus by Marianne Larsen, Wellington NSW. TE NATURAL pre ORY MUSEUM 11 MAY 2004 GED PROCEEDINGS ; MIGRARY of the LINNEAN SOCIETY NEW SOUTH WALES EXCHA GENERAL For information about the Linnean Society of New South Wales, its publications and activities, see the Society’s homepage http://www.acay.com.au/~linnsoc/welcome.htm VOLUME 125 February 2004 mai W boon widte Fest. nso» froNc EDITORIAL This volume consists of three parts. The first part contains general contributions. The second part contains research papers and review papers arising from the symposium “Monotreme IIT” held by the Linnean Society of NSW and the Australian Mammal Society at the University of Sydney in July 2003. The third section contains book reviews and an obituary to Merv Griffiths, the pre-eminent monotreme biologist of all time. The publication of this volume has been delayed by the preparation of the papers from “Monotreme III” and is covered by subscriptions and membership fees for 2003. Intending authors should read the summary of “Instructions for Authors” at the back of this volume carefully. More details are available in the full version available at the Society’s web site or from the Secretary. The preparation of this volume has been prolonged and made difficult by the failure of some authors to provide figures and tables in the format required. In order to keep the costs at a level which allows our Society to continue publication, we set the journal completely ourselves. Therefore we do not have the flexibility of large commercial publishers. We can only deal with figures as photographs, original line drawings or .TIF files. Jpeg files for example are useless in our system. Auto-formatting and track changes are a disaster, as are tables and/ or figures that have been put inside the text. To date we have taken the time to re-set and sometimes re-scan figures, however in future we will apply the policy that final copy not prepared in accordance with the instructions will simply be returned and held over if necessary until the next issue. M.L. Augee Editor + sq hee ia ey mauled sete: isi IS ctaNEGS nes Sa ant * ‘ inplow Jo} 2 108 perc eld NG iad ere ody jit bai: asta plete ros . ia plano> noiiase i ail wit fone lat at eset We ty saith ait ite Rie ‘ening cuaheeA, oll Bo ‘a ne ret ad ite’ to et we ns oii ae maak Pa uw cme + a iy rch tine Sui ha bom fad bys it apnea iin simu ran te 4: oii a Sal _ 0 Gk anbiewriaa 0: rte ‘ais wes ialaatn " adt: cane ‘eet. Bia. 3 pia seca tata? ot ue aldstin ve ner owl hfe Yak cole juli re e. aan foal =e eee veoh pi rat dang nifiga ated musi, saanins Aaepeetie Leah 09 yh —: <9) te: < OF (kao hp sini doit tows # agers a peel ot) eo iz ; os rn tas aici ae Sypmel soi DL Sow snarl erator 9 Yaseen: ren a AANT. so-aghi ib oa 4 gH: i yaad fre sft. ste os en (gta ae wat oyun “pt aved yi, ind ¢ or en a iat ay ae Wai cine 4 “ wnt i Review of Australian Cave Guano Ecosystems with a Checklist of Guano Invertebrates | TimotHy MOouLps Centre for Evolutionary Biology and Biodiversity, School of Earth and Environmental Sciences, University of Adelaide, North Terrace, Adelaide 5005. timothy.moulds @ adelaide.edu.au Moulds, T. (2004). Review of Australian cave guano ecosystems with a checklist of guano invertebrates. Proceedings of the Linnean Society of New South Wales 125, 1-42. This work provides a check-list of all invertebrate species known, or believed to be, associated with cave guano in Australia. A total of 240 species in 121 families, representing 25 orders is listed. These species inhabit 60 karst areas in all mainland states of Australia and Christmas Island (Indian Ocean). Comprehensive assessment of all available records (published and in collections) show that the distribution of ‘several species is more extensive than previously believed. It is unknown whether this is because of inadequate identification of specimens, poorly defined taxonomy or unrecognised intra-species variation due to a lack of specimens. Twenty species from five orders show restricted distributions and guano dependence, although endemic status can not yet be assigned. Amongst these species, eight pseudoscorpions and eight Coleoptera are distributed across several mainland states and Christmas Island. Manuscript received 2 July 2003, accepted for publication 22 October 2003. Keywords: Arthropoda, Australia, biospeleology, cave, checklist, ecosystem, guano, invertebrate. INTRODUCTION Australian cave guano ecosystems are poorly known with only a few communities studied in detail (e.g. Richards 1971; Harris 1973; Bellati et al. 2003). Previous studies concerned with the terrestrial cavernicolous fauna of Australia have mentioned species associated with guano, but have provided little in the way of detail with regard to the ecology of specialised guano communities and species. This paper seeks to synthesise the knowledge of guano ecosystems and communities in Australian caves. A review of guano ecosystems and habitats precedes a checklist of all species known to be associated with Australian cave guano deposits. Populations of cavernicolous animals are usually small because of limited food supplies. However, caves containing guano differ fundamentally because there is a virtually unlimited food supply, commonly resulting in large animal populations. Guano in caves is deposited by bats, birds, orthopterans (crickets and grasshoppers), and small mammals, with each type of guano sustaining a unique assemblage of taxa. Guano deposits are extremely variable, unlike other cave habitats, and consist of numerous micro- habitats differentiated by fluctuating temperature, moisture, and pH. Guano ecosystems contain obligate guano-dwelling organisms (guanobites), opportunistic guano-dwelling animals (guanophiles), and transient guano-using animals (guanoxenes) (Gnaspini and Trajano 2000). The basis for many guano ecosystems is the numerous species of fungi and bacteria that can grow on guano, even in complete darkness. Cave food sources Cavernicolous populations are dependant for their survival upon energy inputs into cave systems. These inputs can vary widely, with availability of food usually being the primary limiting factor (Peck 1976). Inflowing streams and periodic floodwaters introduce significant amounts of zooplankton, accidentals, and organic debris that, for many cave ecosystems, represent the main energy inputs (Peck and Christiansen 1990; Humphreys 1991). Tree roots penetrating the roofs and walls are another important energy source found commonly in tropical caves and lava tubes (Hoch 1988; Hoch and Howarth 1999). Dead animals can be a source of food for scavengers near cave entrances (Richards 1971). Accidentals wandering in from cave entrances also provide a food CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST source, although this is generally periodic in nature and inconsistent in quantity. For the most part cave environments are generally depaurporate in food and consequently are sparsely populated with cavernicolous animals. However, caves containing guano deposits differ substantially because they have a virtually unlimited food supply. When present, guano from bats, birds, and Orthoptera (crickets and grasshoppers) generally forms the major energy source (Park and Barr 1961; Poulson 1972; Martin 1977), with large, varied and unique ecosystems often revolving around such deposits. SOURCES AND DIVERSITY OF CAVE GUANO Cave guano deposits from specific sources can each possess a unique assemblage of taxa (Horst 1972; Poulson 1972). Throughout the world’s biogeographic provinces different taxa are responsible for being the most important guano producers. The most widespread and common guano is that produced by bats and these deposits are generally the largest in volume. The spatial and temporal deposition of bat guano differs from tropical to temperate caves. Cave- dwelling bats in temperate regions show an annual cycle of occupancy over summer months when pups are born, before colonies disperse to cooler, wintering caves where they enter torpor. This annual cycle results in large amounts of guano deposited over summer months and then a cessation of guano input for approximately seven months. In contrast, tropical caves generally show constant bat occupancy rather than an annual cycle and less congregation of individuals due to warmer ambient temperatures. Gnaspini and Trajano (2000) note that many bat populations in tropical Brazil are commonly nomadic, resulting in roaming colonies varying their location in an irregular and non-seasonal fashion. This results in non-continuous deposition. The diet of bats (either haematophagous, insectivorous, frugivorous, or nectarivorous) also influences the composition of guano piles and hence the associated guanophilic communities (Gnaspini 1992; Ferreira and Martins 1998, 1999). Large populations of the common vampire bat (Desmodus rotundus Geoffroy) predominate in Brazilian karst near inhabited areas, due to large numbers of domestic livestock resulting in haematophagous guano deposits. Guano from non- haematophagous bats is absent, or greatly reduced as vampire bats exclude other bat species, thus changing the guanophilic communities present. Birds are common guano producers in the northern parts of South America, the Caribbean and Le) tropical caves of south-east Asia. Cave-dwelling birds nest in the dark zone, providing an important energy resource for many cavernicolous animals. Cave- dwelling birds in South American and Caribbean caves include guacharos (Steatornis caripensis Humboldt) (Snow 1975; Gnaspini and Trajano 2000). This bird discards palm seeds, sometimes with flesh still attached, and deposit droppings in caves, thus providing a wide range of organic matter for cavernicolous animals. Because of the presence of discarded seeds, some taxa associated with seeds and detritus, such as lygaeid bugs are found only in guano of this type. Swiftlets (Aerodramus spp) nest in the entrance and dark zones of tropical caves in south- east Asia, northern Australia and the Pacific and are insectivorous (Medway 1962). These birds also support a range of guanophilic taxa in the caves of Christmas Island (Humphreys and Eberhard 2001). Richards (1971) reported that droppings from several species of birds nesting in the entrance zone of Nullarbor Plain caves support a wide variety of cavernicolous animals. Rhaphidophorid crickets are often important producers of guano in temperate caves such as those of the Nullarbor Plain (Richards 1971). The sometimes large populations of these crickets can accumulate sizeable guano deposits in caves. These deposits are important as few other food sources exist in areas such as the Nullarbor Plain because the low mean rainfall limits organic flood debris and bat populations are generally small. Rhaphidophorid guano is also utilised in Mammoth Cave, Kentucky, where it is widely dispersed through the cave system (Howarth 1983). Small mammals are often significant guano producers in temperate zones of North America. The guano of porcupines (Erethizon dorsatum L.) is reported by Calder (1965) to support a community of collembolans and mites active throughout the year in Frenchman’s Cave (Hants County, Nova Scotia, Canada). Cave rats (Neotoma spp), navigate using urine trails (Howarth 1983). Although common in the caves of the Canadian Rockies and Vancouver Island, their faeces are mostly unusable as a food source due to the high ammonia content from systematic urination at these sites (Trapani 1997). GUANO ECOSYSTEMS AND FOOD WEBS Guanobites are animals that require the presence of guano for survival. They will only feed on guano and will not use other food sources within caves. Although guanobitic species are occasionally found on other substrates in caves as they move between discontinuous guano deposits, they do not feed Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS or reproduce on these substrates (Gnaspini and Trajano 2000). When guano deposition is seasonal (e.g. bat maternity caves), guanobites will commonly become quiescent until bats return and restore fresh guano input. Other guanobite populations crash when guano input ceases and then quickly reproduce when guano input recommences. Guanophiles use guano resources opportunistically and are able to complete their entire life cycle using the guano substrate. Guanophiles will however utilise other cave food resources when available and do not have to rely upon guano to feed or reproduce. Abundance of guanophilic animals will decrease if fresh guano is not available, simply due to food limitation, but individuals will attempt to exploit other food resources to survive until fresh guano is available. Troglobites and troglophiles that have a generalist role in epigean ecosystems are classified as guanophiles if they utilise guano when available, even though they are capable of surviving subterranean habitats without this resource. Guanoxenes will exploit a guano resource for feeding or reproduction but require other substrates within a cave to complete their life cycle (Gnaspini and Trajano 2000). Guanoxenes can be either troglobites, troglophiles or trogloxenes (Gnaspini and Trajano 2000). The cyclical nature of many guano deposits resulting from the annual breeding cycle of bats, leads to a similar cycle in arthropod abundances. Low population numbers of many species reflect changes in micro-habitat conditions resulting from the cessation of fresh guano deposition and lower air and guano temperatures. Guano communities decrease in numbers as many species stop breeding until the food supply (i.e. fresh guano) is restored. This has been observed in the mite Uroobovella coprophila Womersley, which is quiescent during winter months in Carrai Bat Cave, northern New South Wales (Harris 1971). Arthropods in guano communities feed either directly on guano or fungus growing on guano deposits and these in turn support a number of predators scavengers and omnivores (Gillieson 1997). Generalised guano food webs have a guano source directly supporting a range of guanivores including Phoridae (Diptera), Anobiidae (Coleoptera), Tineidae (Lepidoptera), Collembola and mesostigmatid mites (Acarina). Predators that prey upon these consumers include spiders, pseudoscorpions, beetles and opiliones. Specialised parasites and parasitoids are also active in many guano ecosystems. Braconid wasps (Hymenoptera) are found in many Australian guano caves and parasitise the larvae of Monopis spp (Lepidoptera: Tineidae). The larvae of the guanobite Proc. Linn. Soc. N.S.W., 125, 2004 Derolathrus sp. (Coleoptera: Jacobsoniidae) are. parasitised by small myrmarid wasps (Hymenoptera). Parasitic relationships in guano ecosystems are generally poorly understood and further research will undoubtedly reveal many more examples. Some of the most numerous taxa associated with guano deposits are mites (Acarina), particularly from the families Gamasidae, Actinedidae, Oribatidae and Armadillidae (Womersley 1963a, b; Gnaspini and Trajano 2000). Extremely high numbers (>33 million/m?) have been recorded on fresh guano (Harris 1973; Bellati 2001). Guanivores from all biogeographic regions are taxonomically similar, usually belonging to the same families. Differences, however, are found among the predators of guanivore communities and are often represented by taxa from different families depending on the biogeographical region (Gnaspini and Trajano 2000). Bat guano micro-habitat variation Guano environments are extremely variable, consisting of numerous micro-habitats when compared with the majority of subterranean habitats (Harris 1970). Bat guano deposits have been found to exhibit variable temperature of both the ambient air above deposits and within deposits (Harris 1970). In addition, the relative humidity, CO concentration, and ammonia concentration also change when bats occupy a cave due to their breathing and urine (Decu 1986). Variations in pH can be extreme, resulting in strong differentiation between fresh and old guano deposits. The annual cycle of bat roosting adds a temporal component to many guano deposits and also serves to alter air temperature in roosting chambers. Bat maternal chambers are especially variable when extremely large numbers of bats enter a chamber on an annual basis to birth young (Harris 1970). Large numbers of bats can raise the air temperature in a chamber by up to 10°C. This effect is most prevalent in high-domed chambers where heated air is trapped, but Harris (1970) also noted small increases in air temperature close to guano piles of up to 1.4°C due to heat released from guano breakdown. Increased air temperature of up to 12°C has also been noted in Cuban caves where large numbers of the leaf- nosed bat, Phyllonycteris poeyi Gundlach, roost (Decu 1986). This temperature increase can act as a barrier for colonisation by generalist cavernicolous invertebrate species, but allows guanophilic and guanobitic populations to reach large numbers. Temperature within a guano pile can increase significantly with depth. Temperatures 5 cm below the surface of guano piles in Carrai Bat Cave, New South Wales are 1.7°C higher compared with surface CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST temperatures, and 15 cm below the surface temperatures are 3.0°C higher (Harris 1970). Surface guano temperatures have also been reported to increase by 9.3°C, and these increases in both surface and subsurface temperatures were attributed by Harris (1970) to the increase in the metabolic rate of the organisms inhabiting the guano pile. The initiation of growth and reproduction of mites in guano may be linked to the increase in temperature associated with bat occupation of a chamber (Harris 1971). Varying water content of guano due to desiccation with increasing age, results in noticeable micro-habitat differentiation. Fresh guano collected from the tops of piles in Bat Cave (U2), Naracoorte, South Australia, has been measured at up to 85% water by weight (Moulds 2003). Guano from the base of piles is a lighter grey colour due to desiccation and can contain as little as 6% water by weight (Moulds 2003). Guano moisture content increases with the birth of pups as their faecal matter is predominately liquid prior to Protochelifer cavernarum (Pseudoscorpionida) Ptinus exulans (Coleoptera) being weened (approximately 6-8 weeks after birth for the large bent-wing bat Miniopterus schreibersii bassanii Cardinal and Christidis) (T. Moulds unpublished data). The surface of guano deposits commonly exhibit a patchwork appearance of dark moist areas and light grey drier areas. Different species within guano ecosystems prefer different micro- habitats. Richards (1971) noted the majority of guanophilic arthropods in Nullarbor Plain caves were only found in completely or partially dry guano. Guano shows a marked difference in pH between fresh and old deposits. Fresh guano is commonly basic, with the pH varying according to the volume of urine deposited with faeces. Fresh guano commonly has a pH of 8.5-9.0 that rapidly becomes acidic (5.0-5.5) with age and depth, although the centre of guano piles has a stable pH of around 4 (Harris 1971). In bat maternity caves the pH of piles will gradually decrease over winter as no fresh guano is deposited. Data from Bat Cave (U2) (Naracoorte, Shawella douglasi (Blattodea) Uroobovella coprophila (Acarina) Figure 1. Different distribution patterns of guano associated species across Australia. Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Table 1. Possibly endemic, guano dependent species in Australia. State Order Genus and Species Dependance Cave Cave Guano QLD Pseudoscorpionida Sathrochthonius webbi Tb Gp Holy Jump Lava Cave (BM1) QLD Coleoptera Choleva australis Tp Gp Royal Arch Cave (CH9) QLD Coleoptera Dermestes uter Tp Gp Royal Arch Cave (CH9) QLD Coleoptera Alphitobius diaperinus Tp? Gp? Bat Cleft (E6) QLD Coleoptera Omorgus costatus Tp Gp? Johannsens Cave (J1-2) QLD Coleoptera Anomotarus subterraneus Tp Gp Riverton Main Cave (RN1) NSW Pseudoscorpionida Oratemnus cavernicola Tp Gp? Jump Up Cave, Gray Range NSW Pseudoscorpionida Sundochernes guanophilus Tp2 Gb Fig Tree Cave (W148) NSW Pseudoscorpionida Tyrannochthonius cavicola Tp2 Gb Grill Cave (B44) NSW Acarina Neotrombidium gracilipes Tp2 Gb Fig Tree Cave (W148) NSW Acarina Hypoaspis annectans Tp Gp Carrai Bat Cave (SCS) Nullarbor Pseudoscorpionida Cryptocheiridium australicum Tp2 Gp Murra-E]-Elevyn Cave (N47) Nullarbor Isopoda Abedaioscia troglodytes Tb Gp? Pannikin Plain Cave (N49) Nullarbor Coleoptera Quedius luridipennis Tp? Gp Abrakurrie Cave (N3) VIC Pseudoscorpionida Pseudotyrannochthonius Tp2 Gp Mount Widderin Cave (H1) hamiltonsmithi VIC Coleoptera Achosia lanigera Tp? Gp Wilson Cave (EB4) SA Pseudoscorpionida Austrochthonicus cavicola Tp2 Gp Cathedral Cave (U12) SA Pseudoscorpionida Protochelifer naracoortensis Tp2 Gp Bat Cave (U2) WA Blattodea Paratemnopteryx atra Tb Gp Mines nr Marble Bar Christmas I Coleoptera Alphitobius laevigatus Unknown Gp Upper Daniel Roux Cave (C156) South Australia), show that late in spring, before guano deposition recommences, tops of guano piles can become acidic, occasionally as low as pH 5.0 (Moulds 2003). The ever changing pH of guano piles due to age and urine content creates marked micro-habitats used by differing species. Micro-habitat variation of bat chambers is further complicated by the movement of bat roosts in a chamber within a breeding season. These movements are a response to avoiding unfavourable conditions caused by ammonia concentrations and high local temperatures (Poulson 1972). DISTRIBUTION, BIOGEOGRAPHY AND ENDEMISM This is the first checklist for Australian guano- associated invertebrates. The full geographic range of many guanobitic and guanophilic species can now easily be appreciated. Many species have been shown to have unexpectedly wide distributions, sometimes spanning several climatic regions. Several possible explanations exist for these patterns. The lack of systematic searching and collation of published records, and collections has resulted in a poor Proc. Linn. Soc. N.S.W., 125, 2004 understanding of many species distribution and degree of endemism. This is commonly combined with a lack of accurate identification by taxonomic experts leading to the lumping of several similar species into one. Inadequate species definitions from groups requiring systematic revision will also result in species being artificially lumped or split (eg Diptera: Phoridae, David McAlpine, pers. comm. 2002). A lack of collections from most karst areas, both above and below ground, is the greatest problem, resulting in large gaps in distributions and a poor knowledge of variation within species. The paucity of records among some taxa also provides a focal point for future collecting priorities. The collation of this checklist has revealed associations of species across wide geographic regions. Figure 1a shows the extensive range of Protochelifer cavernarum Beier (Pseudoscorpionida) from Jurien Bay, Western Australia, across southern Australia and north to Undara Lava Tubes in northern Queensland. The distribution of Shawella douglasi Princis (Blattodea: Blattellidae) (Fig. 1b) is disjunct with records from northern New South Wales and Jurien Bay, Western Australia. This may be the result of misidentification, poor taxonomic description or a paucity of collecting between these localities, especially throughout northern Australia. Despite a CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST number of invertebrate collections from the Nullarbor karst no individuals have been recorded, possibly due to extremely small populations of troglobitic species and the extremely large size of the karst area concerned. Several species including Ptinus exulans Erichson (Coleoptera: Anobiidae) show very wide distributions from mid-north New South Wales across the Nullarbor Plain to the west coast of Western Australia (Fig. Ic). The distribution of U. coprophila (Acarina: Urodinychidae) (Fig. 1d) is directly linked to the distribution of maternal sites for the large bent-wing bat M. schreibersii. The single record of this species from Undara (north Queensland) may be spurious, a misidentification or an individual transported via phoresy, especially as no records exist between southern and northern Queensland despite large bat maternity caves around Rockhampton. These data raise further questions regarding the colonisation of guano deposits by invertebrates and the boundaries of possibly ill-defined species concepts. Endemic status of guano species, has, in the past been assigned without a full understanding of the distribution of Australian guano fauna. This is apparent for the maternal chamber of Bat Cave (U2), Naracoorte, where previous studies (Hamilton-Smith 2000), identified “several endemic species’ to the maternal chamber or Bat Cave as a whole. This checklist has shown that Bat Cave contains only a single endemic species, Protochelifer naracoortensis Beier, and this pseudoscorpion may possibly be found in other caves in the continuous karst of the Otway Basin. Bat Cave does, however, form the most diverse guanophilic arthropod community in Australia. This highlights the amount of assumed knowledge conceming guano invertebrates in Australia and their distribution. The number of endemic species to specific bat caves is currently unknown but is almost certainly significantly lower than previously believed. Several species have been identified as possessing restricted distributions and guano dependence, although none can yet be positively identified as endemic (Table 1). The restricted distribution status of all species listed in Table 1 is tentative and more extensive collecting, both above and below ground, must be undertaken before distribution can be confirmed. This is especially true for troglophilic species as epigean occurrence of these species will effect their endemic status. The degree of a species’ guano dependence will also affect its endemic status and more ecological knowledge is required to confirm species habits. Species confined to single caves or isolated areas are more likely to be endemic when combined with guano dependence. Only Fig Tree Cave (W148) (Wombeyan, NSW) and Royal Arch Cave (CH9) (Chillagoe, QLD) are found to contain two species showing both restricted distribution and guano dependence (Table 1). The presence of nematodes is almost a certainty in guano caves as they are almost ubiquitous in every other habitat both above and below ground. Despite this the records of nematodes from guano are extremely limited primarily because the majority of caves and karst areas remain completely unsampled for these invertebrates. Nematodes play a potentially important role in the micro-habitat of guano piles and have been recorded in large numbers from overseas caves (Decu 1986). Nematodes are also believed to be one of the first colonisers of new bat caves, being deposited by in urine and faeces (Decu 1986). Further sampling of Australian cave guano will almost certainly reveal a greater diversity of species. Currently no free living nematodes have been recorded by the author from Bat Cave, Naracoorte despite several collection events. Currently no guano invertebrates are recorded from Tasmania, primarily due to the absence of cave- dwelling bats. The possibility remains however, that guano communities occur in orthopteran guano or other invertebrate guano deposits or even bird guano. The guanophilic mite Macrocheles tenuirostris Krantz and Filipponi was first recorded from mutton bird nests in Tasmania and has since been collected from bat guano in Victorian and New South Wales caves. Further field observations within Tasmanian caves may yet reveal these communities. Opportunities for future research in this field are vast with only limited knowledge existing for most karst areas. The ecological classification for many species is poorly known and this will only be achieved through increased observations in situ. The microbiology of guano deposits also remain very poorly known in Australian, as well as in overseas caves. Many karst areas remain completely unstudied biologically, especially with regard to the diversity of invertebrate guano communities. SYSTEMATIC CHECK LIST OF AUSTRALIAN GUANO INVERTEBRATES This checklist includes all Australian cavernicolous species found in association with guano from both caves and mines. Records have been compiled from the speleological literature (both scientific and amateur), unpublished records, and personal observations. Parasites of cave-dwelling mammals (bats) have been included as they are often found in guano, although their potential roles in guano ecosystems is currently unknown. Taxa are arranged Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS systematically by Phylum, Class and Order then alphabetically by Family. Undetermined taxa have been placed at the end of their respective order or family. Due to changes in taxonomy and higher systematics of many taxa the names and position of species can be uncertain. This checklist has adopted the most recent higher classifications attainable and many old names have been updated to reflect changes in the literature. Many groups in this checklist are in need of revision and so some species concepts may be altered in the future resulting in the splitting of some species and the lumping of others. This will obviously affect the distribution of species as presented in this work. Cave names and numbers following the Australian Karst Index (Mathews 1985), and are listed for all species’ records along with appropriate references. Records from caves in the Nullarbor Plain, southern Australia, have not been divided along state boundaries in order to reflect the extremely large and continuous nature of this karst area. Taxa previously considered to be obvious accidentals to cave environments have been excluded from this checklist. The following ecological classification is modified from Hamilton-Smith (1967), and Gnaspini and Trajano (2000), and is based on the degree of cave and guano dependence of taxa. Abbreviations are those used in the checklist. Trogloxene (Tx): an organism that regularly uses the cave environment for part of its lifecycle or as shelter but must leave the cave to feed and or breed. 1* order Troglophile (Tp1): an organism that can complete its entire lifecycle within a cave but possess no specific adaptations to the cave environment and : recorded in both epigean and hypogean habitats. 24 order Troglophile (Tp2): an organism that can complete its entire lifecycle within a cave but possess no specific adaptations to the cave environment and recorded only from hypogean habitats. Troglobite (Tb): obligate cavernicolous organisms that possess specific adaptations to the cave environment. Guanoxene (Gx): an organism that may use guano for reproduction and/or feeding but requires other substrates to complete its life cycle. Guanophile (Gp): an organism that inhabits and reproduces both in guano piles as well as other substrates within a cave. Guanobite (Gb): an organism that requires guano deposits to complete its entire life cycle. Bat Parasite (P): an animal that is an obligate bat parasite requiring bats to complete its lifecycle. Ecological classifications have been assigned to taxa wherever possible. These designations were made using available knowledge concerning behaviour, life history, and distribution within caves. However, information regarding species’ ecology was found to be lacking or minimal in most cases. Because of such constraints some taxa have not been assigned a guano classification. Further, information on other taxa was insufficient to confirm their association with guano ecosystems. Thus, taxa previously recorded only from guano caves, but without a confirmed association with guano, have been included for completeness even though some of these species may be unassociated with guano. Phylum Platyhelminthes Class Tubellaria Order undetermined Undetermined genus and species, Tx, Gx?. VICTORIA: Dickson Cave (M30), Murrindal (Yen and Milledge 1990). Phylum Nemathelminthes Class Nematoda Order Strongyloidea Trichostrongylidae Nycteridostrongylus unicollis Baylis, Tx, Gx, P. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data). Molinostrongylus dollfusi Mawson, Tx, Gx, P. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data). Proc. Linn. Soc. N.S.W., 125, 2004 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Order Undetermined ?Rhabditida Undetermined genus and species, Gp?. VICTORIA: Starlight Cave (W5), Warrnambool (T. Moulds unpublished data), bacterial feeder (K. Davies pers. comm. 2003). Undetermined Family Undetermined genus and species, Gp. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (Harris 1970). Undetermined genus and species, Gp. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998). Phylum Mollusca Class Gastropoda Order Stylommatophora Charopidae Elsothera funera Cox, Gx?. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); VICTORIA: Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Shades of Death Cave (M3), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990). Undetermined Family Undetermined genus and species, Tx, Gx?. NORTHERN TERRITORY: Cutta Cutta Cave (K1), Katherine (Hamilton-Smith unpublished data); QUEENSLAND: Carn Dum (E15), Mount Etna (Hamilton-Smith unpublished data). Phylum Annelida Class Oligochaeta Order Haplotaxida Lumbricidae Undetermined genus and species, Tp, Gx?. VICTORIA: Wilson Cave (EB4), East Buchan (Yen and Milledge 1990). Order Undetermined Undetermined genus and species, Tp?, Gx?. QUEENSLAND: Four Mile Cave (C14), Camooweal (Hamilton-Smith unpublished data). Phylum Arthropoda Class Arachnida Order Scorpionida Undetermined Family Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (M11), Murrindal (Yen and Milledge 1990). Order Araneae Agelenidae Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990). Amaurobiidae Undetermined genus and species, Gx?. VICTORIA: Spring Creek Cave (B1), Buchan (Yen and 8 Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990). Ctenizidae Misgolas sp., NEW SOUTH WALES: Yessabah Bat Cave (YE1), Yessabah (Gray 1973b). Cyatholipidae Undetermined genus and species, Gx?. VICTORIA: Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990). Cycloctenidae Cyclotenus abyssinus Urquhart, Tp. VICTORIA: Shades of Death Cave (M3), Murrindal (Hamilton- Smith unpublished data). Toxopsioides sp., Tp. NEW SOUTH WALES: Carrai Bat Cave (SCS), Stockyard Creek (Gray 1973b); Yessabah Bat Cave (YE1), Yessabah (Gray 1973b). Undetermined genus and species, Gx?. VICTORIA: Moon Cave (B2), Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Shades of Death Cave (M3), Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990). Desidae Badumna socialis Rainbow, Tp, Gx?. NEW SOUTH WALES: Chalk Cave (B26), Bungonia (Hamilton-Smith unpublished data). Colcarteria carrai Gray, Tp?. NEW SOUTH WALES: Carrai Bat Cave (SCS), Stockyard Creek (Gray 1992). Colcarteria yessabah Gray, Tp. NEW SOUTH WALES: Carrai Bat Cave (SCS), Stockyard Creek (Gray 1992). Dictynidae Undescribed genus and species, Gx?. VICTORIA: Moon Cave (B2), Buchan (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990). Filistatidae Undescribed genus and species, Tp2. WESTERN AUSTRALIA: Cape Range peninsula (Gray 1994). Gradungulidae Progradungula carraiensis Forster and Gray, Tp1, Gp . NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (Forster et al. 1987). Linyphiidae Laetesia weburdi Urquhart, Gx?. NEW SOUTH WALES: Jenolan Caves (Hamilton-Smith unpublished data). Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (M11), Murrindal (Yen and Milledge 1990). Lycosidae Lycosa speciosa Koch, Tp1. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (Gray 1973b). Proc. Linn. Soc. N.S.W., 125, 2004 9 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Mimetidae Australomimetus maculosus Rainbow, Tp. NEW SOUTH WALES: Yessabah Bat Cave (YE1), Yessabah (Gray 1973b); Colong Main Cave (CG1), Colong (Hamilton-Smith unpublished data); Jenolan Caves (Hamilton-Smith unpublished data). Undetermined genus and species, Gx?. VICTORIA: Spring Creek Cave (B1), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990). Pholcidae Physocyclus sp., NEW SOUTH WALES: Carrai Bat Cave (SCS), Stockyard Creek (Gray 1973b); Colong Main Cave (CG3), Colong (Gray 1973b). Psilochorus sp., NEW SOUTH WALES: Yessabah Bat Cave (YE1), Yessabah (Gray 1973b) . Pisauridae Undetermined genus and species, NEW SOUTH WALES: Comboyne C4 Cave, Comboyne (Gray 1973b); Carrai Bat Cave (SC5), Stockyard Creek (Gray 1973b). Salticidae Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (M11), Murrindal (Yen and Milledge 1990). Segestriidae Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (M11), Murrindal (Yen and Milledge 1990). Stiphidiidae Stiphidon sp., Gx?. NEW SOUTH WALES: Colong Cave (CG1), Colong (Hamilton-Smith unpublished data). Theridiidae Theridon sp., Tp, Gp. NEW SOUTH WALES: Colong Cave (CG1), Colong (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Steatoda sp., Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Theridiosomatinae Undetermined genus and species, Gp?. NEW SOUTH WALES: Colong Cave (CG1), Colong (Hamilton-Smith unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data). Uloboridae Philoponella patherinus Keyserling, Tp. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data). Undetermined Family Undetermined genus and species, Gp?. NEW SOUTH WALES: Cave C4, Comboyne (Hamilton- Smith unpublished data); The Drum Cave (B13), Bungonia (Hamilton-Smith unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Colong Cave (CG1), Colong (Hamilton-Smith unpublished data); Gable Cave (CL7), Cliefden (Hamilton-Smith unpublished data); Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith unpublished data); Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave (T1), Tuglow (Hamilton-Smith unpublished data); Punchbowl Cave (WJ8), Wee Jasper (Hamilton- 10 Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Smith unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); Basin Cave (W4), Wombeyan (Hamilton-Smith unpublished data); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data); NORTHERN TERRITORY: Cutta Cutta Cave (K1), Katherine (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Abrakurrie Cave (N3) (Hamilton-Smith unpublished data); Madura Cave (Madura 6 Mile Cave) (N62) (Hamilton-Smith unpublished data); QUEENSLAND: Four Mile Cave (C14), Camooweal (Hamilton-Smith unpublished data); Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith unpublished data); Holy Jump Lava Cave (BM1), Bauer’s Mountain (Hamilton-Smith unpublished data); Barker’s Cave (U34), Undara (Hamilton-Smith unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith unpublished data); Speaking Tube (E7), Mount Etna (Hamilton-Smith unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data); Piglet Help! Help! Cave (E17), Mount Etna (Hamilton-Smith unpublished data); Ilium Cave (E31), Mount Etna (Hamilton-Smith unpublished data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Snowflake Cave (L1), Glenelg River (Hamilton-Smith unpublished data); Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Moon Cave (B2), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Shades of Death Cave (M3), Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); SSS Cave (M44), Murrindal (Hamilton-Smith unpublished data); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Bat Cave (P6), Portland (Hamilton-Smith unpublished data); Mt Widderin Cave (H1), Skipton (Hamilton-Smith unpublished data); Panmure Cave (H5), Mount Napier (Hamilton- Smith unpublished data); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); Grassmere Cave (W6), Warrnambool (Hamilton-Smith unpublished data). Order Opilionida Triaenoncychidae Holonuncia cavernicola Forster, Tp2. NEW SOUTH WALES: Basin Cave (W4), Wombeyan (Hamilton-Smith 1967); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data). Holonuncia seriata Roewer, Tp1, Gx. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998). Undetermined genus and species, Tp, Gp. VICTORIA: Moon Cave (B2), Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Shades of Death Cave (M3), Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990). Undetermined Family Undetermined genus and species, Tp, Gp? NEW SOUTH WALES: The Drum Cave (B13), Bungonia (Hamilton-Smith unpublished data); Chalk Cave (B26), Bungonia (Hamilton-Smith unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Cliefden Main Cave (CL1), Cliefden (Hamilton-Smith unpublished data); Gable Cave (CL7), Cliefden (Hamilton- Smith unpublished data); Colong Main Cave (CG1), Colong (Hamilton-Smith unpublished data); Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith unpublished data); Moparabah Cave (Temagog Cave) (MP1), Moparabah (Hamilton-Smith unpublished data); Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave (T1), Tuglow (Hamilton-Smith unpublished data); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data); Yessabah Bat Cave (YE1), Yessabah (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Lynch Cave (N60) (Hamilton-Smith unpublished data); QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); Proc. Linn. Soc. N.S.W., 125, 2004 11 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST VICTORIA: Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Unnamed Cave (NG1), New Guinea Ridge (Hamilton-Smith unpublished data). Order Pseudoscorpionida Atemnidae Oratemnus cavernicola Beier, Tp, Gp?. NEW SOUTH WALES: Jump Up Cave, Gray Range (Beier 1976). Cheiridiidae Cryptocheiridium australicum Beier, Tp2, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 1971). Cheliferidae Protochelifer naracoortensis Beier, Tp2, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Beier 1968; Bellati et al. 2003). Protochelifer cavernarum Beier, Tp2, Gb. NEW SOUTH WALES: Murder Cave (CL2), Cliefden (Beier 1967, 1968); Island Cave (CL6), Cliefden (Hamilton-Smith unpublished data); Belfry Cave (TR2), Timor (Beier 1967); Ashford Caves, Ashford (Beier 1968); NULLARBOR PLAIN: Warbla Cave (N1) (Richards 1971); Weebuddie [Weebubbie, sic] Cave (N2) (Beier 1975); Abrakurrie Cave (N3) (Richards 1971); Murrawijinie No.3 Cave (N9) (Richards 1971); Mullamullang Cave (N37) (Richards 1971); Lynch Cave (N60) (Richards 1971); QUEENSLAND: Taylor Cave (4U4), Undara (Howarth 1988); Collins Cave No.1, Undara (Howarth 1988); VICTORIA: Clogg’s Cave (EB2), East Buchan (Beier 1968); WESTERN AUSTRALIA: Gooseberry Cave (J1), Jurien Bay (Beier 1968); Eneabba Caves (E1-3), Eneabba (Lowry 1996); Arramall Cave (E22), Eneabba (Lowry 1996); River Cave (E23), Eneabba (Lowry 1996); Weelawadji Cave (E24) Eneabba (Lowry 1996); Super Cave (SH1), South Hill River (Hamilton-Smith unpublished data). Protochelifer sp. Tp, Gp. SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Bellati et al. 2003). Chernetidae Sundochernes guanophilus Beier, Tp2, Gb. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Beier 1967). Troglochernes imitans Beier, Tp, Gp. NULLARBOR PLAIN: Murra-El-Elvyn Cave (N47) (Beier 1975); Cocklebiddy Cave (N48) (Beier 1975); Pannikin Plain Cave (N49) (Beier 1975); Dingo Cave (Dingo-Donga) (N160) (Richards 1971). Chthoniidae Austrochthonius cavicola Beier, Tp, Gp. SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Beier 1968). Paraliochthonius cavicolus Beier, Tp2, Gp. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998). Pseudotyrannochthonius hamiltonsmithi Beier, Tp2, Gp. VICTORIA: Mt Widderin Cave (H1), Skipton (Beier 1968). Sathrochthonius tuena Chamberlin, Tp2, Gp. NEW SOUTH WALES: Basin Cave (W4), Wombeyan (Beier 1967), Deua Cave (DE1), Deua (Eberhard and Spate 1995); Punchbowl Cave (WJ8), Wee Jasper (Beier 1968); Imperial Cave (J4), Jenolan (Hamilton-Smith 1967; Gibian et al. 1988); Southern Limestone, Jenolan (Hamilton-Smith 1967; Beier 1968; Gibian et al. 1988); Paradox Cave (J48), Jenolan (Hamilton-Smith unpublished data). Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Sathrochthonius webbi Muchmore, Tb, Gp. QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer’s Mountain southern Queensland (Muchmore 1982). Tyrannochthonius cavicola Beier, Tp2, Gb. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Beier 1967; Harvey 1989). Undetermined Family Undetermined genus and species, Tp, Gp?. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); QUEENSLAND: Royal Arch Cave (CH9), Royal Arch Tower, Chillagoe (Matts 1987). Undetermined genus and species, Tp?, Gx. VICTORIA: Anticline Cave (M11), Murrindal (Yen and Milledge 1990). Mites The mites have been arranged according to the higher classification used by Halliday (1998). Many changes to nomenclature have occurred since previous checklists of cavernicolous fauna have been published so the family placement of some species has been updated to reflect this. Previous family placements have not been recorded but where synonymy has occured the old name (either family or genus) has been included in brackets. Previous generic placements have been recorded in brackets with the prefix “=”. Order Acariformes Suborder Astigmata Histiostomatidae Histiostoma sp. NULLARBOR PLAIN: Mullamullang Cave (N37) (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Rosensteiniidae Nycteriglyphus (Coproglyphus) dewae Zakhvatkin, Tp2, Gb. NEW SOUTH WALES: Basin Cave (W4), Wombeyan (Womersley 1963a; Richards 1967b); Fig Tree Cave (W148), Wombeyan (Womersley 1963a; Richards 1967b); Railway tunnel, North Sydney (Womersley 1963a); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Womersley 1963a). Nycteriglyphus sp., Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 1971); Dingo Cave (160) (Richards 1971). Glycyphagus sp., Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 1971); Dingo Cave (N160) (Richards 1971). Suborder Prostigmata Labidostomidae Undetermined genus and species. NEW SOUTH WALES: Island Cave (CL6), Cliefden (Hamilton- Smith unpublished data). Neotrombidiidae Neotrombidium gracilare Womersley, Tp2, Gb. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Womersley 1963a); Murder Cave (CL2), Cliefden (Womersley 1963a); Punchbowl Cave (WJ8), Wee Jasper (Womersley 1963a); VICTORIA: O’Rourke’s Cave (B12), Buchan (Hamilton-Smith 1967); Wilson Cave (EB4), East Buchan (Hamilton-Smith 1967). Neotrombidium gracilipes Womersley, Tp2, Gb. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Hamilton-Smith 1967). Proc. Linn. Soc. N.S.W., 125, 2004 13 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Neotrombidium neptunium Southcott, VICTORIA: Clogg’s Cave (EB2), East Buchan (Hamilton- Smith unpublished data). Neotrombidium sp., Tp, Gb. NULLARBOR PLAIN: Firestick Cave (N70) (Richards 1971); Dingo Cave (N160) (Richards 1971). Trombiculidae Rudnicula barbarae Domrow (= Trombicula), Tx, Gx, P. NORTHERN TERRITORY: Kuhinoor Mine, Pine Creek (Hamilton-Smith unpublished data). Trombicula thomsoni Womersley, Tx, Gx, P. NEW SOUTH WALES: Bonalbo Colliery (Hamilton- Smith unpublished data); Riverton (Hamilton-Smith unpublished data); NORTHERN TERRITORY: Kuhinoor Mine, Pine Creek (Hamilton-Smith unpublished data). Trombicula dewae Domrow, Tx, Gx, P. NORTHERN TERRITORY: Kuhinoor Mine, Pine Creek (Hamilton-Smith unpublished data). Order Parasitiformes Suborder Ixodida Argasidae Argas sp., Tx, Gx, P, QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data). Ixodidae Amblyomma moreliae Koch, Gx, P. QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data). Ixodes simplex simplex Neumann, Gx, P. Bat parasite in eastern Australia (Hamilton-Smith 1966b; Eberhard 1998); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); Spring Creek Cave (B1), Buchan (Hamilton-Smith unpublished data); Slocombe’s Cave (BA1), The Basin (Hamilton-Smith unpublished data); Anticline Cave (M11), Murrindal (Hamilton-Smith unpublished data); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); Grassmere Cave (W6), Warrnambool (Hamilton-Smith unpublished data). Undetermined genus and species, Gx, P,. QUEENSLAND: Clam Cavern (CH26), Walkunder Tower, Chillagoe (Matts 1987); Spatial Cavern (CH41), Walkunder Tower, Chillagoe (Matts 1987); Royal Arch Cave (CH9), Royal Arch Tower, Chillagoe (Matts 1987); VICTORIA: Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data) Suborder Mesostigmata . Ameroseiidae Ameroseius plumosus Oudemans, Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Laelapidae Cosmolaelaps sp., Tp2, Gb. NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper (Hamilton- Smith 1967); QUEENSLAND: Railway tunnel, Samford (Hamilton-Smith 1967). Hypoaspis (Gaeolaelaps) annectans Womersley, Tp, Gp. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (Harris 1971). Hypoaspis (Gaeolaelaps) sp.1, SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Hypoaspis (Gaeolaelaps) sp.2, SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Hypoaspis (Gaeolaelaps) sp., Tp2, Gb. NEW SOUTH WALES: Cave C4, Comboyne (Hamilton- Smith 1967). Ichoronyssus (Pleisiolaelaps) miniopteri (Zumpt and Patterson 1952) (= Neospinolaelaps, Spinolaelaps), Tp, Gx, P. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998); Bonalbo Colliery (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data). Ichoronyssus (Pleisiolaelaps) aristippe Domrow, NEW SOUTH WALES: Cheitmore Cave, Cheitmore (Hamilton-Smith unpublished data); Wombeyan Caves (Hamilton-Smith unpublished data); Bonalbo Colliery (Hamilton-Smith unpublished data). Macrochelidae Macrocheles spatei Halliday, Tp1, Gp. NEW SOUTH WALES: Deua Cave (DE1), Deua National Park (Halliday 2000). Macrocheles tenuirostris Krantz and Filtpponi, Tp1, Gp. NEW SOUTH WALES: Paradox Cave (J48), Jenolan Caves (Halliday 2000); Cleatmore Cave, Deua National Park (Halliday 2000); Colong Cave, Woof’s Cavern (CG1), Colong (Halliday 2000); Church Cave (WJ31), Wee Jasper (Halliday 2000); TASMANIA: Fisher Island, in nests and burrows of muttonbird (Krantz and Fillipponi 1964); VICTORIA: Panmure Cave (H5), Warrnambool (Hamilton-Smith 1967). Macronyssidae Macronyssus aristippe Domrow (= Ichoronyssus), Tp, Gx, P. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998). Trichonyssus australicus Womersley, Tx, Gx, P. NULLARBOR PLAIN: Warbla Cave (N1) (Hamilton-Smith unpublished data). Parantennulidae Micromegistus gourlayi Womersley. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne (Hamilton-Smith unpublished data). Parasitidae ?Eugamasus sp., Tp, Gp. NULLARBOR PLAIN: Dingo Cave (N160) (Richards 1971). Sejidae (Ichthyostomatogastridae) Asternolaelaps australis Womersley and Domrow, Tp, Gb. SOUTH AUSTRALIA: Bat Cave (U2) Naracoorte (Womersley and Domrow 1959; Hamilton-Smith 1967); VICTORIA: O’Rourkes Cave (B12), Buchan (Hamilton-Smith 1967). Spinturnicidae Spinturnix psi Kolenati, Tp, Gx, P. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998). Undetermined genus and species, Tp, Gx, P. NEW SOUTH WALES: Colong Main Cave (CG1), Colong (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Weebubbie Cave (N2) (Hamilton-Smith unpublished data); Murra-El-Elevyn Cave (N47) (Hamilton-Smith unpublished data); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); Flogged Horse Cave (Cammoo Cave) (J83), Limestone Ridge, Rockhampton (Hamilton-Smith Proc. Linn. Soc. N.S.W., 125, 2004 15 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (B1), Buchan (Hamilton-Smith unpublished data); WESTERN AUSTRALIA: Stockyard Cave (E3), Eneabba (Hamilton-Smith unpublished data). Urodinychidae Uroobovella (Austruropoda) coprophila Womersley (= Cilliba), Tp2, Gp. NEW SOUTH WALES: Cave C4, Comboyne (Smith 1982b); Carrai Bat Cave (SC5), Stockyard Creek (Harris 1973); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data); Church Cave (WJ31), Wee Jasper (Hamilton-Smith 1966b, 1967); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith 1966b, 1967); Cheitmore Cave, Cheitmore (Hamilton-Smith unpublished data); QUEENSLAND: Arch Cave (U22), Undara (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); VICTORIA: Anticline Cave (M11), Murrindal (Hamilton-Smith 1967). Genus and species undetermined, NEW SOUTH WALES: Deua Cave (DE1), Deua (Eberhard and Spate 1995). Undetermined Family Undetermined sp. 1, Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 1971). Undetermined sp. 2, Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 1971); Undetermined Acarina Undetermined Family Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Grimes Cave (CI53) (Humphreys and Eberhard 2001). Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Gable Cave (CL7), Cliefden (Hamilton-Smith unpublished data); NORTHERN TERRITORY: Kintore Cave (K2), Katherine (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Weebubbie Cave (N2) (Hamilton- Smith unpublished data); Murra-El-Elevyn Cave (N47) (Hamilton-Smith unpublished data); QUEENSLAND: Flogged Horse Cave (Cammoo Cave) (J83), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Asbestos mine near Arkaba, Flinders Ranges (Hamilton-Smith unpublished data); Drop Drop Cave (L29), Lower south east (Hamilton- Smith unpublished data); Joanna Bat Cave (U38), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (B1), Buchan (Yen and Milledge 1990); O’Rourkes Cave (B12), Buchan (Hamilton-Smith unpublished data); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson’s Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Bat Cave (P6), Portland (Hamilton-Smith unpublished data); Grassmere Cave (W6), Warrnambool (Hamilton-Smith unpublished data). Class Crustacea Order Isopoda Armadillidae Merulana sp. nov., Tp. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Dennis 1986). Oniscidae Plymophiloscia sp. Vandel, Tp, Gp. NULLARBOR PLAIN: Pannikin Plain Cave (N49) (Richards 1971; Gray 1973a). 16 Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Undetermined genus and species, Tp, Gp. QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); Speaking Tube (E7), Mount Etna (Hamilton-Smith unpublished data); Carn Dum (E15), Mount Etna (Hamilton-Smith unpublished data); VICTORIA: Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data). Philosciidae Abebaioscia troglodytes Vandel, Tb, Gp?. NULLARBOR PLAIN: Pannikin Plain Cave (N49) (Vandel 1973). Eurygastor montanus troglophilus Vandel, Tp?. VICTORIA: Anticline Cave (M11), Murrindal (Vandel 1973). Laevophiloscia dongarrensis Wahrberg, Tp, Gx?. WESTERN AUSTRALIA: Yanchep Cave (YN16), Yanchep (Vandel 1973); Minnie’s Grotto (YN28), Yanchep (Vandel 1973); Gooseberry Cave (J1), Jurien Bay (Vandel 1973). Laevophiloscia hamiltoni Vandel, Tp, Gx. WESTERN AUSTRALIA: Weelawadji Cave (E24), Eneabba (Vandel 1973); Labyrinth Cave (AU16), Augusta (Vandel 1973) Laevophiloscia michaelseni Vandel, Tp. NULLARBOR PLAIN: Cocklebiddy Cave (N48) (Vandel 1973). Porcellionidae Porcellio scaber Latreille, Tp1. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Undetermined Family Undetermined genus and species, Tp, Gx. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith unpublished data); The Drum Cave (B13), Bungonia (Hamilton-Smith unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Cliefden Main Cave (CL1), Cliefden (Hamilton-Smith unpublished data); Cave C4, Comboyne (Hamilton-Smith unpublished data); Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith unpublished data); Moparabah Cave (Temagog Cave) (MP1), Moparabah (Hamilton-Smith unpublished data); Main Cave (Ballroom Cave) (TR1), Timor (Hamilton-Smith unpublished data); Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave (T1), Tuglow (Hamilton-Smith unpublished data); Piano Cave (Long Cave) (WA12), Walli (Hamilton-Smith unpublished data); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); Yessabah Bat Cave (YE1), Yessabah (Hamilton-Smith unpublished data); NORTHERN TERRITORY: Cutta Cutta Cave (K1), Katherine (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Abrakurrie Cave (N3) (Hamilton-Smith unpublished data); Cocklebiddy Cave (N48) (Hamilton-Smith unpublished data); QUEENSLAND: Barker’s Cave (U34), Undara (Hamilton-Smith unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); Fox Cave (U22), Naracoorte (Hamilton- Smith unpublished data); Cave Park Cave (U37), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (B1), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Shades of Death Cave (M3), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); WESTERN AUSTRALIA: Drovers Cave (J2), Jurien Bay (Hamilton-Smith unpublished data); Stockyard Cave (E3), Eneabba (Hamilton-Smith unpublished data). Proc. Linn. Soc. N.S.W., 125, 2004 17 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Order Amphipoda Undetermined Family Undetermined genus and species, Tp, Gx. VICTORIA: Wilson Cave (EB4), East Buchan (Yen and Milledge 1990). Class Myriapoda Order Diplopoda Undetermined Family Undetermined genus and species, Tp2, Gx. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998). Undetermined genus and species, Tp?, Gx. NEW SOUTH WALES: Island Cave (CL6), Cliefden (Hamilton-Smith unpublished data); The Drum Cave (B13), Bungonia (Hamilton-Smith unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Paradox Cave (J48), Jenolan (Hamilton-Smith unpublished data); Moparabah Cave (Temagog Cave) (MP1), Moparabah (Hamilton-Smith unpublished data); Carrai Bat Cave (SC5), Stockyard Creek (Hamilton-Smith unpublished data); Belfry Cave (TR2), Timor (Hamilton-Smith unpublished data); Tuglow Cave (T1), Tuglow (Hamilton-Smith unpublished data); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); Yessabah Bat Cave (YE1), Yessabah (Hamilton-Smith unpublished data); QUEENSLAND: Barker’s Cave (U34), Undara (Hamilton-Smith unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data); Piglet Help! Help! Cave (E17), Mount Etna (Hamilton-Smith unpublished data); Jolly Roger Cave (E29), Mountt Etna (Hamilton-Smith unpublished data); Glen Lyon River Cave (GL1), Glen Lyon (Hamilton-Smith unpublished data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (B1), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data). Order Chilopoda Scolopendromorpha Undetermined genus and species. NULLARBOR PLAIN: Mullamullang Cave (N37) (Richards 1971). Undetermined Family Undetermined genus and species, Gp?. NEW SOUTH WALES: Cave C4, Comboyne (Hamilton- Smith unpublished data); Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith unpublished data);:Carrai Bat Cave (SC5), Stockyard Creek (Hamilton-Smith unpublished data); Moparabah Cave (MP1), Moparabah (Hamilton-Smith unpublished data); Belfry Cave (TR2), Timor (Hamilton-Smith unpublished data); NORTHERN TERRITORY: Cutta Cutta Cave (K1), Katherine (Hamilton-Smith unpublished data); Kintore Cave (K2), Katherine (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Cocklebiddy Cave (N48) (Hamilton-Smith unpublished data); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Superclass Hexapoda Class Insecta Order Collembola Armadillidae Buddelundia albomarginata Wahrberg, Tp, Gx?. NULLARBOR PLAIN: Murrawyinee [sic] No.1 Cave (N7) (Vandel 1973); Cocklebiddy Cave (N48) (Vandel 1973); Lynch Cave (N60) (Vandel 1973); Madura Cave (N62) (Vandel 1973); Old Homestead Cave (N83) (Vandel 1973); Unnamed cave (N140) (Vandel 1973). Entomobryidae Lepidocyrtus sp., Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Lepidosira australica Schott, Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Undetermined genus and species, Gp?. NEW SOUTH WALES: Belfry Cave (TR2), Timor (James et al. 1976); Chalk Cave (B26), Bungonia (Hamilton-Smith unpublished data). Hypogastruridae Hypogastrura sp., NEW SOUTH WALES: Grill Cave (B44), Bungonia (Wellings 1977); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Isotomidae Folsomia candida Willem, Tp. NEW SOUTH WALES: Paradox Cave (J48), Jenolan (Eberhard 1993); Imperial Cave (J4), Jenolan (Eberhard and Spate 1995); Tuglow Main Cave (T1), Tuglow (Eberhard 1993); Jillebean Cave (Y22), Yarrangobilly (Eberhard 1993). Paronellidae Undetermined genus and species, NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Eberhard and Spate 1995). Undetermined Family Undetermined genus and species, Tp, Gp. NULLARBOR PLAIN: Cocklebiddy Cave (N48) (Richards 1971); Lynch Cave (N60) (Richards 1971); Dingo Cave (N160) (Richards 1971); VICTORIA: SSS Cave (M44), Murrindal (Hamilton-Smith unpublished data); Mt Widderin Cave (H1), Skipton (Hamilton-Smith unpublished data). — Undetermined genus and species, Tp?, Gp?. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Colong Main Cave (CG1), Colong (Hamilton-Smith unpublished data); Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); NORTHERN TERRITORY: 16 Mile Cave, Katherine (Hamilton-Smith unpublished data); QUEENSLAND: Speaking Tube (E7), Mount Etna (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Moon Cave (B2), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson’s Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). Order Diplura Undetermined family Undetermined genus and species, SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Proc. Linn. Soc. N.S.W., 125, 2004 19 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Order Blattodea Blattellidae Blattidae Neotemnopteryx australis Saussure, Tp, Gp. NEW SOUTH WALES: Moparabah Cave (Temagog Cave) (MP1), Moparabah (Hamilton-Smith 1967); Cave C4, Comboyne (Hamilton-Smith 1967); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Neotemnopteryx fulva Saussure (= Gislenia australica Brunner), Tp, Gb. NEW SOUTH WALES: Glen Dhu Cave (Allston Cave) (TR15), Timor (Richards 1967a); Murder Cave (CL2), Cliefden (Richards 1967a); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Richards 1967a); Haystall Cave (U23), Naracoorte (Richards 1967a); VICTORIA: Mabel Cave (EB1), East Buchan (Richards 1967a). Neotemnopteryx sp., Tp, Gp?. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith unpublished data); QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith unpublished data). ?Neotemnopteryx (?Gislenia sp.), Tp, Gp?. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Richards 1967a); Cave 4, Comboyne (Richards 1967a); Hill Cave (TR7), Timor (Richards 1967a); Moparabah Cave (Temagog Cave) (MP1), Moparabah (Richards 1967a); Swallow Cave (CU1), Cudgegong (Richards 1967a); QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Richards 1967a); Riverton Main Cave (RN1), Riverton, southern Queensland (Richards 1967a); Viator Cave (VR4), Viator Hill, southern Queensland (Richards 1967a); Johannsen’s Cave (J1), Limestone Ridge, Rockhampton (Hamilton-Smith 1967); Winding Stairway Cave (4E2), Mt Etna (Hamilton-Smith 1967); SOUTH AUSTRALIA: Alexandra Cave (5U3), Naracoorte (Richards 1967a); Bat Cave (U2), Naracoorte (Richards 1967a). Paratemnopteryx atra Princis, Tb, Gp. WESTERN AUSTRALIA: Mines near Marble Bar (Princis 1963; Richards 1967a; Moore et al. 2001). Paratemnopteryx rufa Tepper, Gb?. NULLARBOR PLAIN: Murrawijinie No.3 Cave (N9) (Richards 1971); Abrakurrie Cave (N3) (Richards 1971). Paratemnopteryx sp., Tp, Gb?. QUEENSLAND: Pinwill Cave (4U17), Undara (Howarth 1988). Shawella douglasi Princis, Tp, Gb?. NEW SOUTH WALES: River Cave (SC1), Stockyard Creek (Hamilton-Smith 1967); WESTERN AUSTRALIA: Drovers Cave (J2), Jurien Bay (Hamilton-Smith unpublished data); Jurien Bay caves (Princis 1963; Richards 1967a); Eneabba Caves (E1-3), Eneabba (Lowry 1996); Weelawadji Cave (E24), Eneabba (Lowry 1996). Trogloblattella nullarborensis Mackerras, Tb, Gp. NULLARBOR PLAIN: Abrakurrie Cave (N3) (Mackerras 1967; Richards 1971); Koonalda Cave (N4) (Mackerras 1967); Mullamullang Cave (N37) (Mackerras 1967); Roaches Rest Cave (N58) (Mackerras 1967); Arubiddy Cave (N81) (Mackerras 1967). Polyzosteria mitchelli Angas, Tp. NULLARBOR PLAIN: Warbla Cave (N1), (Mackerras 1965); Kestrel Cavern (N40) (Mackerras 1965; Richards 1967a). Polyzosteria pubescens Tepper, Tp. NULLARBOR PLAIN: Weebubbie Cave (N2) (Hamilton- Smith unpublished data). Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Zonioploca medilinea Tepper, Tp?. NULLARBOR PLAIN: Warbla Cave (N1) (Richards 1967a). Order Orthoptera Rhaphidophoridae Australotettix carraiensis Richards, Tp, Gx. NEW SOUTH WALES: Barnett’s Cave (SC6), Stockyard Creek (Richards 1964); Carrai Bat Cave (SC5), Stockyard Creek (Richards 1964); Col’s Cave, Stockyard Creek (Richards 1964); Lot’s Mansion, Stockyard Creek (Richards 1964); River Cave (SC1) Stockyard Creek (Richards 1964). Cavernotettix buchanensis Richards, Tx, Gx. VICTORIA: Wilson Cave (EB4), East Buchan (Richards 1966; Yen and Milledge 1990); Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Spring Creek Cave (B1), Buchan (Richards 1966; Yen and Milledge 1990); Shades of Death Cave (M3), Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990); Nargun’s Cave (NN1), Nowa Nowa Caves (Richards 1966; Yen and Milledge 1990); Weta Cave (NN2), Nowa Nowa Caves (Richards 1966; Yen and Milledge 1990). Cavernotettix montanus Richards, Tx, Gx. NEW SOUTH WALES: small cave nr Glory Cave, Yarrangobilly (Richards 1966); Jersey Cave (Y23), Yarrangobilly (Richards 1966); Restoration Cave (Y50), Yarrangobilly (Richards 1966); Unnamed cave, Yarrangobilly (Richards 1966); Cooleman Cave (CP1), Cooleman Plains (Richards 1966); Unnamed cave opp. Blue Waterhole, Cooleman Plains (Richards 1966); Unnamed cave nr Murray Cave, Cooleman Plains (Richards 1966). Cavernotettix wyanbenensis Richards, Tx, Gx. NEW SOUTH WALES: Wyanbene Cave (WY 1), Wyanbene (Richards 1966); Bat Cave, Cheitmore (Richards 1966). Pallidotettix nullarborensis Richards, Tx, Gx. NULLARBOR PLAIN: Warbla Cave (N1) (Richards 1971); Weebubbie Cave (N2) (Richards 1971); Murra-El-Elevyn Cave (N47) (Richards 1971); Cocklebiddy Cave (N48) (Richards 1971); Pannikin Plain Cave (N49) (Richards 1971); Tommy Grahams Cave (N56) (Richards 1971). Undetermined genus and species, Tx, Gx. NEW SOUTH WALES: Gmill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Colong Main Cave (CG1), Colong (Hamuilton-Smith unpublished data); QUEENSLAND: Danes Four Cave (C4), Camooweal (Hamilton-Smith unpublished data); Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data); Haunted Cave (CH1), Chillagoe (Hamilton-Smith unpublished data); VICTORIA: Starlight Cave (W5), Warrnambool (T. Moulds unpublished data). Order Psocoptera Liposcelidae Liposcelis corrodens Broadhead, Tp1, Gp. WESTERN AUSTRALIA: Arranmall [sic] Cave (E22), Eneabba (Smithers 1975); undetermined caves (Smithers 1975). Psyllipsocidae ?Psyllipsocus ramburi Selys-Longcamp, Tp1, Gp. NEW SOUTH WALES: Murder Cave (CL2), Cliefden (Hamilton-Smith 1967); Island Cave (CL6), Cliefden (Smithers 1964); Hill Cave (TR7), Timor (James et al. 1976); Basin Cave (W4), Wombeyan (Smithers 1964); Fig Tree Cave (W148), Wombeyan (Smithers 1975); Punchbowl Cave (WJ8), Wee Jasper (Smithers 1964); Church Cave (WJ31), Wee Jasper (Smithers 1964); NULLARBOR PLAIN: Weebubbie Cave (N2) (Richards 1971); Abrakurrie Cave (N3) (Hamilton-Smith 1967; Richards 1971); Koonalda Cave (N4) (Richards 1971); Madura Cave (N62), (Richards 1971); QUEENSLAND: Riverton Main Cave (RN1), Riverton, southern Queensland (Hamilton-Smith 1967); SOUTH AUSTRALIA: Bat Cave Proc. Linn. Soc. N.S.W., 125, 2004 21 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST (U2), Naracoorte (Smithers 1964; Bellati et al. 2003); Blackberry Cave, Naracoorte (Smithers 1964); VICTORIA: Clogg’s Cave (EB2), East Buchan (Smithers 1964); O’Rourkes Cave (B12), Buchan (Smithers 1964); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith 1967). Trogiidae Lepinotus inquilinus Heyden, Tp1, Gp. WESTERN AUSTRALIA: Arranmall (sic) Cave (E22), Eneabba (Smithers 1975). ?Lepinotus reticulatus Enderlein, Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Smithers 1964; Bellati et al. 2003) Undetermined genus and species, NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Dennis and Mayhew 1986). Undetermined Family Undetermined genus and species, Tp, Gx. NEW SOUTH WALES: Gable Cave (CL7), Cliefden (Hamilton-Smith unpublished data); QUEENSLAND: Viator Main Cave (VR1), Viator Hill (Hamilton-Smith unpublished data); VICTORIA: Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990). Order Hemiptera Cixiidae Undetermined genus and species, Tp. QUEENSLAND: Mount Etna Main Cave (E1), Mount Etna (Hamilton-Smith unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data). Lygaeoidea Undetermined family and genus, Gp?. VICTORIA: Starlight Cave (W5), Warrnambool (T. Moulds unpublished data). Reduviidae Armstrongula sp. Tp, Gp. SOUTH AUSTRALIA: McKinley’s Daughter’s Cave (F175), Flinders Ranges (T. Moulds unpublished data); Unnamed mine, Weetootla Gorge, Gammon Ranges (T. Moulds unpublished data). Centrogonus sp. Tp, Gp. NORTHERN TERRITORY: Kintore Cave (K2), Katherine (Hamilton- Smith unpublished data). Undetermined Emesinae genus and species, Tp, Gp. QUEENSLAND: Crazy Cracks Cave, Jacks Gorge, Broken River (T. Moulds unpublished data); Not Another Frig Tree Crave, Jacks Gorge, Broken River (T. Moulds unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data). Undetermined genus and species, Tp, Gp. NORTHERN TERRITORY: Cutta Cutta Cave (K1), Katherine (Hamilton-Smith unpublished data); QUEENSLAND: Queenslander Cave (CH15), Queenslander Tower (CH5246) Chillagoe (T. Moulds unpublished data); Trezkinn Cave (CH14), Chillagoe (T. Moulds unpublished data); Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data). Undetermined genus and species, Tp, Gp?. QUEENSLAND: Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data). 22 Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Undetermined Family Undetermined genus and species, QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton- Smith unpublished data). Order Neuroptera Myrmeleontidae Aeropteryx sp., Tp, Gp. SOUTH AUSTRALIA: McKinley’s Daughter’s Cave (F175), Flinders Ranges (T. Moulds unpublished data); Moro Bat Cave (F47), Flinders Ranges (T. Moulds unpublished data); Unnamed cave, Brachina Gorge, Flinders Ranges (T. Moulds unpublished data); Unnamed bat cave, Chambers Gorge, Flinders Ranges (T. Moulds unpublished data); Unnamed cave, Chambers Gorge, Flinders Ranges (T. Moulds unpublished data); Unnamed mine, Weetootla Gorge, Gammon Ranges (T. Moulds unpublished data). Myrmeleontinae sp., Tp?. QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith unpublished data). Undetermined Family Undetermined genus and species, QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer’s Mountain (Hamilton-Smith unpublished data). Order Coleoptera Anobtidae (Ptinidae) Ptinus exulans Erichson, Tp1, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith 1967); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Island Cave (CL6), Cliefden (Hamilton-Smith 1967); Jenolan Caves (Hamilton-Smith 1967); Willi Willi Bat Cave (WW1), Willi Willi (Hamilton-Smith 1967); Bungonia various caves (Eberhard 1998); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Colong Main Cave (CG1), Colong (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Warbla Cave (N1) (Richards 1971); Murrawijinie No. 1 Cave (N7) (Richards 1971); Murra-El-Elevyn Cave (N47) (Hamilton-Smith 1967; Richards 1971); Firestick Cave (N70) (Richards 1971); Dingo Cave (N160) (Richards 1971); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith 1967; Bellati et al. 2003); Blanche Cave (U4), Naracoorte (Hamilton-Smith 1967); VICTORIA: Starlight Cave (WS), Warrnambool (Hamilton-Smith 1967); Clogg’s Cave (EB2), East Buchan (Hamilton-Smith 1967); WESTERN AUSTRALIA: Goosebury Cave (J1), Jurien Bay (Hamilton-Smith 1967). Carabidae Anomotarus subterraneus Moore, Tp, Gp. QUEENSLAND: Riverton Main Cave (RN1), Riverton, southern Queensland (Moore 1967). Cratogaster melus Laporte, Tp?. QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data). Darodilia sp., Tp?. QUEENSLAND: Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith unpublished data). Gnathaphanus pulcher Dejean, Tp?. NORTHERN TERRITORY: Cutta Cutta Cave (K1), Katherine (Hamilton-Smith unpublished data); Kintore Cave (K2), Katherine (Hamilton-Smith unpublished data); QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data). Lecanomerus sp., Gp?. NEW SOUTH WALES: Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith unpublished data). Mecyclothorax ambiguus Erichson, VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton- Smith unpublished data). Proc. Linn. Soc. N.S.W., 125, 2004 23 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Meonis sp., Tp, Gp. QUEENSLAND: Main Mount Etna Cave (E1), Mount Etna (Hamilton-Smith unpublished data). Mystropomus subcostatus Chaudoir, Tp?. QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith unpublished data); Speaking Tube (E7), Mount Etna (Hamilton-Smith unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data); Piglet Help! Help! Cave (E17), Mount Etna (Hamilton-Smith unpublished data). Notonomus angustibasis Sloane, Tp?, Gx. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne (Hamilton-Smith unpublished data). Notospeophonus castaneus castaneus Moore, Tp2. SOUTH AUSTRALIA: Blanche Cave (U4), Naracoorte (Hamilton-Smith 1967); Blackberry Cave (U8), Naracoorte (Hamilton-Smith 1967); Stick Cave (U11), Naracoorte (Moore 1964); Cathedral Cave (U12), Naracoorte (Moore 1964); Fox Cave (U22), Naracoorte (Hamilton-Smith 1967); Haystall Cave (U23), Naracoorte (Hamilton-Smith 1967); Cave Park Cave (U37), Naracoorte (Hamilton-Smith unpublished data); Tantanoola Caves (Hamilton-Smith 1967); VICTORIA: Bat Cave (P6), Portland (Moore 1962); Byaduk Caves, Byaduk (Moore 1962); Panmure Cave (H5), Mount Napier (Moore 1964); Mt Widderin Cave (H1), Skipton (Hamilton-Smith 1967); Snowflake Cave (L1), Glenelg River (Hamilton-Smith 1967); Curran’s Creek Cave (G4), Glenelg River (Hamilton-Smith 1967). Notospeophonus castaneus consobrinus Moore, Tp, Gp. VICTORIA: Spring Creek Cave (B1), Buchan (Hamilton-Smith unpublished data); Moon Cave (B2), Buchan (Hamilton-Smith unpublished data); Mabel Cave (EB1), East Buchan (Hamilton-Smith unpublished data); Wilson’s Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Slocombe’s Cave (BA1), The Basin (Hamilton-Smith unpublished data); Shades of Death Cave (M3), Murrindal (Hamilton-Smith unpublished data); Anticline Cave (M11), Murrindal (Hamilton-Smith unpublished data); SSS Cave (M44), Murrindal (Hamilton-Smith unpublished data). Notospeophonus jasperensis jasperensis Moore, Tp2, Gp. NEW SOUTH WALES: Punchbowl! Cave (WJ8), Wee Jasper (Moore 1964); Pylon 58 Cave (WJ99), Wee Jasper (Moore 1964); Basin Cave (W4), Wombeyan (Hamilton-Smith unpublished data). Notospeophonus jasperensis vicinus Moore, Tp2, Gp. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998). Notospeophonus pallidus Moore, Tp2, Gp?. NEW SOUTH WALES: Childrens Cave (CL12), Cliefden (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Myponga (Moore 1964); NULLARBOR PLAIN: Warbla Cave (N1) (Hamilton-Smith 1967; Richards 1971); Weebubbie Cave (N2) (Richards 1971); Abrakurrie Cave (N3) (Hamilton-Smith 1967; Richards 1971); Koonalda Cave (N4) (Hamilton-Smith 1967; Richards 1971); Koomooloobooka Cave (N6) (Richards 1971); Murrawijinie No.3 Cave (N9) (Richards 1971); Knowles Cave (N22) (Hamilton- Smith 1967; Richards 1971); Mullamullang Cave (N37) (Richards 1971); Joe’s Cave (N39) (Hamilton-Smith 1967; Richards 1971); Moonera Tank Cave (N53) (Richards 1971); Madura Cave (Madura 6 Mile South Cave) (N62) (Richards 1971); Lynch Cave (N60) (Richards 1971). Notospeophonus sp., Tp, Gp?. QUEENSLAND: Viator Main Cave (VR1), Viator Hill (Hamilton- Smith unpublished data). Phloeocarabus sp. Tp?, Gp?. QUEENSLAND: Haunted Cave (CH1), Chillagoe (Hamilton-Smith unpublished data). Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Pogonoglossus sp., Tp, Gp?. NORTHERN TERRITORY: Cutta Cutta Cave (K1), Katherine (Hamilton-Smith unpublished data). Pseudoceneus sp. Tp, Gp?. WESTERN AUSTRALIA: Stockyard Cave (E3), Eneabba (Hamilton- Smith unpublished data). Speotarus lucifugus Moore, Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Moore 1964; Bellati et al. 2003); NULLARBOR PLAIN: Warbla Cave (N1) (Richards 1971); Weebubbie Cave (N2) (Richards 1971); Abrakurrie Cave (N3) (Richards 1971); Koonalda Cave (N4) (Richards 1971); Winbirra Cave (N45) (Richards 1971); Murra-El-Elevyn Cave (N47) (Richards 1971); Cocklebiddy Cave (N48) (Richards 1971); Moonera Tank Cave (N53) (Richards 1971); Lynch Cave (N60) (Richards 1971); Unnamed cave (N139) (Richards 1971). Speotarus princeps Moore, Tp2, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Moore 1964); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). Speotarus sp., Tp, Gp. NULLARBOR PLAIN: Warbla Cave (N1) (Hamilton-Smith unpublished data); Weebubbie Cave (N2) (Hamilton-Smith unpublished data); Murra-El-Elevyn Cave (N47) (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Mount Sims Cave (F7), Walpunda Creek, Flinders Ranges (Hamilton-Smith unpublished data); WESTERN AUSTRALIA: Gooseberry Cave (J1), Jurien Bay (Hamilton-Smith unpublished data). Thenarotes speluncarius Moore, Tp, Gp. NULLARBOR PLAIN: Abrakurrie Cave (N3) (Richards 1971); Koonalda Cave (N4) (Richards 1971); New Cave (N11) (Richards 1971); Lynch Cave (N60) (Richards 1971); Decoration Cave (N84) (Richards 1971); SOUTH AUSTRALIA: Cave No. 1, Buckalowie, Flinders Ranges (Hamilton-Smith unpublished data). Trechimorphus diemenensis Bates, Tp1, Gx. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Jenolan Caves (Moore 1964); VICTORIA: Dalley’s Sinkhole (M35), Murrindal (Hamilton-Smith 1967). Trichosternus vigorsi Gory, Tp? Gx. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne (Hamilton-Smith unpublished data). Undetermined genus and species, NEW SOUTH WALES: Grill Cave (B44), Bungonia (Eberhard and Spate 1995); Belfry Cave (TR2), Timor (James et al. 1976); Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave (T1), Tuglow (Hamilton-Smith unpublished data); QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data); Mount Etna Main Cave (E1), Mount Etna (Hamilton-Smith unpublished data); Cave with the thing that went thump! (E5), Mount Etna (Hamilton-Smith unpublished data). % Undetermined genus and species, Tp, Gp. QUEENSLAND: Barker’s Cave (U34), Undara (Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (B1), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson’s Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Shades of Death Cave (M3), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990). Cryptophagidae Anchicera sp., Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Atomaria sp., Gp. Southern Australia (Hamilton-Smith 1968). Proc. Linn. Soc. N.S.W., 125, 2004 25 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Basin Cave (W4), Wombeyan (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Fox Cave (U22), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Wilson’s Cave (EB4), East Buchan (Hamilton- Smith unpublished data); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data). Curculionidae Mandalotus sp. Gp?. NEW SOUTH WALES: Chalk Cave (B26), Bungonia (Hamilton-Smith unpublished data). Talaurinus sp. Gp?. QUEENSLAND: Johannsen’s Cave (J1-2), Mount Etna (Hamilton-Smith unpublished data). Dermestidae Dermestes ater DeGeer, Tp, Gp. QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton- Smith unpublished data). Undetermined genus and species, Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer’s Mountain (Hamilton- Smith unpublished data); Unidentified cave in southern Queensland (Hamilton-Smith 1967); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). Endomychidae Undetermined genus and species, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith unpublished data). Histeridae Carcinops sp., Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 2001). Saprinus sp., Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith unpublished data); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Clogg’s Cave (EB2), East Buchan (Hamilton-Smith unpublished data). Tomogenius ?ripicola Marseul, Tp, Gb. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Lynch Cave (N60) (Richards 1971); Thylacine Hole (N63) (Richards 1971); Dingo Cave (N160) (Richards 1971). Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998); Ashford Main Cave (AS1), Ashford (Hamilton-Smith unpublished data); Carrai Bat Cave (SC5), Stockyard Creek (Hamilton-Smith unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer’s Mountain (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Sand Cave (Joanna) (U16), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Chimney Cave (BR1), Bat Ridges, Portland (Hamilton-Smith unpublished data); Clogg’s Cave (EB2), East Buchan (Hamilton-Smith unpublished data); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Bat Cave (P6), Portland (Hamilton-Smith unpublished data); WESTERN AUSTRALIA: Gooseberry Cave (J1), Jurien Bay (Hamilton-Smith unpublished data). Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Jacobsoniidae Derolathrus sp., Tp, Gb. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); Various caves in southern Australia (Hamilton-Smith 1967). Undetermined genus and species, Tp, Gb. VICTORIA: Bat Cave (P6), Portland (Hamilton-Smith unpublished data); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). Lathridiidae Corticaria sp., Gp. Southern Australia (Hamilton-Smith 1968); NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Weebubbie Cave (N2) (Hamilton-Smith unpublished data); Abrakurrie Cave (N3) (Hamilton-Smith unpublished data): VICTORIA: Skipton Cave (Mount Widderin Cave) (H1), Mount Napier (Hamilton-Smith unpublished data). Leiodidae Choleva australis, Tp, Gp. QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith unpublished data). Choleva sp., Tp, Gp. NULLARBOR PLAIN: Cocklebiddy Cave (N48) (Richards 1971); Lynch Cave (N60) (Richards 1971). Nargomorphus minusculus Blackburn, Tp1, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith 1967; Bellati et al. 2003); VICTORIA: Anticline Cave (M11), Murrindal (Hamilton-Smith 1967). Pseudonemadus adelaidae Blackburn, Tp, Gp. NEW SOUTH WALES: Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data). Pseudonemadus australis Erichson, Gp. VICTORIA: Chimney Cave (BR1), Bat Ridge, Portland (Hamilton-Smith unpublished data); Bat Cave (P6), Portland (Hamilton-Smith unpublished data); Panmure Cave (H5), Mt Napier (Hamilton-Smith unpublished data). Pseudonemadus integer Portevin, Gp. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne (Hamilton-Smith unpublished data); QUEENSLAND: Speaking Tube (E7), Mount Etna (Hamilton- Smith unpublished data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Mt Widderin Cave (H1), Skipton (Hamilton-Smith unpublished data); Panmure Cave (H5), Mt Napier (Hamilton- Smith unpublished data). Pseudonemadus sp., Gp. Southern Australia (Hamilton-Smith 1968). ?Leiodidae Undetermined genus and species, NEW SOUTH WALES: Basin Cave (W4), Wombeyan (Smith 1982a). Melyridae Heteromastix sp. Tx?, Gx?. NEW SOUTH WALES: Colong Main Cave (CG1), Colong (Hamilton- Smith unpublished data). Merophysiidae Undetermined genus and species, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith 1967). Proc. Linn. Soc. N.S.W., 125, 2004 27 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Pselaphidae Ptilidae Rybaxis? sp., Tp, Gp. NEW SOUTH WALES: Basin Cave (W4), Wombeyan (Hamilton-Smith 1966a); Bungonia various caves (Eberhard 1998). Tyromorphus speciosus King, Tp1. NEW SOUTH WALES: Unidentified cave, Southern Limestone, Jenolan (Hamilton-Smith 1966a); Paradox Cave (J48), Jenolan (Hamilton-Smith unpublished data); QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith 1966a); VICTORIA: Anticline Cave (M11), Murrindal (Hamilton-Smith 1966a). Undetermined genus and species, Gp. QUEENSLAND: Rope Ladder Cave, Mingella (Weinstein and Slaney 1995). Undetermined genus and species, Tp, Gp. VICTORIA: Wilson’s Cave (EB4), East Buchan (Hamilton-Smith unpublished data). Achosia lanigera Deane, Tp?, Gp. VICTORIA: Wilsons Cave (EB4), East Buchan (Hamilton-Smith unpublished data). Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne (Hamilton-Smith unpublished data). Rhizophagidae Undetermined genus and species, Gp. QUEENSLAND: Rope Ladder Cave (FR2), Mingella, Fanning River (Weinstein and Slaney 1995). Scarabaeidae Aulacopris maximus Matthews, Tp1, Gb. NEW SOUTH WALES: Yessabah Bat Cave (YE1), Yessabah (Waite 1898); Unknown cave in Coorabakh National Park (formerly part Lansdowne State Forest), Taree (Williams 2003). Aulacopris reichei White, Tp1, Gp. NEW SOUTH WALES: Yessabah Bat Cave (YE1), Yessabah (Lea 1923); Unknown cave, Mosman (Fricke 1964). Amphistomus accidatus Matthews, Tx, Gp. QUEENSLAND: Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data). Saprosites mendax Blackburn, Gp. SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data). Undetermined genus and species, Gp. NEW SOUTH WALES: Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data). Silphidae Ptomaphila lachrymosa Schreibers, VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton- Smith unpublished data). Staphylinidae 28 Myotyphlus jansoni Matthews, Tp1, Gp. NEW SOUTH WALES: Unidentified cave, Southern Limestone, Jenolan (Hamilton-Smith and Adams 1966); Paradox Cave (J48), Jenolan (Hamilton- Smith unpublished data); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith and Adams 1966); Bat Cave (P6), Portland (Hamilton-Smith 1967). Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Philonthus parcus Sharp, Gp?. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (Moore 1964); Church Cave (WJ31), Wee Jasper (Moore 1964); VICTORIA: Starlight Cave (W5), Warmambool (Hamilton-Smith unpublished data). Quedius luridipennis Macleay, Tp?, Gp. NULLARBOR PLAIN: Abrakurrie Cave (N3) (Richards 1971). Quedis sp., Gp. Southern Australia (Hamilton-Smith 1968); NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); Signature Cave (WJ7) (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Wilsons Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998); Colong Main Cave (CG1), Colong (Hamilton-Smith unpublished data); Dip Cave (WJ1), Wee Jasper (Hamilton-Smith unpublished data); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Abrakurrie Cave (N3) (Hamilton- Smith unpublished data); QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data); Camooweal Four Mile East Cave (Camooweal Cave, Four Mile Cave) (C13), Camooweal (Hamilton-Smith unpublished data); VICTORIA: Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Grassmere Cave (W6), Warrnambool (Hamilton-Smith unpublished data). Undetermined genus and species , Tp?, Gx?. QUEENSLAND: Four Mile Cave (C14), Camooweal (Hamilton-Smith unpublished data). Tenebrionidae Adelium sp. Tp? Gp?. SOUTH AUSTRALIA: Sand Cave (Joanna) (U16), Naracoorte (Hamilton- Smith unpublished data). Alphitobius laevigatus Fabricius, Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 2001). Alphitobius diaperinus Panzer, Tp?, Gp?. QUEENSLAND: Bat Cleft (E6), Mount Etna (Hamilton- Smith unpublished data). Brises acuticornis Pascoe, Tp1, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith 1967); NORTHERN TERRITORY: Unknown bat caves, 15 km south of Alice Springs (Mathews 1986); NULLARBOR PLAIN: Warbla Cave (N1) (Richards 1971); Weebubbie Cave (N2) (Hamilton-Smith 1967; Richards 1971); Abrakurrie Cave (N3) (Richards 1971); Koonalda Cave (N4) (Richards 1971); Koomooloobooka Cave (N6) (Richards 1971); Murrawijinie No. 1 Cave (N7) (Richards 1971); Murrawijinie No. 2 Cave (N8) (Richards 1971); Murrawijinie No.3 Cave (N9) (Richards 1971); White Wells Cave (N14) (Hamilton-Smith 1967; Richards 1971); Unnamed cave (N33) (Mathews 1986); Mullamullang Cave (N37) (Hamilton-Smith 1967; Richards 1971); Winbirra Cave (N45) (Richards 1971); Nurina Cave (N46) (Richards 1971); Murra-El- Elevyn Cave (N47) (Hamilton-Smith 1967; Richards 1971); Cocklebiddy Cave (N48) (Richards 1971); Pannikin Plain Cave (N49) (Richards 1971); Moonera Tank Cave (N53) (Richards 1971); Tommy Grahams Cave (N56) (Richards 1971); Lynch Cave (N60) (Richards 1971); White Wells Blowhole (N61) (Hamilton-Smith 1967); Madura Cave (Madura Six Miles South Cave) (N62) (Hamilton-Smith 1967; Richards 1971); Thylacine Hole (N63) (Richards 1971); Firestick Cave (N70) (Richards 1971); Old Homestead Cave (N83) (Richards 1971); Diprose No.1 Cave (N96) (Hamilton-Smith 1967); Diprose No.3 Cave (N98) (Hamilton-Smith 1967); Snake Pit (N133) (Richards 1971); Unnamed cave (N149) (Richards 1971); Dingo Cave (N160) (Richards 1971); Swallow Cave, Cocklebiddy (Mathews 1986); SOUTH AUSTRALIA: Punyelroo Cave (M1), Swan Proc. Linn. Soc. N.S.W., 125, 2004 29 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Reach Murray Plains (Hamilton-Smith 1967); Clara St. Dora Cave (F4), Buckalowie, Flinders Ranges (Hamilton-Smith 1967); Unknown bat cave, Wilpena Pound, Flinders Ranges (Mathews 1986); Wooltana Cave (F9), Flinders Ranges (Mathews 1986). Brises katherinae Matthews, Tp, Gp. NORTHERN TERRITORY: Cutta Cutta Cave (K1), Katherine (Mathews 1986; Hamilton-Smith et al. 1989); Kintore Cave (K2), Katherine (Mathews 1986); Three Mile Cave, Katherine (Mathews 1986); 16 Mile Cave, Katherine (Mathews 1986). Helea catenulatus? Mail., Gp?. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton- Smith unpublished data). Pterohelaeus piceus Kirby, Tp, Gp?. NEW SOUTH WALES: Island Cave (CL6), Cliefden (Hamilton-Smith unpublished data); Timor Caves, Timor (Hamilton-Smith unpublished data). Pterohelaeus sp., Tp, Gp?. NEW SOUTH WALES: Main Cave (Ballroom Cave) (TR1), Timor (Hamilton-Smith unpublished data); QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); Ilium Cave (E31), Mount Etna (Hamilton-Smith unpublished data). Undetermined genus and species, Gp. NEW SOUTH WALES: Colong Main Cave (CG1), Colong (Hamilton-Smith unpublished data); Moparabah Cave (MP1), Moparabah (Hamilton-Smith unpublished data). Trogidae Omorgus costatus Wiedemann, Tp, Gp?. QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data). Trox alatus Macleay, Tp, Gp. NORTHERN TERRITORY: various caves of the Kimberley region (Hamilton-Smith et al. 1989); Kintore Cave (K2), Katherine (Hamilton-Smith unpublished data). Trox amictus Haaf, Tp, Gp. NULLARBOR PLAIN: Abrakurrie Cave (N3) (Richards 1971); Murrawijinie No. 1 Cave (N7) (Richards 1971); Murrawijinie No.3 Cave (N9) (Richards 1971); Mullamullang Cave (N37) (Richards 1971); Lynch Cave (N60) (Richards 1971); Skink Hole (N82) (Richards 1971); Old Homestead Cave (N83) (Richards 1971); Decoration Cave (N84) (Richards 1971). Trox sp., Tp, Gp. NEW SOUTH WALES: guano caves from northern NSW (Hamilton-Smith 1967); Comboyne C4 Cave, Comboyne (Hamilton-Smith unpublished data); NORTHERN TERRITORY: various bat caves (Hamilton-Smith 1967); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); Arch Cave (U22), Undara (Hamilton-Smith unpublished data); Barker’s Cave (U34), Undara (Hamilton-Smith unpublished data); various bat caves (Hamilton-Smith 1967); Bat Cleft (E6), Mount Etna (Hamilton-Smith unpublished data). Undetermined Family Undetermined genus and species, Tp, Gp. QUEENSLAND: Clam Cave (CH26), Walkunder Tower, Chillagoe (Matts 1987); Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith unpublished data); Bat Cleft (E6), Mount Etna (Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (B1), Buchan (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990). Order Siphonaptera Ischnopsyllidae Porribius sp., Tx, Gx, P. NULLARBOR PLAIN: Mullamullang Cave (N37) (Richards 1971); Warbla Cave (N1) (T. Moulds unpublished data). 30 Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Order Diptera Anthomyiidae Undetermined genus and species, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith unpublished data); Southern Australia (Hamilton-Smith 1968). Cecidomyidae Undetermined genus and species, Gp. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data). Ceratopogonidae Culicoides sp., Tp, Gx?. QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data). Cypselosomatidae Clisa (Cypselosoma) australe McAlpine, Tp1, Gb. NEW SOUTH WALES: Carrai Bat Cave (SCS), Stockyard Creek (McAlpine 1966, 1993). Chloropidae Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 2001). Chironomidae Diplocladius multiserialis Freeman, Tx, Gx?. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data). Podonomus sp., Tx, Gx?. NEW SOUTH WALES: Mammoth Cave (J13), Jenolan (Hamilton-Smith unpublished data); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data). Polypedilum watsoni Freeman, Tx, Gx?. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data). Tanytarus sp. Tx, Gx?. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data). Undetermined genus and species,Tx, Gx?. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Carrai Bat Cave (SC5), Stockyard Creek (Hamilton-Smith unpublished data); Main Cave (Ballroom Cave) (TR1), Timor (Hamilton-Smith unpublished data); QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data). Chyromyidae Aphaniosoma sp., Tp, Gp. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (McAlpine 1966); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data). Dolicopodidae Sympycnus sp., Tx?, Gx?. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data). ?Drosophilidae Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 2001). Proc. Linn. Soc. N.S.W., 125, 2004 31 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Fanniidae Fannia sp., Tp, Gx. VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). Milichiidae Undetermined genus and species, Tp? Gx?. NEW SOUTH WALES: Church Cave (W31), Wee Jasper (Hamilton-Smith unpublished data); QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer’s Mountain (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), Riverton (Hamilton- Smith unpublished data). Muscidae Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Grimes Cave (CI53) (Humphreys and Eberhard 2001); Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 2001); NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Bat Cave (P6), Portland (Hamilton-Smith unpublished data). Undetermined genus and species, Gx?. QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data). Mycetophilidae Exechia pullicauda Skuse, Tp?, Gp?. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data). Exechia sp., Tp?, Gp?. NEW SOUTH WALES: The Drum Cave (B13), Bungonia (Hamilton-Smith unpublished data). Nycteribiidae Basilia halei Musgrave, Tp. Gx, P. WESTERN AUSTRALIA: Wilgie-Mia Ochre Mine (MIS34), Cue (Hamilton-Smith unpublished data). Basilia musgravei Theodor, Tp, Gx, P. QUEENSLAND: Chillagoe Caves (Maa 1971); Pink’s Cave (CH20), Chillagoe (Maa 1971); Viator Cave (VR4), Viator Hill (Maa 1971); Mount Etna Main Cave (E1), Mount Etna (Maa 1971). Basilia techna Maa, Tp, Gx, P. NORTHERN TERRITORY: Red Bank Mine (Maa 1971); QUEENSLAND: Lawn Hill (Maa 1971); Olsen’s Caves (Maa 1971). Basilia troughtoni Musgrave, Tp, Gx, P. WESTERN AUSTRALIA: Brown Bone Cave (SH17), South Hill River (Hamilton-Smith unpublished data). Basilia sp., Tp, Gx, P. QUEENSLAND: Tea-Tree Cave (CH43), Chillagoe (Hamilton-Smith unpublished data). Eremoctenia vandeuseni Maa, Tp, Gx, P. QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Joanna Bat Cave (U38), Naracoorte (Hamilton-Smith unpublished data). Nycteribia allotopa meridiana Maa, Tp, Gx, P. QUEENSLAND: Pink’s Cave (CH20), Chillagoe (Maa 1971); Mount Etna Main Cave (E1), Mount Etna (Maa 1971). Nycteribia alternata Maa, Tp, Gx, P. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Maa 1971); Back Creek Mine (Maa 1971); Bonalbo Colliery (Maa 1971); Rise and Shine Mine (Maa 1971); Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); QUEENSLAND: Phoenician Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Mine (Maa 1971); Pink’s Cave (CH20), Chillagoe (Maa 1971); Chillagoe Caves (Maa 1971); Mount Etna Main Cave (E1), Mount Etna (Maa 1971). Nycteribia parilis vicaria Maa, Tp, Gx, P. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Maa 1971); Back Creek Mine (Maa 1971); Bonalbo Colliery (Maa 1971); Bullio Cave (W2), Wombeyan (Maa 1971); Carrai Bat Cave (SC5), Stockyard Creek (Maa 1971); Chietmore Cave (Maa 1971); Colong Caves (Maa 1971); Endless Cave (Maa 1971); Fig Tree Cave (W148), Wombeyan (Maa 1971); Gable Cave (CL7), Cliefden (Maa 1971); North Sydney railway tunnel (Maa 1971); Piano Cave (WA12), Walli (Maa 1971); Prospect Tunnel (Maa 1971); Puchbowl Cave (WJ8), Wee Jasper (Maa 1971); Wee Jasper Caves (Maa 1971); Rise and Shine Mine (Maa 1971); Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); Wombeyan Caves (Maa 1971); Yessabah Bat Cave (YE1), Yessabah (Maa 1971); Bungonia various caves (Maa 1971; Hamilton-Smith 1972; Eberhard 1998); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith unpublished data); Pink’s Cave (CH20), Chillagoe (Maa 1971); Tea-Tree Cave (CH43), Chillagoe (Hamilton-Smith unpublished data); Chillagoe Caves (Maa 1971); Phoenician Mine (Maa 1971); Pilkington Cave, Rockhampton (Maa 1971); Mount Etna Main Cave (E1), Mount Etna (Maa 1971); Cowie Bay Cave, Cape York Peninsula (Maa 1971); SOUTH AUSTRALIA: Hodges Cave (Joanna Bat Cave) (U38), Naracoorte (Maa 1971); Tomato-Stick Cave (U10), Naracoorte (Maa 1971); Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (B1), Buchan (Maa 1971); Grassmere Cave (W6), Warrmambool (Maa 1971); Mabel Cave (EB1), East Buchan (Hamilton- Smith unpublished data). Penicillidia oceanica Bigot, Tp, Gx, P,. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Maa 1971); Back Creek Mine (Maa 1971); Belfery [sic] Cave (TR2), Timor (Maa 1971); Bonalbo Colliery (Maa 1971); Carrai Bat Cave (SC5), Stockyard Creek (Maa 1971); Chietmore Cave (Maa 1971); Colong Main Cave (CG3), Colong (Maa 1971); Drum Cave (B13), Bungonia (Maa 1971); Endless Cave (Maa 1971); Fig Tree Cave (W148), Wombeyan (Maa 1971); Gable Cave (CL7), Cliefden (Maa 1971); Prospect Tunnel (Maa 1971); Punchbowl Cave (WJ8), Wee Jasper (Maa 1971); Rise and Shine Mine (Maa 1971); Timor Main Cave (TR1), Timor (Maa 1971); Waterfall Gold Mine (Maa 1971); Yessabah Bat Cave (YE1), Yeassabah (Maa 1971); Bungonia various caves (Maa 1971; Hamilton-Smith 1972; Eberhard 1998); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); QUEENSLAND: Chillagoe (Maa 1971); Holy Jump Lava Cave (BM1), Bauer’s Mountain (Maa 1971); Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Maa 1971); VICTORIA: Greenhouse Cave (B3), Buchan (Maa 1971); Spring Creek Cave (B1), Buchan (Hamilton-Smith unpublished data). Penicillidia setosala Maa, Tp, Gx, P. NEW SOUTH WALES: Fingal Point Cave (Maa 1971); Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); QUEENSLAND: Phoenician Mine (Maa 1971). Penicillidia tectisentis Maa, Tp, Gx, P. NEW SOUTH WALES: Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); QUEENSLAND: Mount Etna Main Cave (E1), Mount Etna (Maa 1971); SOUTH AUSTRALIA: Tomato-Stick Cave (U10), Naracoorte (Maa 1971); Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Grassmere Cave (W6), Warrnambool (Maa 1971). Penicillidia vandeuseni Maa, Tp, Gx, P. NEW SOUTH WALES: Fingal Point Cave (Maa 1971); Rise and Shine Mine (Maa 1971); Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); QUEENSLAND: Chillagoe Caves (Maa 1971); Royal Arch Cave (CH9), Chillagoe (Maa 1971); Tea-Tree Cave (CH43), Chillagoe (Hamilton-Smith unpublished data); Mount Etna Main Cave (E1), Mount Etna (Maa 1971); SOUTH AUSTRALIA: Hodges Cave (Maa 1971). Phthiridium curvatum Theodor, Tp, Gx, P. NEW SOUTH WALES: Bonalbo Colliery (Maa 1971); Bullio Cave (W2), Wombeyan (Maa 1971); Cliefden (Maa 1971); Junction Cave (W152), Wombeyan (Maa 1971); Rise and Shine Mine (Maa 1971); Tanja Gold Mine (Maa 1971); Proc. Linn. Soc. N.S.W., 125, 2004 . 33 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Morapabah Cave (Temagog Cave) (MP1), Morapabah (Maa 1971); Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); QUEENSLAND: Mount Etna Main Cave (E1), Mount Etna (Maa 1971); VICTORIA: Mabel Cave (EB1), East Buchan (Maa 1971). Undetermined genus and species, Tp, Gx, P,. QUEENSLAND: Flogged Horse Cave (Cammoo Cave) (J83), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Asbestos mine near Arkaba, Flinders Ranges (Hamilton-Smith unpublished data); Drop Drop Cave (L29), Lower south east (Hamilton-Smith unpublished data). Phoridae Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998); The Drum Cave (B13), Bungonia (Hamilton-Smith unpublished data); Colong ‘Main Cave (CG3), Colong (Eberhard and Spate 1995); River Cave (CP6), Cooleman Plains (Eberhard and Spate 1995); Main Cave (Ballroom Cave) (TR1), Timor (Hamilton-Smith unpublished data); Basin Cave (W4), Wombeyan (Hamilton-Smith unpublished data); Urinary Tract Cave (W78), Wombeyan (Eberhard and Spate 1995); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data); Signature Cave (WJ7), Wee Jasper (Hamilton-Smith unpublished data); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data); Dogleg Cave (WJ10), Wee Jasper (Eberhard 1993); Humicrib Cave (WJ34), Wee Jasper (Eberhard 1993); Pylon 58 Cave (WJ99), Wee Jasper (Eberhard 1993); NULLARBOR PLAIN: Abrakurrie Cave (N3) (Richards 1971); Murra-El-Elevyn Cave (N47) (Richards 1971); QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); Fox Cave (U22), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Wilson’s Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). Platypezidae Undetermined genus and species, Gp. NEW SOUTH WALES: The Drum Cave (B13), Bungonia (Hamilton-Smith unpublished data). Psychodidae Phlebotumus sp., Tp, Gx?. QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data). Sergentomyia queenslandi Quate, Tp, Gx?. QUEENSLAND: Haunted Cave (CH1), Chillagoe (Hamilton-Smith unpublished data). Sergentomyia sp., Tp, Gx?. QUEENSLAND: Haunted Cave (CH1), Chillagoe (Hamilton-Smith unpublished data); Donna Cave (CH2), Chillagoe (Hamilton-Smith unpublished data); Royal Arch Cave (CH9), (Hamilton-Smith unpublished data); Royal Arch Cave (CH9), Chillagoe (Hamilton- Smith unpublished data); Trezkinn Cave (CH14), Chillagoe (Hamilton-Smith unpublished data); Keef’s Cave (CH24), Chillagoe (Hamilton-Smith unpublished data); Tea-Tree Cave (CH43), Chillagoe (Hamilton-Smith unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data). Undetermined genus and species, Tp. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); VICTORIA: Cloggs Cave (EB2), East Buchan (Hamilton-Smith unpublished data). Sciaridae Bradysia sp., Tp, Gp?. NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); Willi Willi Bat Cave (WW1), Willi Willi (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Panmure Cave (H5), 34 Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Mount Napier (Hamilton-Smith unpublished data). Lycoriella sp., Tp?, Gp?. NEW SOUTH WALES: Signature Cave (WJ7), Wee Jasper (Hamilton- Smith unpublished data); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Cloggs Cave (EB2), East Buchan (Hamilton-Smith unpublished data). Undetermined genus and species, Tp, Gp?. NEW SOUTH WALES: The Drum Cave (B13), Bungonia (Hamilton-Smith unpublished data); Chalk Cave (B26), Bungonia (Hamilton-Smith unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Colong Main Cave (CG1), Colong (Hamilton-Smith unpublished data); Paradox Cave (J48), Jenolan (Hamilton- Smith unpublished data); Main Cave (Ballroom Cave) (TR1), Timor (Hamilton-Smith unpublished data); Signature Cave (WJ7), Wee Jasper (Hamilton-Smith unpublished data); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); Basin Cave (W4), Wombeyan (Hamilton-Smith unpublished data); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Abrakurrie Cave (N3) (Hamilton-Smith unpublished data); QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); VICTORIA: Clogg’s Cave (EB2), East Buchan (Hamilton-Smith unpublished data); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). Sphaeroceridae Leptocera sp., Tp, Gp. VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Paradox Cave (J48), Jenolan (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); VICTORIA: Wilson’s Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Panmure Cave (HS), Mount Napier (Hamilton-Smith unpublished data); Grassmere Cave (W6), Warrnambool (Hamilton-Smith unpublished data). Streblidae Ascodipteron archboldi Maa, Tp, Gx, P. QUEENSLAND: Chillagoe Caves (Maa 1971); Gordon Mine (Maa 1971). Ascodipteron australiense Muir, Tp, Gx, P. QUEENSLAND: Mount Etna Main Cave (E1), Mount Etna (Maa 1971). Brachytarsina amboinensis uniformis Maa, Tp, Gx, P. NEW SOUTH WALES: Bungonia various caves (Maa 1971; Hamilton-Smith 1972; Eberhard 1998); Ashford Main Cave (AS1), Ashford (Maa 1971); Back Creek Mine (Maa 1971); Belfery [sic] Cave (TR2), Timor (Maa 1971); Carrai Bat Cave (SC5), Stockyard Creek (Maa 1971); Drum Cave (B13), Bungonia (Maa 1971); Endless Cave (Maa 1971); Fig Tree Cave (W148), Wombeyan (Maa 1971); Prospect Tunnel (Maa 1971); Rise and Shine Mine (Maa 1971); Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); Yessabah Bat Cave (YE1), Yessabah (Maa 1971); QUEENSLAND: Chillagoe (Maa 1971); Viator Cave (VR4), Viator Hill, southern Queensland (Maa 1971); Mount Etna Main Cave (E1), Mount Etna (Maa 1971). Brachytarsina verecunda Maa, Tp, Gx, P. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Maa 1971); Bonalbo Colliery (Maa 1971); Cliefden (Maa 1971); Drum Cave (B13), Bungonia (Maa 1971); Humicrib Cave (WJ34), Wee Jasper (Maa 1971); Tanja Gold Mine (Maa Proc. Linn. Soc. N.S.W., 125, 2004 35 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 1971); Morapabah Cave (Temagog Cave) (MP1), Morapabah (Maa 1971); Timor Caves (Maa 1971); Wee Jasper (Maa 1971); Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); Bungonia various caves (Maa 1971; Hamilton-Smith 1972; Eberhard 1998); QUEENSLAND: Mount Etna Main Cave (E1), Mount Etna (Maa 1971). Undetermined genus and species, Tp, Gx, P. QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); Flogged Horse Cave (Cammoo Cave) (J83), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data). ?Therevidae Tipulidae Undetermined genus and species, Gx?. NEW SOUTH WALES: Colong Main Cave (CG1), Colong (Hamilton-Smith unpublished data). Undetermined genus and species, Gx?. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data). Trichoceridae Undetermined genus and species, Tp, Gp?. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Snowflake Cave (L1), Genelg River (Hamilton-Smith unpublished data); Fox Cave (U22), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Mount Widderin Cave (H1), Skipton (Hamilton-Smith unpublished data). Undetermined Family Undetermined genus and species, Tp, Gx. NEW SOUTH WALES: Cliefden Main Cave (CL1), Cliefden (Hamilton-Smith unpublished data); Gable Cave (CL7), Cliefden (Hamilton-Smith unpublished data); Tuglow Cave (T1), Tuglow (Hamilton-Smith unpublished data); QUEENSLAND: Bat Cleft (E6), Mount Etna (Hamilton-Smith unpublished data); Speaking Tube (E7), Mount Etna (Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (B1), Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990). Undetermined genus and species, Tp, Gp. VICTORIA: Bat Cave (P6), Portland (Hamilton-Smith unpublished data); Mt Widderin Cave (H1), Skipton (Hamilton-Smith unpublished data). Order Lepidoptera Noctuidae Agrotis infusa Boisduval, Tx. VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). Persectania ewingii Westwood, Tx. VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton- Smith unpublished data). Pseudaletia australis Feaud., Tx. VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). Pyralidae 36 Pyralis manihotalis Guenée, Tp1, Gp. QUEENSLAND: Rope Ladder Cave (FR2), Mingella, Fanning River (Weinstein and Edwards 1994). Pyralinae or Epipaschiinae sp., Gp. CHRISTMAS ISLAND (Indian Ocean): Smiths Cave (CI9) (Humphreys and Eberhard 2001). Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS Tineidae Lindera tessellatella Blanchard, Gb?. NEW SOUTH WALES: Humicrib Cave (WJ34), Wee Jasper (Eberhard and Spate 1995). Monopis crocicapitella Clemens, Tp, Gb. NEW SOUTH WALES: Drum Cave (B13), Bungonia (Eberhard 1998); Grill Cave (B44), Bungonia (Eberhard 1998); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton- Smith unpublished data). Monopis sp., Gb. NEW SOUTH WALES: Gable Cave (CL7), Cliefden (Eberhard and Spate 1995); Colong Main Cave (CG3), Colong (Eberhard and Spate 1995); Jenolan undetermined cave (Gibian et al. 1988); Basin Cave (W4), Wombeyan (Smith 1982b); Undetermined caves, Wombeyan (Dew 1963); Signature Cave (WJ7), Wee Jasper (Hamilton-Smith unpublished data); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data); Dogleg Cave (WJ10), Wee Jasper (Eberhard 1993); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); Humicrib Cave (WJ34), (Eberhard 1993); Carey’s Cave (WJ100), Wee Jasper (Eberhard 1993); NULLARBOR PLAIN: Abrakurrie Cave (N3) (Richards 1971); Koonalda Cave (N4) (Richards 1971); Mullamullang Cave (N37) (Richards 1971); Cocklebiddy Cave (N48) (Richards 1971); Moonera Tank Cave (N53) (Richards 1971); Thylacine Hole (N63) (Richards 1971); Old Homestead Cave (N83) (Richards 1971); Dingo Cave (N160) (Richards 1971). Undetermined genus and species, Gb. CHRISTMAS ISLAND (Indian Ocean): Smiths Cave (CI9) (Humphreys and Eberhard 2001); Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 2001); NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (Hamilton-Smith unpublished data); Cliefden Main Cave (CL1), Cliefden (Hamilton-Smith unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); QUEENSLAND: Rope Ladder Cave (FR2), Mingella, Fanning River (Weinstein and Slaney 1995); Queenslander Tower (CH5246), Chillagoe (Matts 1987); Spring Tower (CH5223-5), Chillagoe (Matts 1987); Donna Tower (CH5155), Chillagoe (Matts 1987); Royal Arch Tower (CH5158-9), Chillagoe (Matts 1987); Tea Tree Tower (CH5137), Chillagoe (Matts 1987); Ryan Imperial Tower (CH5239), Chillagoe (Matts 1987); Wallaroo Tower (CH5201), Chillagoe (Matts 1987); Tower of London Cave (CH5) Chillagoe (Matts 1987); Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data); Holy Jump Lava Cave (BM1), Bauer’s Mountain (Hamilton-Smith unpublished data); VICTORIA: Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Grassmere Cave (W6), Warrnambool (Hamilton-Smith unpublished data). | Undetermined Family Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Smiths Cave (CI9) (Humphreys and Eberhard 2001); Swiflet Cave (CI30) (Humphreys and Eberhard 2001); Managers Alcove (CI50) (Humphreys and Eberhard 2001); Grimes Cave (CI53) (Humphreys and Eberhard 2001); Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 2001). Undetermined genus and species, NULLARBOR PLAIN: Abrakurrie Cave (N3) (Hamilton-Smith unpublished data). Order Hymenoptera Braconidae Apanteles ?carpatus Say, Tp1, Gp. NEW SOUTH WALES: Humidicrib Cave (WJ34), Wee Jasper (Eberhard and Spate 1995); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). Proc. Linn. Soc. N.S.W., 125, 2004 37 CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Apanteles sp., Tp, Gp. NEW SOUTH WALES: Church Cave (W31), Wee Jasper (Hamilton-Smith unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data). Undetermined genus and species. Tp?. QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer’s Mountain (Hamilton-Smith unpublished data). Formicidae Amblyopone australis Erichson, VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). Iridomyrmex purpureus Smith, Tx, Gx. SOUTH AUSTRALIA: Eregunda Mine near Blinman, Flinders Ranges (T. Moulds unpublished data). Oligomyrmex sp., Tp?, Gx?. QUEENSLAND: Crazy Cracks Cave, Jacks Gorge, Broken River (T. Moulds unpublished data). Pachycondyla sp., Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 2001). Undetermined genus and species, NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith unpublished data); Spring Cave, Mount Surprise (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Ichneumonidae Undetermined Cryptinae genus and species, Gp?. NEW SOUTH WALES: undetermined caves (Hamilton-Smith 1967); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); Spring Creek Cave (B1), Buchan (Hamilton-Smith unpublished data); Wilson’s Cave (EB4), East Buchan (Hamilton-Smith unpublished data). Myrmaridae Gonatocerinae sp., Gp?. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). Undetermined Family 38 Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998). Undetermined genus and species, Gp?. NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); Willi Willi Bat Cave (WW1), Willi Willi (Hamilton-Smith unpublished data); VICTORIA: Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). Proc. Linn. Soc. N.S.W., 125, 2004 T. MOULDS ACKNOWLEDGEMENTS This work was only possible due to the financial support of the Department of Environment and Heritage, South Australia, and the University of Adelaide. Thanks to Stefan Eberhard and Sue White for providing the stimulus for writing this paper. Many thanks to Elery Hamilton-Smith whose critical comments and unlimited access to his personal unpublished records greatly improved this paper. Thank you to Mike Gray, Courtenay Smithers and Judy Bellati who brought additional references to my attention. I also wish to thank John Jennings and Andy Austin for editonal comments. REFERENCES Beier, M. (1967). Some Pseudoscorpionidea from Australia, chiefly from caves. Australian Zoologist 14, 199-205. Beier, M. (1968). Some cave-dwelling Pseudoscorpionidea from Australia and New Caledonia. Records of the South Australian Museum 15, 757-765. Beier, M. (1975). Neue Pseudoskorpione aus Australien und Neu-Guinea. Annalen des Naturhistorischen Museums Wien 78, 203-213. Beier, M. 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Transactions of the Royal Society of South Australia 86, 147- 154. Womersley, H. (1963b). A new larval Neotrombidium (Acarina: Leeuwenhoekidae) from bat guano. Records of the South Australian Museum 14, 473-476. Yen, A. L. and Milledge, G. A. (1990). Invertebrates of the Buchan-Murrindal area cave systems. Melbourne, Department of Conservation and Environment, Pp. 1-38. 4] CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST Zumpt, F. and Patterson, P. M. (1952). Three new parasitic mites from the Ethiopian region (Acarina: Laelaptidae). Journal of the Entomological Society of Southern Africa 15, 159-164. Proc. Linn. Soc. N.S.W., 125, 2004 A Devonian Brachythoracid Arthrodire Skull (Placoderm Fish) from the Broken River Area, Queensland GAVIN C. YOUNG Department of Earth and Marine Sciences, Australian National University, Canberra ACT 0200 gyoung @ geology.anu.edu.au Young, G.C. (2004). A Devonian brachythoracid arthrodire skull (placoderm fish) from the Broken River area, Queensland. Proceedings of the Linnean Society of New South Wales 125, 43-56. An incomplete brachythoracid arthrodire skull acid-prepared from the Devonian limestones of the Broken River area of Queensland is described as Doseyosteus talenti gen. et sp. nov. It supposedly comes from strata dated by conodonts as late Early Devonian in age (Emsian stage), but shows several derived features of the skull, typical of Middle-Late Devonian brachythoracids, and not seen in any arthrodire from the Emsian limestones of the Burrinjuck area of NSW. The alignment with conodont zones of stratigraphic subdivisions of the Burrinjuck sequence is revised. Published information on the provenance and age of all previously described placoderm taxa from Broken River is reviewed and amended. The new taxon may be most closely related to Late Devonian (Frasnian) brachythoracids from Iran and the Gogo Formation of Western Australia. Manuscript received 27 May 2003, accepted for publication 22 Oct 2003. KEYWORDS: Placoderm fishes, Arthrodira, Brachythoraci, Broken River, Devonian, new genus Doseyosteus, Queensland. INTRODUCTION Devonian sedimentary rocks, including many marine limestones, are well exposed in the Broken River area of Queensland (Fig. 1). Conodonts form the basis for dating the sedimentary sequence (Mawson and Talent 1989; Sloan et al. 1995). Vertebrate remains reported from this sequence include microfossils from many horizons (De Pomeroy 1996; Turner, Basden and Burrow 2000), and less well known vertebrate macro-remains. The latter include two genera of antiarch placoderms described by Young (1990), a ptyctodont toothplate ascribed to ?Ptyctodus sp. by Turner and Cook (1997), a new species of the brachythoracid arthrodire Atlantidosteus Leliévre 1984 described by Young (2003a), an isolated suborbital plate of another arthrodire illustrated by Turner et al. (2000, fig. 8.7), and jaw remains of an onychodontid (Turner et al. 2000, fig. 5). Undescribed vertebrate macro-remains include various placoderm bones, most of which belong to brachythoracid arthrodires. The Arthrodira is the most diverse order within the class Placodermi, and its major subgroup, the Brachythoraci, comprises nearly 60% of about 170 genera within the Arthrodira (Carr 1995). The brachythoracid arthrodires were one of the most successful groups of early gnathostome fishes (e.g. Young 1986; Janvier 1996). In marine environments of the Late Devonian they included probably the largest predators of their time. The major radiation of brachythoracid subgroups had apparently already occurred by the Middle Devonian, and primitive representatives were already widespread in shallow marine environments of the Early Devonian (e.g. Young et al. 2001; Mark-Kurik and Young 2003), and are important in considering the origins and interrelationships of major brachythoracid subgroups (e.g. Leliévre 1995). The stratigraphic occurrence of various placoderm remains in the Broken River sequence were reviewed by Young (1993, 1995, 1996), De Pomeroy (1995, 1996), and Turner et al. (2000), and they have been mentioned in relation to conodont studies by Sloan et al. (1995). There has been conflicting information published about the provenance of some of the described placoderm taxa. These were collected from the Broken River area many years ago by Professor John Jell, University of Queensland, and sent to Canberra for acid preparation and study. In this paper I describe a new arthrodire skull from this collection, and review the locality information and age determinations for previously described placoderm taxa. DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND A ‘ \ . Nawagiaspis loc OUTSTATIO Viv ~ JESSEY SPRINGS a {vv .y CAINOZOIC [incl. basalt] T-> } L. DEVONIAN- CARBONIFEROUS MIDDLE DEVONIAN = [Broken River Group] wow disconformity GSo54 EARLY DEVONIAN ~ [4 SJ SILURIAN a lnconformity ("—] ?CAMBRIAN-ORDOVICIAN A. FOSSIL FISH LOCALITIES unconformity wey %, eae A sf wf Figure 1. (A) location of the Broken River area in Queensland, Australia; (B) geological map of the collecting area (modified from Turner, Basden and Burrow 2000, fig. 2), showing localities for previously described placoderm taxa, and the specimen described in this paper (ANU V1026). LOCALITY AND AGE OF DESCRIBED PLACODERM TAXA FROM THE BROKEN RIVER AREA Wurungulepis denisoni Young 1990 According to information provided with this specimen, it came from University of Queensland 44 locality L4399 (not L4339, given in error by Young 1990: 45), on the north bank of the Broken River, Grid Reference 640 460 on the Burges 1:100 000 sheet, and was assigned a Middle Devonian (?Eifelian) age within the Broken River Formation (J.S. Jell, letter of 17 April 1980). Judging by the map of the area Proc. Linn. Soc. N.S.W., 125, 2004 published by Sloan et al. (1995: fig. 2), the locality lies within outcrop referred to as ‘undifferentiated Broken River Group’. A ‘Wurungulepis-Atlantidosteus fauna’, of assumed Eifelian age, was listed in the macrovertebrate zonation of Young (1993, 1995, 1996). However De Pomeroy (1995: 480) assigned Wurungulepis to the late Emsian serotinus Conodont Zone (CZ), citing a personal communication of J.A. Talent. This information was repeated by Turner et al. (2000: 498). Later (pers. comm. 28/8/95) J.A. Talent had advised A. Basden that this specimen was collected from the grid reference cited above, situated on a bend of the Broken River in an anticline, in strata which were pre-Dosey Limestone in the sequence, and equivalent to the Bracteata Formation and Lomandra Limestone (spanning the Emsian-Eifelian boundary; Sloan et al. 1995: fig. 3). No conodont data were obtained from the specimen, so its precise position relative to the standard conodont zonation is uncertain. Wurungulepis is an early representative of the asterolepidoid antiarchs, with a high short trunk armour (Young 1990), and was placed within the asterolepidoid clade adjacent to Sherbonaspis, and as sister group to Stegolepis, Asterolepis, Remigolepis and Pambulaspis, by Zhu (1996: fig. 29). As earlier discussed (Young 1990: 48) the initially suggested Eifelian age was consistent with the oldest asterolepid (pterichthyodid) occurrence in Europe, cited as Gerdalepis from the Eifelian of Germany by Denison (1978), although this occurrence is slightly younger (early Givetian) according to Otto (1998: 118). However Gardiner (1994) cited Young (1974) for an older record (Emsian) of the asterolepid antiarchs, but the ‘cf. Pterichthyodes’ mentioned by Young (1974) was based on an erroneous attribution by Hills (1958: 88) to the Early Devonian limestone sequence of an ‘Antiarchan from Taemas’. In fact, the specimen in question came from the overlying Hatchery Creek Formation, of presumed Eifelian age (Fig. 2). This specimen was assigned to the new genus Sherbonaspis by Young and Gorter (1981). Previously, the suggested Emsian age of a pterichthyodid antiarch from the Georgina Basin (Young 1984a) was noted as possibly the oldest occurrence of this group anywhere recorded. G.C. YOUNG costatus partitus patulus serotinus inversus- laticostatus perbonus- gronbergi dehiscens pireneae z < z O > tu a a Oo < wi kindlei sulcatus Figure 2. Proposed alignment with conodont zones of subdivisions of the Early Devonian limestone sequence (Murrumbidgee Group) around Burrinjuck Dam, N.S.W., revised from Basden et al. (2000: fig. 2). Abbreviations for stratigraphic subdivisions are: B - Bloomfield Limestone Member; CB - Cavan Formation; CR - Crinoidal Limestone Member; CU - Currajong Limestone Member; HC - Hatchery Creek Formation; M - Majurgong Formation; R - Receptaculites Limestone Member; SY - Spirifer yassensis Limestone Member; W - Warroo Limestone Member; 1-6 - units of Upper Reef Formation. V1370 — horizon for highest known arthrodire in the sequence. New evidence now indicates that two assemblages may have been mixed in this region (Burrow and Young, in press), with the limestone occurrence yielding the antiarch probably younger than the diverse Proc. Linn. Soc. N.S.W., 125, 2004 | 45 DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND Wuttagoonaspis fauna from underlying sandstones (Young and Goujet 2003). The antiarchs are a major subgroup of the class Placodermi, ranging in age from Early Silurian to latest Devonian. In recent years there has been a significant expansion in our knowledge of the group. A cladistic analysis of their distribution in relation to phylogeny by Young (1984b) involved 22 taxa and 40 characters. In a recent review of antiarch phylogeny, Zhu (1996) noted some 45 genera and 154 species, and his data matrix used 66 characters for 40 genera. The original age assessment of Eifelian for Wurungulepis from Broken River is most consistent with our current knowledge of this large and diverse group. Nawagiaspis wadeae Young 1990 This specimen is recorded from locality BRJ68D (University of Queensland locality L4428; ‘small limestone outcrop on eastern side of gully 1 km upstream from Six Mile yard’), Grid Reference 596 442 on the Burges 1:100 000 sheet, which was assigned a Middle Devonian (?Givetian) age within the Broken River Formation (J.S. Jell, letter of 17 April 1980). Apparently this specimen was found by Dr Mary Wade. Again, De Pomeroy (1995: 480) referred this taxon to the significantly older (late Emsian) serotinus CZ, based on its assigned position within the Bracteata Formation in section Br4 of Sloan et al. (1995, fig. 6). This information was repeated by Turner et al. (2000: 498, 506). However Prof. J.A. Talent’s previous advice to the author (pers. comm. 5/8/92), was that this specimen was considerably younger (ensensis — varcus Zones; late Eifelian - Givetian). Clearly, there was some confusion about which fish specimen was being referred to. Subsequent advice given to A. Basden (pers. comm. 28/8/95), was that N. wadeae came from the bank of Dosey Creek (Grid Reference 616 437), the location of section Br2 within outcrop of the Bracteata Formation (Sloan et al. 1995: fig. 2). The different, and presumably correct, locality information provided with the specimen, as cited above, corresponds to the vicinity of the boundary between the Papilio and Mytton Formations on the map of Sloan et al. (1995: fig. 2). This is consistent with the Givetian age first suggested by J.S. Jell. Nawagiaspis wadeae is another antiarch, originally interpreted as possibly a primitive bothriolepidoid (Young 1990), although in Zhu’s (1996) phylogeny it comes out as a basal asterolepidoid. Apart from primitive Chinese antiarchs, and the erroneous Emsian pterichthyodid occurrence discussed above, the stratigraphic record of this group 46 is Middle-Late Devonian (Gardiner 1994, fig. 32.1). The bothriolepidoid clade had an earlier history in Asia, and apparently expanded its range to most regions of the world in the Givetian (Young 2003b). The confusion about the provenance of this specimen may have resulted from the misconception that it was a recognisable ‘skull’ when collected. Turner et al. (2000) used this term to refer to the type, but the specimen as collected was a largely complete trunk armour, and the incomplete skull, missing its central portion, formed a minor part of the specimen. The whole specimen may have appeared to a non- vertebrate worker to represent a ‘skull’. Such fish remains, when collected in the field, are generally not determinable until after acid preparation (e.g. the type specimen of Atlantidosteus pacifica Young 2003a, before preparation, was assumed to be a ventral plate of the trunk armour, rather than a large suborbital bone from the cheek). A summary list of prepared fish remains from the original J.S. Jell collection was provided to J.A. Talent in 1995 to check on age and locality data. This list mentioned only one skull, the brachythoracid specimen described below, of which locality data provided by J.S. Jell are almost the same as stated by Sloan et al. (1995) for NV. wadeae. Thus it seems that the specimen described below, previously listed as a ‘skull’, has been confused with the type of N. wadeae, leading to erroneous locality and age information being given in De Pomeroy (1995), Sloan et al. (1995), and Turner et al. (2000). In the context of the global distribution in time and space of this major placoderm subgroup (see above), it is almost certain that Nawagiaspis is Middle Devonian in age, and a Givetian age, as first suggested by J.S. Jell, is most consistent with other information about the stratigraphic distribution of the more derived antiarchs. Atlantidosteus pacifica Young 2003a This specimen came from locality BRJ 67B (University of Queensland locality L 4472), Grid Reference 675 485 on the Burges 1:100 000 sheet, described as “Top of ridge to three-quarters way down western slope, west of road between Six Mile Dam and Diggers Creek’ (J.S. Jell, letter of 17 April 1980). This is the locality (with a slightly different grid reference) referred to as ‘Fish Hill’ by Turner et al. (2000: 507). They assigned it a middle Eifelian age (costatus - australis conodont zone), but noted that Sloan et al. (1995) gave a slightly longer partitus - early kockelianus zonal range for the Fish Hill section. This is consistent with the original assignment of a Middle Devonian (?Eifelian) age within the Broken River Formation by Prof. J.S. Jell. This occurrence is Proc. Linn. Soc. N.S.W., 125, 2004 G.C. YOUNG part of the evidence for proposing an Eifelian ‘Wurungulepis-Atlantidosteus fauna’ in the macrovertebrate zonation of Young (1993, 1995, 1996). Doseyosteus talenti gen. et sp. nov. This specimen, described below, was the only one in the J.S. Jell collection lacking a sample number at the time of preparation. It is highly probable that it was a sample collected the year before the other material, and was taken to Canberra separately by Dr P. Jell (J.S. Jell, letter of 17 April 1980). The following locality details, provided by Prof. J.S. Jell (letter of 17 April 1980), indicate that it is the specimen collected from the alternative erroneous locality for Nawagiaspis just discussed: ‘BRJ34 = L 4054. Grid Reference 616 438 Burges 1:100,000 sheet. Western bank of Dosey Creek, 750 m upstream from its junction with Broken River. Base of thick limestone lens in Broken River Formation, Middle Devonian. ? Eifelian’. In a published listing of University of Queensland locality numbers (Turner et al. 2000: 506), UQL4054 is assigned to ‘basal part of limestone, Lomandra/Dosey Limestone, Broken River Group’, with a slightly different grid reference (615 438), but the same locality description as above. However, it is assigned to the Emsian serotinus CZ, citing Sloan et al. (1995). Again, no conodonts were obtained from the sample, and section Br4 through the Bracteata Formation at this locality did not produce identifiable conodonts (Sloan et al. 1995: caption to fig. 6). Nevertheless, these authors (p.5) considered the entire formation to belong to the serotinus CZ, making it equivalent to the upper part of the Burrinjuck (NSW) limestone sequence, which extends from the top of the pirenae CZ (latest Pragian) into the serotinus CZ (the second youngest zone of the late Emsian). It is therefore relevant to make comparisons with the stratigraphic distribution of the diverse arthrodire assemblage described from the Burrinjuck limestone sequence. The described arthrodire fauna from the Burrinjuck sequence (White 1952, 1978; White and Toombs 1972; Young 1979, 1981, in press a, b; Young et al. 2001; Mark-Kurnik and Young 2003) includes 10 genera of brachythoracids, amongst which the most derived taxa (Cathlesichthys and Dhanguura) come from the upper part of the Wee Jasper limestone sequence. Basden et al. (2000, fig. 2) showed the youngest arthrodire skull from the Wee Jasper section (ANU V1370; the holotype of Dhanguura) to come from the uppermost unit 6 of the ‘Upper Reef Formation’ of Young (1969). This specimen is more Proc. Linn. Soc. N.S.W., 125, 2004 advanced than other arthrodires known from the Burrinjuck sequence in possessing several derived characters of the skull, the most obvious being the T- shaped rostral plate, a feature of more derived eubrachythoracids (character 5 of Carr 1991; character 4 of Leliévre 1995). Eubrachythoracids were the most diverse fish group of the Middle and Late Devonian, and the new Broken River brachythoracid described below clearly belongs to this group, with a skull which is more advanced in several respects than any of the known Burrinjuck arthrodires (see below). Gardiner (1994) lists the first occurrence of this grouping (his family Coccosteidae) as Coccosteus Miller 1841 from the Middle Devonian (Eifelian) of Scotland, for which a late Eifelian age is indicated by spores of the devonicus-naumovae zone (V.T. Young 1995). The same species (Coccosteus cuspidatus) is recorded from the Kernave Member of the Narva Formation in the Baltic sequence, although a related brachythoracid ‘Protitanichthys’ occurs a little earlier, and in equivalent strata (costatus CZ) in the Rhenish sequence (Mark-Kurik 2000). However Otto (1997: 115) suggested that remains of early eubrachythoracids (coccosteids) first occur in the early Eifelian of Scotland, Germany, and the Baltic sequence. Dhanguura johnstoni Young (in press a) comes from a horizon about 420 m stratigraphically above the boundary equivalent of the Bloomfield and Receptaculites Members of the Taemas Limestone. A similar horizon high in the limestone sequence has produced the large lungfish Dipnorhynchus cathlesae Campbell and Barwick 1999. The lungfish locality is close to localities L537 and L538 of Pedder et al. (1970) which yielded tetracorals Vepresiphyllum dumosum, Sulcorphyllum pavimentum, Chalcidophyllum vesper and C. gigas. This represents the uppermost ‘tetracoral teilzone’ of the Murrumbidgee Group (Pedder et al. 1970: fig. 4), and is Coral Fauna F in the scheme of Garratt and Wright (1989). These authors considered the succeeding G and H Coral Faunas to overlap, and belong to the late Emsian, rather than Eifelian as previously assessed. Garratt and Wright (1989) also aligned Coral Fauna F from Wee Jasper (and the Sulcor Limestone of northern NSW) with the mid-Emsian inversus CZ (see column 13 of Young 1995, 1996). However Basden et al. (2000: fig. 2) showed the uppermost beds of the limestone sequence at Wee Jasper (containing Coral Fauna F) extending well into the next youngest serotinus CZ. Evidence supporting this (summarised by Basden 2001, table 2.1) derives from reassignment of some of the conodonts from the highest productive sample (C62) in Pedder et al.’s (1970) section 2, referred by them to Polygnathus linguiformis 47 DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND linguiformis, but reassigned to Polygnathus inversus by Klapper and Johnson (1975), and to Polygnathus serotinus (delta morphotype) by Mawson (1987). On the other hand, the age in terms of conodont zone alignment of several constituent members of the Taemas Limestone, as indicated by Basden et al. (2000, fig. 2), seem to be too young, and should be revised downwards on the following evidence. Lindley (2002a: 275) noted that the occurrence of the index species of Coral Fauna D (Chalcidophyllum recessum) in the Currajong Limestone Member indicates that it should be aligned with the dehiscens rather than the perbonus CZ. The overlying Bloomfield Limestone Member may also have lower beds of dehiscens rather than the perbonus CZ age (Basden 2001: table 2.1). The Warroo Limestone Member contains perbonus CZ elements (Nicoll, in Lindley 2002b), and the uppermost Crinoidal Limestone Member in the Taemas sequence may align with both the inversus and the serotinus CZ (Basden 2001: table 2.1). These revised alignments are summarised in Fig. 2. Correlation with the upper part of the Wee Jasper sequence is unclear, because the constituent members of the Taemas Limestone are difficult to recognise in the thicker upper part of the sequence, represented by units 1-6 of Young (1969). If the new arthrodire skull described below from Broken River is of serotinus CZ age, as proposed by Sloan et al. (1995), it is still considerably more derived (see below) than any arthrodire from the Burrinjuck sequence. If correctly dated, this would indicate that derived features characterising the Middle-Late Devonian eubrachythoracid arthrodires had originated at least by late Emsian time. To summarise, it is emphasised that there is no overlap in the arthrodire skull characters just discussed between the Burrinjuck and Broken River limestone sequences, even though the youngest occurrences in the former sequence are also the most derived taxa within the better-documented Burrinjuck arthrodire fauna. For the new taxon described below, this evidence would support either a latest Emsian age (but younger than the Burrinjuck sequence), or an Eifelian age as originally suggested by Prof. J.S. Jell. ABBREVIATIONS The specimen described below (prefix ANU V) is housed in the Earth and Marine Sciences Department, Australian National University, Canberra (GCY Vertebrate Collection). Standard abbreviations for placoderm dermal bones are used in the text and figures, and together with other morphological abbreviations are listed as follows: 48 anth, anterior nuchal thickening; Ce, central plate; cf.Ce, area overlapping Ce plate; cf.M, area overlapping marginal plate; cf.PM, area overlapping postmarginal plate; cf.PtO, area overlapping postorbital plate; cr.im, inframarginal crista; csc, central sensory line canal; d.end, openings of dermal tube for endolymphatic duct; dep, depression; gr.M, groove on Ce plate which received the edge of the marginal plate; ifc.ot, otic branch of infraorbital sensory groove; if.r, infranuchal ridge; if.pt, infranuchal pit; kb, knob-like thickening of inframarginal crista; Iep, lateral consolidated part of skull roof; llc, main lateral line sensory canal; M, marginal plate; mp, middle pitline; mppr, posterior median process of nuchal plate; Nu, nuchal plate; oa.Ce, area overlapped by Ce plate; oa.M, area overlapped by M plate; oa.Nu, area overlapped by Nu plate; orb, orbital notch; Pi, pineal plate; plpr, posterolateral process or lobe on Ce plate; PM, postmarginal plate; pmc, postmarginal sensory groove; pnp, postnuchal process of paranuchal plate; PNu, paranuchal plate; Pp, posterior pitline; PrO, preorbital plate; PtO, postorbital plate; R, rostral plate; soa, subobstantic area; soc, supraorbital sensory canal; th.end, endolymphatic thickening; th.pre, pre-endolymphatic thickening; tnth, transverse nuchal ridge or thickening; vg, vascular grooves. SYSTEMATIC PALAEONTOLOGY Class PLACODERMI McCoy, 1848 Order ARTHRODIRA Woodward, 1891 Suborder BRACHYTHORACI Gross, 1932 Doseyosteus talenti gen. et sp. nov. Name From Dosey Creek, the type locality, and the Greek osteus (bone). The species name recognises Proc. Linn. Soc. N.S.W., 125, 2004 G.C. YOUNG Professor John A. Talent, Macquarie University, who has had a long and distinguished career in Devonian research, including extensive work in the Broken River area of Queensland. Diagnosis A eubrachythoracid arthrodire in which the skull shows an embayed anterior margin of the nuchal plate resulting from overlap by the central plates, the central plates have strong posterolateral lobes separating the nuchal and paranuchal, and a mesial process of the marginal plate extends to the anterior angle of the paranuchal. Subobstantic area of skull extending onto marginal plate. Dermal bones smooth, or ornamented with fine tubercles. Remarks Since only the skull is known, and it is incomplete, several features characterising the derived subgroup “Eubrachythoraci’ are for the present inferred for this new taxon. Definition of the eubrachythoracid arthrodires is discussed by Carr (1991: 379-381) and Long (1995: 55). Thus Doseyosteus talenti gen. et sp. nov. is assumed to have had a T-shaped rostral plate, a posteriorly placed pineal plate separating the preorbitals, a dermal process of the preorbital plate forming the anterodorsal margin of the orbit, and trilobate central plates. The holotype shows a strongly developed posterior thickening of the skull roof, which in the midline is represented by the anterior nuchal thickening. This is much more prominent than the transverse ridge on the posterior margin of the nuchal plate, and is a derived feature seen in coccosteomorph and pachyosteomorph brachythoracids, but generally lacking in Early Devonian taxa, for example the genus Cathlesichthys from Burrinjuck, NSW (Young, in press a). The embayed anterior margin and inferred proportions of the nuchal plate, and the strong posterolateral lobe of the Ce plate, are resemblances to the Late Devonian taxa Eastmanosteus and Golshanichthys, but the former differs in having the posterior pitline well developed on the posterolateral lobe of the central plate, and both forms lack the mesial process of the marginal plate inferred for this new taxon. Material ANU V1026 (holotype), an incomplete skull preserved as two unconnected portions. Locality and Horizon Locality BRJ34 (University of Queensland locality L4054), Grid Reference 616 438, Burges 1:100 000 sheet; western bank of Dosey Creek, 750 m Proc. Linn. Soc. N.S.W., 125, 2004 upstream from its junction with Broken River (J.S. Jell, letter of 17 April 1980; see discussion above). Horizon was described as the ‘base of thick limestone lens in Broken River Formation’, assigned to the Bracteata Formation (Sloan et al. 1995) or the “Lomandra/Dosey Limestone, Broken River Group (Turner et al. 2000). Age: ?late Emsian - Eifelian (see discussion above). Description ANU V1026 represents a large part of the posterolateral region of a brachythoracid skull roof, preserved as two separate portions. The larger portion (Fig. 3A,D) includes parts of the Nu, PNu and Ce plates (Fig. 4A,B), and the right postmarginal corner of the skull is preserved as a separate portion (Figs. 3B,C, 4C,D). The specimen was extracted from the rock in six pieces, but they are well preserved, suggesting that it was broken up before incorporation in the sediment. The nuchal (Nu) plate is represented by most of its right half, including the midline, so its overall shape can be estimated. Midline length of the Nu is about 70 mm. It has an embayed posterior margin, with a prominent posterior median process (mppr, Fig. 4). Except for the posterior lateral corner the right lateral margin of the Nu plate is fairly well displayed on the external surface. The bone is fractured in its middle region, and shows anteriorly that it was both overlapped and underlapped by the central (Ce) plate, a condition also reported in Holonema (Miles 1971). Along the anterior margin of the plate a thin overlapping lamina of the Ce plate has broken away to reveal an extensive overlap area (oa.Ce, Fig. 4B). In unbroken condition the anterior margin of the Nu plate would have been deeply embayed (Fig. 5). On its visceral surface extensive contact faces for the central plates are developed in the normal manner (cf.Ce, Fig. 4A). Other features shown are the prominent infranuchal pits (if-pt) and ridge (if.r) and the transverse nuchal thickening or ridge (tnth). Noteworthy is the strong development of the anterior nuchal thickening (anth), which is relevant to the question of the age of this specimen (see discussion above). This is a derived feature of brachythoracids, and in ANU V1026 is more pronounced than in any Emsian brachythoracid from the Burrinjuck fauna. These have Nu plates which are fairly flat in front of the infranuchal pits. This is the case even in a form like Cathlesichthys, which is derived in having a very strong transverse nuchal ridge (Young in press a). In posterior view ANU V1026 shows that the anterior nuchal thickening is more pronounced than the transverse nuchal ridge, the reverse of the condition in Cathlesichthys. This advanced character is also seen in most Middle-Late Devonian brachythoracids, such 49 DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND Figure 3. Doseyosteus talenti gen. et sp. nov. Holotype (ANU V1026). Larger (A,D) and smaller (B,C) skull portions in external (B,D) and internal (A,C) views. as Golshanichthys, Tafilalichthys, and various Gogo forms (e.g. Leliévre et al. 1981; Leliévre 1991; Miles and Dennis 1979; Long 1988, 1995; Dennis-Bryan 1987). These taxa all resemble the giant Famennian form Dunkleosteus, where the ‘posterior consolidated arch’ of the skull roof (‘PCA’ of Heintz 1932: fig. 13) is a broad thickening running in front of the infranuchal pits, as the main transverse thickening of the skull. In contrast, in the Early Devonian form Cathlesichthys from Burrinjuck, the transverse nuchal ridge located behind the infranuchal pits forms the main thickening supporting the posterior skull margin. The right paranuchal (PNu) plate of Doseyosteus gen. nov. is represented externally by an elongate portion including the mesial margin forming sutures with the Nu and Ce plates (PNu, Fig. 4B). There is also a small broken part of the postnuchal process (pnp). The PNu and Ce plates were also connected by a complex interlocking suture; a broken part around 50 the anterior end of the PNu exposes an overlap area (oa.Ce, Fig. 4B), and the edge of a more extensive contact face is shown on the visceral surface (cf.Ce, Fig. 4A). The endolymphatic thickening forms a broad thickened area mesially (th.end), combining with the thickened portion of the Nu plate (anth). This thickened part of the skull is much more prominent than in primitive brachythoracids like Buchanosteus or Taemasosteus (White 1978; Young 1979). Along the broken edge of the specimen, maximum bone thickness (in the part enclosing the endolymphatic duct) is almost 15 mm, which is three times the bone thickness at the anterior preserved extremity of the Nu. The exoskeletal division of the right endolymphatic duct opens on the visceral skull roof surface at the anterior edge of the area of thickened bone (th.end), and is also visible on the broken margin of the specimen (d.end, Fig.4A).This is also an advanced character of the brachythoracid skull — in large Emsian brachythoracids Proc. Linn. Soc. N.S.W., 125, 2004 G.C. YOUNG Figure 4. Doseyosteus talenti gen. et sp. nov. Holotype (ANU V1026). A,B. Larger portion of skull in internal (A) and external (B) views. C,D. Smaller skull portion in internal (C) and external (D) views. from Burrinjuck the endolymphatic duct is not within the bone, but anteriorly forms a bony tube attached to or projecting from the inner surface of a much thinner PNu plate (Young in press a: figs. 3, 4, 7A, 9B). A similar condition occurs in Holonema from Gogo (J.A. Long, pers. comm.; Miles 1971: fig. 53). The preserved part of the right Ce plate is crossed by a prominent sensory groove (csc), which Proc. Linn. Soc. N.S.W., 125, 2004 must be the central sensory canal rather than the supraorbital sensory canal, because of its oblique orientation to the midline. Middle and posterior pitlines are represented by faint markings in the region of the ossification centre (mp, pp). Anterolateral and posterolateral margins of the preserved part of the Ce plate are somewhat fractured, but appear to approximate natural margins. The former is bevelled 51 DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND N p 9 Sey SF Figure 5. Doseyosteus talenti gen. et sp. nov. Attempted skull roof reconstruction, preserved portions shaded. externally, and internally shows a contact face for the postorbital plate (cf.PtO), showing that it overlapped the PtO extensively, as in most other brachythoracids (e.g. Miles and Westoll 1968: fig. 2; Young 1979: fig. 1; 1981: fig. 5). Holonema is an exception in this respect (Miles 1971: fig. 12). Subdivisions of the posterior part of this contact face suggest that it also overlapped the marginal (M) plate (cf.M, Fig. 4A). The posterolateral margin of the Ce plate is somewhat thicker, and carries a deep groove (gr.M) for an interlocking suture, the Ce plate providing external and internal laminae to enclose the margin of the contiguous bone. The nature of the preserved margins suggests that they approximate the suture position. Since the anterior end of the PNu is well shown on the specimen, and is most unlikely to have extended to this margin of the Ce plate, it seems that the intervening space must have been occupied by a mesial projection of the M plate (M, Fig. 5). This arrangement has not previously been recorded in brachythoracids. A similar but smaller process of the M intrudes the Ce plate of Buchanosteus, but this is some distance in front of the PNu (Young 1979: fig. 1): . There is a long posterolateral projection of the Ce plate partly separating the Nu and PNu plates (plpr, Fig. 4B), a feature seen in various other brachythoracids. An early example with this Proc. Linn. Soc. N.S.W., 125, 2004 G.C. YOUNG morphology is Ulrichosteus Leliévre, 1982a from the Givetian of Germany, but this form has the Nu plate extending anteriorly in front of the PNu, whereas in Doseyosteus the PNu is slightly longer. Ardennosteus Leliévre, 1982b also has a strong posterolateral lobe of the Ce, but this Famennian form differs in its sinuous interlocking sutures, broader transverse nuchal thickening, and coarse tubercular ornament. Development of a posterolateral lobe of the Ce is one of three features representing the ‘trilobate’ condition of the Ce plates (characters 13, 14, 21 of Carr 1991), a widespread condition amongst Middle-Late Devonian eubrachythoracids which has proved difficult to define. Internally this part of the Ce is more extensive, the overlapped portion extending back to the endolymphatic thickening, again as in other brachythoracids. The visceral surface of the Ce is gently concave laterally, with several shallow grooves (vg) resembling the vascular grooves described in Holonema by Miles (1971: fig.12). This depressed region is flanked mesially by the pre-endolymphatic thickening (th.pre), which forms a low broad ridge with a curved anteromesial orientation. The preserved anteromesial edge of the Ce plate is thickened and abraded (Fig. 3D). Associated with this skull portion was a smaller part of the left preobstantic corner of the skull roof (Fig. 3B,C), assumed to have belonged to the same individual. The specimen includes part of the PNu and M plates (Fig. 4C,D), and is crossed by a section of the main lateral line (llc), and the infraorbital (ifc.ot) and postmarginal (pmc) sensory canals. Unlike forms such as Coccosteus, Holonema and Buchanosteus (Miles and Westoll 1968; Miles 1971; Young 1979), the M plate carries part of the subobstantic area (soa, Fig. 4D). A subobstantic area of similar extent is seen in the Gogo brachythoracid Harrytoombsia Miles and Dennis (1979: fig. 4), and in all plourdosteids sensu Long (1995). The PM plate is missing, but on the visceral surface there is a clear contact face for this bone (cf.PM, Fig. 4C). The visceral surface also shows the inframarginal crista to be strongly developed, dorsally as a very prominent irregular knob of bone (kb) separated posteriorly by a deep groove from the ventrally directed crista (cr.im), which itself carries a groove. The free ventral margin of the plate is thickened (lcp), representing the “lateral consolidated part’ of the skull, and a depression between the thickening and the inframarginal crista (dep) may correspond to similar structures in Coccosteus and Buchanosteus Young (1979: 314). The external ornament on both specimens comprises fine tubercles in some areas, sometimes only faintly discernible on a generally smooth surface (Fig. Proc. Linn. Soc. N.S.W., 125, 2004 3B,D). The fine ornament is similar to that on the SO plate of Atlantidosteus pacifica, but that form displays affinity with the homostiid arthrodires in a range of features (Young 2003a), whereas the skull of Doseyosteus talenti gen. et sp. nov. lacks various specialised characters of Homostius and related forms (e.g. elongate Nu and PNu plates, small dorsal orbits, etc.). The reduced ornament also distinguishes this new form from various ‘coccosteomorph’ arthrodire remains known from the early Middle Devonian of northern Germany and the Baltic sequence (Otto 1997, 1999). An attempted reconstruction of the skull roof of the new taxon based on available information is presented in Fig. 5. The skull could have been broader across the preobstantic corners than shown, since the gap between the two preserved portions is based only on a general alignment of sutures and sensory grooves. The anterior part of the skull is unknown, and restored shape of bones is generally based on various coccosteomorph arthrodires (e.g. Denison 1978: fig. 57). Advanced features depicted (T-shaped R plate, Pi plate separating PrO plates, trilobate Ce plates) are based on their co-occurrence with preserved skull characters in all other known taxa. They need to be confirmed with additional material. On the larger preserved portion, the breadth and anterior embayment of the Nu plate, and the marked posterior lobe of the Ce plate separating the Nu and PNu plates, are general resemblances to Eastmanosteus and Golshanichthys, as noted above. The M and Ce plates retain extensive contact to separate the PNu from the PtO, the assumed primitive condition for brachythoracids. In contrast, the plourdosteid arthrodires, which were widespread in the Late Devonian, and apparently replaced the largely Middle Devonian coccosteids (Long 1995), have a much enlarged PtO reaching back to contact both the Ce and PNu plates. In consequence the M plate is reduced in size, whereas in Doseyosteus gen. nov., although not completely preserved, the M plate was Clearly a more extensive bone, which apparently shows a unique feature in the large mesial process embaying the Ce plate in front of the PNu. In summary, this new but poorly known brachythoracid shows a range of advanced characters otherwise only seen in Middle or Late Devonian taxa, and it resembles the Frasnian taxa Eastmanosteus and Golshanichthys in several features which might indicate a close relationship. Eastmanosteus yunnanensis Wang, 1991 from the Givetian of China would otherwise be the earliest known member of this group (family Dinichthyidae). Kiangyousteus Liu, 1955, also from China (Givetian of Szechuan), may be another primitive dinichthyid (Denison 1978). Both 53 DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND taxa differ from the new form described here in their well-developed coarse tubercular ornament, presumably a primitive feature. Doseyosteus talenti gen. et sp. nov. displays an unusual shape of the M plate which is apparently unique to this new genus and species. More material, including the unknown trunk armour, which in brachythoracids comprises 17 separate bones, will clarify the affinities of this new taxon. ACKNOWLEDGMENTS Professor J.S. Jell (University of Queensland) and Professor K.S.W. Campbell (ANU) are thanked for making the specimen available for study. Mr R.W. Brown (Geoscience Australia) assisted in acid preparation. Professor J.A. Talent and Dr A. Basden (Macquarie University) advised and discussed at length the provenance and age of Broken River placoderms, and Dr S. Turner (Queensland Museum) provided comparative material. Comparison with European and Moroccan arthrodire material was facilitated by a visiting professorship at the Muséum national d’ Histoire naturelle, Paris, in 1999. Professor D. Goujet is thanked for arranging this, and for the provision of facilities, and together with Dr. H. Leliévre and Dr. P. Janvier discussed at length placoderm morphology and relationships. Dr Leliévre arranged for arthrodire casts to be sent to Canberra for comparative study. B. Harrold is thanked for providing essential computer support at ANU, and V. Elder for assistance with specimen curation. Dr E. Mark-Kurik and Dr R. Carr discussed arthrodire phylogeny, and Dr Carr arranged for a visit to Cleveland, Ohio, for study of large arthrodire material. Financial support was provided in Canberra by ANU Faculties Research Fund Grants F01083 and F02059, and overseas by the Alexander von Humboldt Foundation, for a Humboldt Award in Berlin (2000-2001), and assistance with travel to the USA (Flagstaff and Cleveland, 2000). I thank Prof. H.-P. Schultze for provision of facilities in the Museum fiir Naturkunde, Berlin. Dr P. De Deckker is thanked for provision of facilities in the Geology Dept., ANU. This research was a contribution to IGCP Projects 328, 406, 410, and 491. REFERENCES Basden, A. (2001). ‘Early Devonian fish faunas of eastern Australia: documentation and correlation’. Ph.D Thesis, Macquarie University [unpublished], 349 pp., 63 figs. Basden, A., Burrow, C.J., Hocking, M., Parkes, R. and Young G.C. (2000). Siluro—Devonian microvertebrates from south-eastern Australia. In IGCP 328, Final Report, A. Blieck and S. Turner (eds), Courier Forschungsinstitut Senckenberg 223, 201-222. Burrow, C.J. and Young, G.C. (in press). 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Mawson, R. (1987). Early Devonian conodont faunas from Buchan and Bindi, Victoria, Australia. Palaeontology 30, 251-297. Mawson, R. and Talent, J.A. (1989). Late Emsian- Givetian stratigraphy and conodont biofacies - carbonate slope and offshore shoal to sheltered Proc. Linn. Soc. N.S.W., 125, 2004 lagoon and nearshore carbonate ramp - Broken River, north Queensland, Australia. Courier Forschungsinstitut Senckenberg, 117, 205-259. McCoy, F. (1848). On some new fossil fishes of the Carboniferous period. Annals and Magazine of Natural History 2, 1-10. Miles, R. S. (1971). The Holonematidae (placoderm fishes), a review based on new specimens of Holonema from the Upper Devonian of Western Australia. Philosophical Transactions of the Royal Society of London. B. Biological Sciences 263, 101-234. Miles, R.S. and Dennis, K. (1979). A primitive eubrachythoracid arthrodire from Gogo, Western Australia. Zoological Journal of the Linnean Society 66, 31-62. Miles, R.S. and Westoll, T. S. (1968). The placoderm fish Coccosteus cuspidatus Miller ex Agassiz from the Middle Old Red Sandstone of Scotland. Part I. Descriptive morphology. Transactions of the Royal Society of Edinburgh 67, 373-476. Miller, H. (1841). “The Old Red Sandstone’. (Johnstone and Hunter, Edinburgh). Otto, M. (1997). Vertebrate fossils of the Middle Devonian (Eifelian) Mihlenberg Formation in the Bergisches Land, northwestern Germany. Paldontologische Zeitschrift 71, 107-116. Otto, M. (1998). New finds of vertebrates in the Middle Devonian Brandenberg Group (Sauerland, Northwest Germany). Paldontologische Zeitschrift 72, 117-134. Otto, M. (1999). Neues Material von Protitanichthys ? montanus (Vertebrata, Placodermi, Arthrodira) aus dem unteren Mitteldevon des Bergischen Landes. Neues Jahrbuch fur Geologie und Paldontologie Monatshefte 1999 (7), 397-408. Pedder, A.E.H., Jackson, J.H. and Philip, G.M. (1970). Lower Devonian biostratigraphy in the Wee Jasper region of New South Wales. Journal of _ Paleontology 44, 206-251. Sloan, T.R., Talent, J.A., Mawson, R., Simpson, A.J., Brock, G.A., Engelbretsen, M.J., Jell, J.S., Aung, A.K., Pfaffenritter, C., Trotter, J. and Withnall, I.W. (1995). Conodont data from Silurian-Middle Devonian carbonate fans, debris flows, allochthonous blocks and adjacent autochthonous platform margins: Broken River and Camel Creek areas, north Queensland, Australia. Courier Forschungsinstitut Senckenberg 182, 1-77. Turner, S., Basden, A. and Burrow, C.J. (2000). Devonian vertebrates of Queensland. In ‘IGCP 328, Final Report’ (Eds A. Blieck and S. Turner). Courier Forschungsinstitut Senckenberg 223, 487-521. Turner, S. and Cook, A. (1997). Ptyctodont jaw from the Broken River Province, NEQ. Memoirs of the Queensland Museum 42, 80. Wang J.-Q. (1991). A fossil Arthrodira from Panxi, Yunnan. Vertebrata PalAsiatica 29, 264-275 (Chinese, English summary). DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND White, E.I. (1952). Australian Arthrodires. Bulletin of the British Museum (Natural History) (Geology), 1, 249-304. White, E.I. (1978). The larger arthrodiran fishes from the area of the Burrinjuck Dam, N.S.W. Transactions of the Zoological Society of London 34, 149-262. White, E.I. and Toombs, H.A. (1972). The buchanosteid arthrodires of Australia. Bulletin of the British Museum (Natural History) (Geology) 22, 379-419. Woodward, A.S. (1891). “Catalogue of Fossil Fishes. Part 2’. (British Museum (Natural History), London). Young, G.C. (1969). ‘The geology of the Burrinjuck-Wee Jasper area, New South Wales’. B.Sc. Hons thesis, ANU Canberra [unpublished]. Young, G.C. (1974). Stratigraphic occurrence of some placoderm fishes in the Middle and Late Devonian. Newsletters on Stratigraphy 3, 243- 261. Young, G.C. (1979). New information on the structure and relationships of Buchanosteus (Placodermi, Euarthrodira) from the Early Devonian of New South Wales. Zoological Journal of the Linnean Society 66, 309-352. Young, G.C. (1981). New Early Devonian _ brachythoracids (placoderm fishes) from the Taemas - Wee Jasper region of New South Wales. Alcheringa 5, 247-271. Young, G.C. (1984a). An asterolepidoid antiarch (placoderm fish) from the Early Devonian of the Georgina Basin, central Australia. Alcheringa 8, 65-80. Young, G.C. (1984b). Comments on the phylogeny and biogeography of antiarchs (Devonian placoderm fishes), and the use of fossils in biogeography. Proceedings of the Linnean Society of New South Wales 107, 443-473. Young, G. C. (1986). The relationships of placoderm fishes. Zoological Journal of the Linnean Society 88, 1-57. Young, G.C. (1990). New antiarchs (Devonian placoderm fishes) from Queensland, with comments on placoderm phylogeny and biogeography. Memoirs of the Queensland Museum 28, 35-50. Young, G.C. (1993). Middle Palaeozoic macrovertebrate biostratigraphy of Eastern Gondwana. In ‘Palaeozoic Vertebrate Biostratigraphy and Biogeography’. (Ed. J.A. Long) pp. 208-251. (Belhaven Press, London). Young, G.C. (1995). Timescales 4. Devonian. Biostratigraphic charts and explanatory notes. 2™ Series. Australian Geological Survey Organisation, Record 1995/33, 1-47. Young, G.C. (1996). ‘Devonian (chart 4)’, In ‘An Australian Phanerozoic Timescale’ (Eds. Young, G.C. and Laurie, J.R.) pp. 96-109. (Oxford University Press, Melbourne). Young G.C. (2003a). A new species of Atlantidosteus Leliévre, 1984 (Placodermi, Arthrodira, Brachythoraci) from the Middle Devonian of the Broken River area (Queensland, Australia). Geodiversitas 25, 681-694. Young, G.C. (2003b). North Gondwanan mid-Palaeozoic connections with Euramerica and Asia; Devonian vertebrate evidence. Courier Forschungsinstitut Senckenberg 242, 169-185. Young, G.C. (in press a). Large brachythoracid arthrodires (placoderm fishes) from the Early Devonian of Wee Jasper, New South Wales, Australia. Journal of Vertebrate Paleontology Young, G.C. (in press b). Homostiid remains (placoderm fishes; Arthrodira), from the Early Devonian of the Burrinjuck area, New South Wales. Alcheringa Young, G.C. and Gorter, J.D. (1981). A new fish fauna of Middle Devonian age from the Taemas/Wee Jasper region of New South Wales. Bureau of Mineral Resources Geology & Geophysics, Bulletin 209, 83-147. Young, G.C. and Goujet, D. (2003). Devonian fish remains from the Dulcie Sandstone and Cravens Peak Beds, Georgina Basin, central Australia. Records of the Western Australian Museum, Supplement 65, 1-85. Young G.C., Leli¢vre H. and Goujet D. (2001). Primitive jaw structure in an articulated brachythoracid arthrodire (placoderm fish; Early Devonian) from southeastern Australia. Journal of Vertebrate Palaeontology 21, 670-678. Young, V.T. (1995). Micro-remains from Early and Middle Devonian acanthodian fishes from the U.K. and their biostratigraphic possibilities. Ichthyolith Issues, Special Publication 1, 65-68. Zhu, M. (1996). The phylogeny of the Antiarcha (Placodermi, Pisces), with the description of Early Devonian antiarchs from Qujing, Yunnan, China. Bulletin du Museum national d Histoire naturelle, Paris (4,C) 18, 233-347. Proc. Linn. Soc. N.S.W., 125, 2004 Effects of Slashing and Burning on Thesium australe R. Brown (Santalaceae) in Coastal Grasslands of NSW JANET S. COHN Biodiversity Research and Management Division, NSW National Parks and Wildlife Service, PO Box 1967, Hurstville, NSW 2220 (Ganet.cohn@npws.nsw.gov.au) Cohn, J. (2004). Effects of slashing and burning on Thesium australe R. Brown (Santalaceae) in coastal grasslands of NSW. Proceedings of the Linnean Society of New South Wales 125, 57-65. Two studies examined the effects of burning and cutting on aspects of the population dynamics of a nationally vulnerable herb, Thesium australe on the central and north coast of NSW. Study sites were grasslands dominated by Themeda australis with scattered native shrubs (Banksia integrifolia, Acacia sophorae) and the exotic shrub Chrysanthemoides monilifera ssp. rotundata. In the first study (May 1995 to December 1996), Thesium australe occurred at high density (1/m?) on exposed, long-unburnt headlands. In the second study, (December 1996 to December 1998), Thesium australe was at low density (<1/100m7) on more protected and recently burnt hinterland. On the headlands, winter treatments had no significant effect on the survival, density and vigour of Thesium australe. In the hinterland, one year after summer treatments, seedling recruitment resulted in a higher density of Thesium australe in the cut plots than either the burnt or the control. Flowenng and fruiting of Thesium australe were not restricted by season. After winter and summer treatments, flowering and fruiting occurred within 6 months and 1 year, respectively. Although exposed coastal headlands may require no management intervention to increase the occurrence of Thesium australe, except where the possibility of shrub invasion exists, a regime of slashing on less exposed hinterlands may be needed to reduce competition from Themeda australis. Further research is necessary to determine if slashing or burning the more protected hinterland would yield different results if carried out in seasons other than summer. Manuscript received 1 March 2003, accepted for publication 22 October 2003. KEYWORDS: fire, grasslands, headlands, mowing, slashing, Thesium australe. INTRODUCTION Although Thesium australe is a herb with a wide ecological tolerance, extending from tropical to alpine climates, it is confined to widely scattered locations in open woodlands and grasslands where Themeda australis/ T. triandra (Kangaroo Grass) is common in the understorey (Scarlett et al. 1994). On the north coast of NSW, T. australe occurs on grassy headlands used predominantly for passive recreation, often adjacent to residential areas (Griffith 1992; Fig. 1). In south-eastern Australia, open woodland and grassland communities have largely been modified and fragmented by introduced grazers, cultivation and changed fire regimes (Stuwe and Parsons 1977; Scarlett and Parsons 1990; McDougall and Kirkpatrick 1994; Tremont and McIntyre 1994; Prober and Thiele 1995; Lunt 1997). As a consequence T. australe is rated as nationally vulnerable (Briggs and Leigh 1996) and vulnerable in NSW under Schedule 2 of the NSW Threatened Species Conservation Act 1995 Non-coastal, long-unburnt grasslands dominated by Themeda australis / triandra, have been shown to be species poor (Stuwe and Parsons 1977; Kirkpatrick 1986; McDougall 1989), largely as a result of the high competitive ability of this tussock grass (Groves 1974). With a general recent decline in fire frequency on coastal headlands (Griffith 1992), dominance by Themeda australis and the recruitment of native and exotic shrubs are potential threats to the survival of T. australe (Griffith 1992), although Cooper (1986) suggested that headlands exposed to salt-laden winds may be an exception. He cites the persistence of T. australe at Perpendicular Point, 20 years after fire, as an example. Research on Thesium alpinum in Denmark found that it became extinct as a result of shading from trees (Lojtnant and Worsoe 1980). Thesium australe may be similarly sensitive. In coastal Victoria, an increase in native shrub and tree recruitment has been linked to a decline in fire frequency (Bennett 1994; McMahon et al. 1994; Lunt 1998a b). On the north coast of NSW, increased recruitment of native shrubs EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE Figure 1. Grassland habitat of Thesium australe at Look at Me Now Headland on the north coast of NSW. and trees (Acacia, Banksia, Allocasuarina spp.) and an invasive exotic shrub, Bitou bush, (Chrysanthemoides monolifera ssp. rotundata have been observed (Dodkin and Gilmore 1985; Griffith 1987; Griffith 1992). A number of studies have suggested a regime of regular burning and/or mowing to maintain species richness in grasslands and prevent shrub invasion (Groves 1974; Stuwe and Parsons 1977; Kirkpatrick 1986; McDougall 1989; Lunt 1990a, 1998b). Current information on the response of T. australe to fire in the field has been based on observations. While Leigh and Briggs (1989) suggest that survival and recruitment are unaffected by fire, Archer (1984) believed seeds were stimulated to germinate. In laboratory trials, Scarlett (pers. comm.) found that heat did not stimulate seed germination. There have been no studies on the effect of mowing or cutting on T. australe (Griffith 1992). On coastal headlands and conservation reserves where burning or slashing grasslands may be used for conservation or hazard reduction purposes, it is important to establish their effect on native species. Two separate studies examined the effects of a single burning and a single cutting on aspects of the population dynamics of T. australe, namely: 1/ its survival, density, vigour and reproductive status where it occurred at relatively high density on long-unburnt, exposed headlands (winter treatments); 2/ its density and reproductive status where it was at very low density in a more protected and recently burnt hinterland (summer treatments). These were not intended to be comparative studies and indeed the different timing and methods of treatment (see Materials and Methods), driven by 58 the availability of resources, make this not possible anyway. MATERIALS AND METHODS The studies were located at several sites on the north and central coast of NSW (Fig. 2): Perpendicular Point (AMGR Easting 485600, Northing 6499200); Look at Me Now Headland (E 518000, N 6661300); and Old Bar Park (E 461300, N 6462800). Perpendicular Point and Look at Me Now Headland are within respectively, Kattang Nature Reserve (NR) and Moonee Beach NR. Both are managed by the New South Wales National Parks and Wildlife Service (NSW NPWS). Old Bar Park is managed by The Greater Taree City Council. All three sites are used for recreation, predominantly by walkers. Although no motor vehicular access is allowed in the NRs, there was evidence of their past usage at Look at Me Now Headland, where at the time of this study wheel ruts were still very obvious. Vehicles were used on Perpendicular Point as recently as 1986 (Cooper 1986). There is some use of motor vehicles in Old Bar Park, but this is mostly on the pre-existing tracks and the airstrip (author’s personal observations). Perpendicular Point and Look at Me Now Headland are characterised by black headland soils, which are loamy soils high in organic matter (Parbery 1947). Yellow podzolic soils predominate at Old Bar Park (Long 1996). Aspects and slopes of the study sites varied. At Perpendicular Point the site was located on a north-western aspect with a slope of 9°, whilst at Look at Me Now Headland the site was on a more exposed southerly aspect with a slope of 6°. The site at Old Bar Park was flat. The study sites were in grassland communities dominated by Themeda australis. Scattered shrubs at Perpendicular Point included native (e.g. Acacia sophorae, Banksia integrifolia) and exotic (e.g. Chrysanthemoides monilifera ssp. rotundata) taxa. Another nationally endangered herb, Zieria prostrata (Briggs and Leigh 1996) also occurred on a number of the headlands with T. australe (Griffith 1992; NPWS 1998). Thesium australe was found at Perpendicular Point in 1957 (Cooper 1986) and at Look at Me Now Headland and Old Bar Park after 1992 (Griffith 1992). Although at relatively high density at Perpendicular Point and Look at Me Now Headland (approximately 1/m*), at Old Bar Park it occurred mostly as very scattered plants (approximately <1/100 m7). Thus, the focus at this latter site was more on recruitment Proc. Linn. Soc. N.S.W., 125, 2004 J.S.COHN Moonee Beach Nature Reserve P Coffs Harbour kilometres Port Macquarie Reserve Map location ¢ Figure 2. Locality of study sites within Moonee Beach NR (Look at Me Now Headland), Kattang NR (Perpendicular Point) and Old Bar Park on the north and central coast of NSW. Stippled areas represent estate managed by NSW National Parks and Wildlife Service. responses to treatments. There was little information on the fire history at the three sites. Griffith (1992) believed that Perpendicular Point may not have burnt for a considerable period of time. In 1985, Cooper (1986) believed that Perpendicular Point had not burnt for at least 20 years. There was no record of the last fire at Look at Me Now Headland. Old Bar Park was last burnt in 1991 by a low intensity fire (T. Cross pers. comm.). Approximately 1 year prior to this study Old Bar Park was slashed (S. Griffith pers. comm.), presumably for hazard reduction purposes. Headlands (high density plants) Treatments were applied in winter (July 1995) at Perpendicular Point and Look at Me Now Headland (Table 1). There were 15 replicate plots of each treatment (burnt, cut, control). Each treatment was allocated randomly to a 0.5 m x 0.5 m plot laid out in rows, over a total area of 75 m? at Perpendicular Point and 112 m? at Look at Me Now Headland. Plots were Proc. Linn. Soc. N.S.W., 125, 2004 burnt using a gas burner. Because of the heavy dew, each burnt plot was subjected to heat for 5 minutes, until all of the grasses and herbs had been burnt and the bare ground had been heated and scorched. This simulated a high intensity burn (R. Bradstock pers. comm.). In the cutting treatment all grasses and herbs, including T. australe were cut to within 0.5 cm of the ground with shears. At both sites, in all plots, individual T. australe plants were tagged and numbered and the fates of the original and emerged plants were surveyed over 1.5 years (Table 1). Data on plant vigour (number of stems/ plant; Perpendicular Point only) and the incidence of flowering or fruiting were also collected. Analyses of the proportion of T. australe plants surviving 6 and 16 months after treatment, were made using Generalised Linear Modelling (GLIM), with a binomial error structure (Crawley 1993). The effects of the factors, treatment (burnt, cut, control) and site (Perpendicular Point, Look at Me Now Headland) and their interactions were examined using the chi-squared 59 EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE Table 1. The dates of treatment applications and monitoring at the study sites. Study Site Treatment (date) burn, cut (26/7/95) burn, cut (27/7/95) Perpendicular Point Look at Me Now Headland Old Bar Reserve statistic. The density of T. australe plants (0.25 m*) was examined using fully factorial analyses of variance (ANOVA) and Tukey tests for pairwise comparisons. The effects of treatments (burnt, cut, control) and sites (Perpendicular Point, Look at Me Now Headland) were examined at pre- and post-treatment dates (0, 6 and 16 months). To satisfy Cochran’s test of homogeneity of variances, data were square root transformed and if necessary a more conservative level of significance (p<0.01) was applied (Underwood 1981). Analyses of the vigour of T. australe plants (number of stems/plant) at Perpendicular Point were made using one-way ANOVAs. The effects of treatments (burnt, cut, control) were examined at pre- and post-treatment dates (0, 6 and 16 months). Data from all plots and cohorts within each treatment were pooled. Hinterland (low density plants) At Old Bar Park, where T. australe occurred at very low density, large plots were subjected to burning or slashing. Each treatment (burnt, slashed, control) was allocated to a 10 m x 10 m plot within an overall area of 40 m x 50 m. There were 2 replicates of each treatment. Whilst for practical purposes the two burnt plots were placed together, replicates of the cut treatment and control were randomly allocated to the remaining plots. Burning took place in hot conditions during summer (December 1996). Two plots were slashed the next day to within 5 cm of the ground. The resulting cuttings were removed from the plots. Individual T. australe plants were tagged, numbered and followed for 2 years (Table 1). Although not measured quantitatively at Perpendicular Point and Look at Me Now Headland, observations indicated that the measurement of bare ground may be useful in discussing trends in the data. The cover of grasses/herbs and bare ground were measured at each census (<5 replicates) in classes (1=1- 10%, 2=11-20%...... ,10=91-100%) within randomly allocated quadrats (1 m*), located within each treatment 60 burn, slash (16/12/96) Monitoring Dates (pre and post treatment) 12/5/95, 14/2/96, 4/12/96 26/7/95, 11/2/96,18/12/96 3/12/96, 2/12/97, 16/12/98 plot. Rock cover was negligible. Analysis of the density of T. australe plants in each plot (number/100 m?) was made using fully factorial ANOVAs and Tukey tests for post hoc comparisons. The effects of the treatments (burnt, cut, control) were examined on each day of sampling. Analyses of the cover classes of bare ground were made using a two-way fully factorial ANOVA. The effects of treatment (burnt, cut, control) and sampling date (pre-treatment, 1 and 2 years post- treatment) were examined. RESULTS Headlands (high density plants) Site, but not treatment, had a significant effect on the proportion of plants surviving 6 and 16 months after the start of the study (Fig. 3). At both times survival was higher at Perpendicular Point than at Look at Me Now Headland (respectively X7=10, df=1, p<0.005; X?=8.5 df=1, p<0.025). By the end of the study between 80% and 100% of the original plants had suffered mortality. Six months after the application of treatments there was no significant difference in the density of T. australe (0.25 m) with respect to treatment and site (p>0.05; Fig. 4). Sixteen months after treatment, however, there was a significant effect of site (F=4.72, df = 1,84, p<0.05). Look at Me Now Headland had a higher density of plants than Perpendicular Point. At Perpendicular Point there was no significant difference in the vigour of T. australe (number of stems/plant) with respect to treatment either prior to treatment, or 6 months and 16 months after treatment (p>0.05; Fig. 5). There appeared to be a general increase in plant vigour over this period. Within 6 months of applying treatments at Perpendicular Point and Look at Me Now Headland, flowering and fruiting of original plants and new recruits of 7. australe were recorded in summer (11 February 1996). Proc. Linn. Soc. N.S.W., 125, 2004 J.S.COHN Hinterland (low density plants) The density of T. australe plants at Old Bar Park was_ significantly affected by treatment 2 years after application. The slashed areas had a higher density than either the burnt or the control which were not significantly different from one another (F=15.5, df=2,3, p<0.05; Fig. 6). Pre-treatment year) and | year after treatment, there was no significant difference in the density of T. Treatments : . applied Jul-95 + Sep-95 | Proportion of plants alive Jan-96 | May-95 Nov-95 Mar-96 May-96 Jul-96 Sep-96 Nov-96 =n a. Date (month an Figure 3. Proportional survival of T. australe (mean, se) at Perpendicular Point (#) and Look at Me Now Headland (™) following treatments (pooled; #“S‘ra/e between treatments burnt, cut, control) applied in July 1995. (p>0.05). There was a significant interactive effect of treatment and time of a/ Perpendicular Point sampling on the cover of bare ground at Old Bar Park (F=3.12, df=4, 27, p<0.05; Fig. 7). Whilst there was no significant difference between the plots prior to the imposition of treatments, 1 and 2 years (no significant Treatments applied mo FP NY WwW N Qn On ava NG NOP NO PND | KO ONS & ‘Ley lg og ae ge em difference) after burning, the N Zw Hae ahGiaes. oe oe iS So b d ignificantl = 3 o oO are ground was significantly S 3 SS = Da = het Dai Zot x = LPC higher than in the slashed or — =| the control at any time (except eo b/ Look at Me Now Headland | bum at 2 years = pre-burn and Siwas cut at 2 years). A Within 1 year of 2 treatment application, 1 flowering and fruiting of new recruits of 7. australe were 0 recorded in summer (2 Dec 1998). S$ GR Se RF REG S Piteems 8 2 S 2 fF Ss 2 A 2 DISCUSSION Date (month and year) Headlands (high density plants) Figure 4. The density (mean, se) of 7. australe plants (0.25 m7) before Burning or cutting T. and after treatments at Perpendicular Point and Look at Me Now australe plants in winter, did Headland. Treatments (burnt @, cut @, control A) were applied in July —-‘0t significantly affect their 1995. survival. Similarly, Leigh and Briggs (1989) found the survival of a population of T. australe near Canberra, was unaffected by a trial burn in autumn. Indeed, the Proc. Linn. Soc. N.S.W., 125, 2004 61 EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE = 4 Treatments applied ei LKB ow | AY 2, 2 one0 Z May-96 Jul-96 Sep-96 Nov-96 OD YD WY NH OO OO Cogt Site meat SR Sine Mie oe = PI Zae e Date (month and year) Figure 5. The size of T. australe plants before and after treatments at Perpendicular Point. Size was measured as the number of stems per plant (mean, se). Treatments (burnt @, cut @, control A) were applied in July 1995. — Nn Treatments applied — i=) No. plants/1 00m” N >) 1998 1997 Date (December in year) 1996 Figure 6. The density (100 m7) of 7. australe plants (mean, s.e.) in the hinterland at Old Bar Park before and after treatments were applied (burnt @, slashed ™, control A). Treatments were applied in December 1996. se) 5 a Treatments 8 BS 6 OD oO i, Sinisb.c Seen) 1997 Date (December in year) Figure 7. Cover classes (mean, se) of bare ground before and after treatments (burnt , slashed ™, control A), in the hinterland at Old Bar Park. Treatments were applied in December 1996. Cover classes:1=0-10%.....10=11-100%. existence of buds in the immediate vicinity of the soil surface (McIntyre et al. 1995) allows the species to resprout after disturbance. In subalpine and tableland climates, it is the habit of T. australe to die back to the rootstock during winter and resprout in spring (Cooper 1986; Archer 1987; Gross et al. 1995; Cohn 1999). This is not the case in coastal areas, where the species persists all year round (Cohn 1999). Whilst a study on the southern tablelands of NSW (Leigh and Briggs 1989), describes T. australe as an annual or a biennial, this study suggests that the species may live longer on the coast. After 6 and 16 months, respectively, approximately 30% and 17% of plants were still alive. Since it is likely that these plants originated at least 9 months previously in spring, their ages were more than likely 15 months and 25 months, respectively. Certainly, Prober and Thiele (1998) believe it possible that T. australe lives longer in less severe climates. Although there was no significant effect of treatment on the density of 7. australe, there was a higher density at the more exposed Look at Me Now Headland than at Perpendicular Point 2 years after treatments. This agrees with Cooper’s (1986) hypothesis that competition from Kangaroo Grass (T. australis) on exposed headlands is reduced by salt laden winds. It is also possible that the experimental burn, which was hotter than would be experienced naturally, even in extreme conditions (R. Bradstock pers. comm.), could have led to some mortality of T. australe seeds near the soil surface, thus reducing the effectiveness of this treatment. The small size of the plots may also have reduced 62 Proc. Linn. Soc. N.S.W., 125, 2004 J.S.COHN treatment effectiveness. Finally, more time may have been required for the T. australe populations to respond to a reduction in competition brought about by the experimental treatments. Thesium australe is able to grow and reproduce very quickly following disturbance in winter. In December, 6 months after burning or cutting, there was no significant difference in the vigour of plants in the treated plots and the control. At the same time resprouting plants and new recruits were flowering and beginning to fruit. Indeed, flowering and fruiting of T. australe at both Perpendicular Point and Look at Me Now Headland occurred throughout the year (Cohn 1999). By contrast, flowering and fruiting has been found to be seasonal at inland locations, occurring from spring to autumn (Stanley and Ross 1983; Briggs and Leigh 1985; Gross et al. 1995; Cohn 1999). Hinterland (low density plants) At the more protected Old Bar Park, where T. australe was mostly absent from the plots prior to treatment, summer slashing rather than burning led to significant seedling recruitment of T. australe, 2 years after treatment (Fig. 6). Although it is generally recognised that burning provides the bare ground for seedling establishment that slashing does not (Lunt 1990a), other factors seemed to be at play in this study. The comparable cover of bare ground in all treatments at the time of the high numbers of T. australe in the slashed plots (Fig. 7), indicates that a reduction in grass height may have been responsible. In Victoria, Lunt (1990b) believed that selective grazing of tussocks to a height of 5 cm by rabbits and kangaroos may have contributed to the maintenance of species richness by reducing competition from perennial grasses. Thus, it is probable that a reduction in the height of the dominant species Themeda australis rather than an increase in bare ground, led to significant recruitment of 7. australe seedlings in the slashed treatment. Given that the post-fire conditions reduced competition from Themeda australis, it is curious that there was not a significant effect on T. australe numbers. In the same summer following burning, seedlings of T. australe were observed in these plots (S. Long pers. comm.). Their low numbers throughout the study, however, may have resulted from the more exposed conditions, reflected by higher cover of bare ground, experienced during the first and second summers (Fig. 7). In addition, T. australe’s hemiparasitic dependence on other herbs and grasses, (Scarlett et al. 1994), may have made it difficult for T. australe to survive, given that its hosts were also recovering from the effects of the fire. Indeed summer Proc. Linn. Soc. N.S.W., 125, 2004 ‘dying back’ of T. australe in times of water stress has previously been recorded by Leigh and Briggs (1989). Whilst older T. australe plants may have the resources to recover, seedlings, such as those observed in this experiment soon after the burning, may not have had that capability. Management implications The results from this study, coupled with the long-term persistence of 7. australe on exposed headlands in the absence of active management (Cooper 1986; Griffith 1992), indicate that there is apparently no need for a change in this regime, except where shrub recruitment (native or exotic), may be competing with the survival of T. australe. By contrast, in the more protected hinterlands, where T. australe also occurs, active management may be required to reduce competition from Kangaroo Grass (T. australis). Although, results from this study indicate that early summer slashing of a grassland (5 cm height) resulted in recruitment of T. australe plants, further research is required to determine if this is the most appropriate time of the year and method. Of particular concern is that disturbance of the summer growing Kangaroo Grass (T. australis) at this crucial time could result in the introduction of weed species (Gniffith 1992). There is also a need to determine if burning outside summer, especially in autumn or early spring, would yield different results. This is important in the light of other work, which indicates that fire rather than mowing in grasslands is preferable to maintain species richness (Kirkpatrick 1986; Lunt 1991; James 1994). If shrub encroachment becomes a threat to the survival of T. australe, studies have recommended various fire intervals of between 2 and 10 years to reduce dominance of native shrubs (Groves 1974; Lunt 1998b) or a regime of frequent fire and mechanical disturbance to reduce exotic shrub frequency (e.g. Chrysanthemoides monolifera; Kirkpatrick 1986). Although this study did not examine an appropriate disturbance interval for T. australe, its quick growth and reproductive development and its continued presence at Hat Head (S. Griffith pers. comm.), which has burnt every 2 to 4 years for the past 15 years (NSW NPWS Records), indicates it can apparently cope with a relatively frequent disturbance regime. Studies in Victoria (Scarlett and Parsons 1982, 1993) suggest that the absence of T. australe and other late-flowering species along railway lines has resulted from annual, late-season burning. Further research is required in coastal areas to determine an appropriate disturbance interval for the long-term conservation of T. australe. 63 EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE ACKNOWLEDGEMENTS Thanks to Steve Griffith, Shirley Cohn and Steve Clemesha for their assistance in the field. National Parks and Wildlife Service staff from Port Macquarie District office provided me with encouragement, information and equipment. Thanks to Old Bar Bushfire Brigade, who carried out the burn at Old Bar Park. Financial assistance was provided by Environment Australia and New South Wales National Parks and Wildlife Service. Thanks to Andrew Denham and Mark Tozer who kindly provided comments on the manuscript. REFERENCES Archer, W.R. (1984). Thesium australe R. Brown (Santalaceae)-field notes and observations. The Victorian Naturalist 101, 81-85. Archer, W.R. (1987). Additional field notes and observations of Thesium australe R. Brown (Santalaceae). 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Species and ecosystems at risk: a preliminary review. In “Bitou Bush and Boneseed’ (Eds A. Love and R. Dyason) pp.33-52. NSW National Parks and Wildlife Service and NSW Department of Agriculture. Griffith, S. J. (1987). A Survey of the Distribution of Bitou Bush (Chrysanthemoides monilifera ssp. rotundata) on the North Coast of NSW between Diamond Head and Seal Rocks, with Recommendations for Management. Unpublished Report, New South Wales Interdepartmental Working Group on Bitou Bush. Griffith, S. J. (1992). Species Recovery Plan for Thesium australe. Unpublished Report, Endangered 64 Species Program, Australian National Parks and Wildlife Service, Canberra. Gross, C.L., Nano, C., Jones, R., and Harre, C. (1995). Thesium australe (Santalaceae). An Examination of Population Sizes at Selected Sites on the New England Tablelands, NSW-Summer 1994/95. Unpublished Report, Environment Australia, Canberra. Groves, R.H. (1974). Growth of Themeda australis grassland in response to firing and mowing. CSIRO, Australia, Division of Plant Industry Field Station Record 13, 1-7. James, T.A. (1994). Observations on the effects of mowing on native species in remnant bushland, western Sydney. Cunninghamia 3, 515-519. Kirkpatrick, J.B. (1986). The viability of bush in cities-ten years of change in urban grassy woodland. Australian Journal of Botany 34, 691-708. Leigh, J.H. and Briggs, J.D. (1989). Research relating to the conservation of rare or threatened plant species and their habitats in eastern Australia. In “The Conservation of Threatened Species and their Habitats’. (Eds M. Hicks and P. Eiser) pp.192- 201. (Occasional Paper No. 2: Canberra). Lojnant, B. and Worsoe, E. (1980). The genus Thesium Toadflax extinct in Denmark. Flora Fauna 86, 65-71. Long, S.A. (1996). Old Bar Park Plan of Management (Draft). (Greater Taree City Council, Taree). Lunt, I.D. (1990a). The impact of an autumn fire on a long-grazed Themeda triandra (Kangaroo Grass) grassland, implications for management of invaded, remnant vegetation. The Victorian Naturalist 107, 45-51. Lunt, I.D. (1990b). Species-area curves and growth-form spectra for some herb-rich woodlands in western Victoria, Australia. Australian Journal of Ecology 15, 155-161. Lunt, I.D. (1991). Management of remnant lowland grasslands and grassy woodlands for nature conservation: a review. The Victorian Naturalist 108, 56-66. Lunt, I.D. (1997). Effects of long-term vegetation management on remnant grassy forests and anthropogenic native grasslands in south-eastern Australia. Biological Conservation 81, 287-297. Lunt, I.D. (1998a). Two hundred years of land use and vegetation change in a remnant coastal woodland in southern Australia. Australian Journal of Botany 46, 629-647. Lunt, I.D. (1998b). Allocasuarina (Casuarinaceae) invasion of an unburnt coastal woodland at Ocean Grove, Victoria: structural changes 1971-1996. Australian Journal of Botany 46, 649-656. McDougall, K.L. (1989). The Re-establishment of Themeda triandra (Kangaroo Grass): Implications for the Restoration of Grassland. Technical Report Series No. 89, Arthur Rylah Institute for Environmental Research, Melbourne. McDougall, K.L., and Kirpatrick, J.B. (1994). (Eds) ‘Conservation of Lowland Native Grasslands in Proc. Linn. Soc. N.S.W., 125, 2004 J.S.COHN South-eastern Australia’. (World Wide Fund for Nature: Sydney). McIntyre, S., Lavorel, S., and Tremont, R.M. (1995). Plant life-history: their relationship to disturbance in herbaceous vegetation. Journal of Ecology 83, 31-44. McMahon, A.R.G., Carr, G.W., Bedggood, S.E., Hill, R.J., and Pritchard, A.M. (1994). Prescribed fire and control of coast wattle (Acacia sophorae (Labill.) R. Br.) invasion in coastal heath south- west Victoria. In ‘Fire and Biodiversity: The Effects and Effectiveness of Fire Management’. pp. 87-96. Biodiversity Series Paper No. 8, Department of Environment, Sport and Territories, Melbourne. NPWS (1998). ‘Zieria prostrata Recovery Plan’. (NPWS: Sydney). Parbery, N.H. (1947) Black headland soils of the south coast. Unusual process of formation. Agricultural Gazette of New South Wales 58, 123-125. Prober, S.M. and Thiele, K.R. (1995). Conservation of the grassy white box woodlands: relative contributions of size and disturbance to floristic composition and diversity of remnants. Australian Journal of Botany 43, 349-366. Prober, S.M. and Thiele, K-R. (1998). Ecology and Management of the Austral Toad-flax (Thesium australe) at ‘Open Grounds’, Gillingal Station, East Gippsland. Unpublished Report, Department of Natural Resources and Environment, Victoria. Scarlett, N.H., Bramwell M. and Earl, G. (1994). Austral Toad-flax, Thesium australe. Action Statement No. 56. Unpublished Report, Department of Conservation and Natural Resources, Victoria. Proc. Linn. Soc. N.S.W., 125, 2004 Scarlett, N.H. and Parsons, R.F. (1982). Rare plants of the Victorian Plains. In “Species at Risk: Research in Australia’. (Eds R.H.Groves and W.D.L Ride) pp. 89-105. (Australian Academy of Science: Canberra). Scarlett, N.H. and Parsons, R.F. (1990). Conservation biology of the southern Australian daisy Rutidosis leptorrhynchoides. In “Management and Conservation of Small Populations’. (Eds T.W. Clark and J.H. Seebeck) pp. 195-205. (Chicago Zoological Society: Illinois). Scarlett, N.H. and Parsons, R.F. (1993). Rare or threatened plants in Victoria. In ‘Flora of Victoria’. Vol. 1. (Eds D.B. Foreman and N.G. Walsh) pp. 227-254. (Inkata Press: Melbourne). Stanley, T.D. and Ross, E.M. (1983). “Flora of South- Eastern Queensland’. Volume 1. (Queensland Department of Primary Industries: Brisbane). Stuwe, J. (1986). An Assessment of the Conservation Status of Native Grasslands on the Western Plains, Victoria and Sites of Botanical Significance. Arthur Rylah Institute Report Series No. 48, Department of Conservation, Forests and Lands, Melbourne. Stuwe, J. and Parsons, R.F. (1977). Themeda australis grasslands on the basalt plains, Victoria: floristics and management effects. Australian Journal of Ecology 2, 467-476. Tremont, R.M. and McIntyre, S. (1994). Natural grassy vegetation and native forbs in temperate Australia: structure, dynamics and life histories. Australian Journal of Botany 42, 641-658. Underwood, A.J. (1981). Techniques of analysis of variance in experimental marine biology and ecology. Annual Reviews of Oceanography and Marine Biology 19, 513-605. 65 wnt i E v ay “Sy ; | ee ih aie ; “syat b ip & ni tomenst: ‘aa ig agpege. al ante nena. : “i ns 5 a (Soe LLM see HB she) eibeptasils ies: ee > Clkenniiga, Serre We nao estan RL Packs, atic Wee Set reper ah Port Me ATM wFicd ry real hhh ocr eee a a ie cL ee Yk Ta ceumeekdas°Oe tay ceacted Ah Hi es : 3} 5 ae ee) atari ofan cenit, : oi ist ppp MT 4282) AR sateen bas, ~ aaa x, ee Jogwl ak siearkY a snp biecreltanaty | OP a ae ge fod < Oe ¢ ie Pan “4 awiey Ma: : 273 >y. i Testbed wi ¥: a ada Bodtagises’ Pied ae sRecees, <* "+ fel Aira BN ee tee TS hy [ Pde, (MM 3; soviet! cS i Twat tu bowen pa A ated anit Fe , ae rast Ww rg ot 43, AS Nagi, Phat an 5 fee meas | ia awe athe tr Sqvite oi: we aed Useanpcatal of “ a a ~ Agro! rine = a ie te gee . EEOC oar, (ales xe ‘Be ere ie os ee + Bisrss Sus i “hey yrds shy owen , eT ofr spa Uh Nien X Ula tread arc Smal chal ines cers pas See ‘ H skaisoe ¢ i, state tin Maneyeamens - sy OTs - eo he, pasa i } A ta aay WW of ia ee ie a oom ‘Chentivis ‘Stes c Pe Siar Pegs irery taser fag Sheds . r : Wee _ Te PY we ” a= os : . pes ; Sea, 7ert, Faas oh A (ce im . Lic ‘ han 4 Ai " “4 ‘ . , 66 | . w £ : ; j my Phe A N ; = J i na ad Trichromothrips veversae sp.n. (Insecta, Thysanoptera), and the Botanical Significance of Insects Host-specific to Austral Bracken Fern (Pteridium esculentum) LAuRENCE Mounp! AND Masami MAsumoro2 ‘CSIRO Entomology, GPO Box 1700, Canberra 2601 (laurence.mound @csiro.au); *MAFF, Yokohama Plant Protection Station, Shin’yamashita, 1-16-10, Yokohama, 238-0801, Japan. Mound, L. and Masumoto, M. (2004). Trichromothrips veversae sp.n. (Insecta, Thysanoptera), and the botanical significance of insects host-specific to Austral bracken fern (Pteridium esculentum). Proceedings of the Linnean Society of New South Wales, 125, 67-71. Austral bracken fern, Pteridium esculentum, differs from its European counterpart in supporting one species of both thrips and aphid. The previously undescribed species of thrips, Trichromothrips veversae sp-n. (Thripidae), is widespread and locally abundant in southern Australia breeding on the youngest fronds of bracken but not on other ferns. It is unique among nearly 30 species of this Old World tropical genus in lacking long setae on the pronotum. Manuscript received 18 June 2003, accepted for publication 17 September 2003. KEYWORDS: aphids, bracken, Preridium, thrips, Trichromothrips, INTRODUCTION Common bracken fern is often considered to be a single, cosmopolitan species Pteridium aquilinum (Dennstaedtiaceae). In retaining this view, the major reference work on botanical nomenclature (Mabberley, 1997) recognised two subspecies, the nominate one from the Northern Hemisphere and Africa, and P. aquilinum caudatum from the Southern Hemisphere. In Australia, in contrast, Brownsey (1989) recognised three species of Pteridium: P. aquilinum introduced to a small area of South Australia in the Adelaide Hills; P. revolutum native to north-eastern Queensland but extending widely across New Guinea and South East Asia; and P. esculentum native to southern and eastern Australia but extending to South East Asia and the Pacific. More recently, Thomson (2000) has concluded from an extensive study of both structural and molecular characters that several of the Pteridium varieties distinguished worldwide, including esculentum, “might best be treated as species’. These differences in opinion concerning the botanical status of bracken fern are not without entomological significance. No species either of aphid (Homoptera) or of thrips (Thysanoptera) is known to live on bracken in Europe, where this plant is widespread and abundant and often an invasive weed. In contrast, the aphid species Shinjia orientalis (Mordwilko) (= S. pteridifoliae Shinji) has been reported widely on Preridium from northern India and Japan to eastern Australia. Moreover, populations of bracken in eastern North America support another aphid species, Mastopoda pteridis Oestlund, and in western North America five aphid species in the genus Macrosiphum have been reported from Pteridium (V. F. Eastop, 2003 pers. comm.). If Pteridium were truly monotypic, comprising one worldwide panmictic species, then different populations might be expected to support similar, if not identical insect species. The description here of a new species of Thripidae that is widespread on bracken in Australia would thus appear to provide further support for the recognition of distinct species within this ubiquitous plant genus. Presumably these insects are reflecting diversity within the genus Pteridium that botanists have been reluctant to acknowledge. The existence of this thrips species had been suspected for many years. In 1967, the wife of the eminent Australian insect ecologist H.G. Andrewartha, Hattie Vevers-Steele after whom the new species described below is named, drew the attention of one of us (LAM) to some specimens of a thrips species taken from bracken near Adelaide during her studies on Australian Thysanoptera (see Mound, 1996). The specimens were in poor condition, and efforts at that time to locate the species in the field were not successful. However, during the past 10 years this thrips has been found to be widespread across southern Australia, but breeding only in the curled apices of the youngest fronds of bracken. This species was listed by Shuter and Westoby (1992) from a population of bracken near Sydney as “Anaphothripinae gen. et sp. THRIPS AND APHIDS ON AUSTRAL BRACKEN FERN indet’, but is here recognised as a new species of the widespread Old World genus Trichromothrips. However, within that genus it exhibits one remarkably deviant autapomorphy — the absence of any long setae on the pronotum. This thrips has been found only on Pteridium esculentum, as defined by Brownsey (1998), even when this has been found growing in association with other ferns that are superficially similar, such as the closely related Hypolepis muelleri (Dennstaedtiaceae), or young specimens of the more distantly related tree fern Dicksonia antarctica (Dicksoniaceae). No thrips have been found on any species of Hypolepis, although Scirtothrips frondis Hoddle and Mound breeds abundantly on the youngest fronds of Dicksonia and has also been taken on a species of Cyathea (Hoddle and Mound, 2003). Trichromothrips Priesner Trichromothrips Priesner, 1930: 9. Type species T. bellus Priesner. Bhatti (2000) has fully defined and reviewed this genus, synonymising the genus Dorcadothrips Priesner and providing a key to identify the 27 included species. Of these, 24 are from the Old World, between Africa and Queensland but mostly from South East Asia. The other three species, two from Hawaii and one widespread, may also have come originally from the Oriental region. The collection data for most of the species are probably not reliable indicators of the plants on which these thrips breed, but two species (T. billeni Strassen and T. bilongilineatus Girault) are associated with ferns (Mound, 2002b), and in the region of Japan around Tokyo and Yokahama, T. alis Bhatti or a closely related species is found on a species of Polystichum (Dryopteridaceae). Finally, three related genera of Thripidae are also associated with ferns, Laplothrips Bhatti, Octothrips Moulton and Pteridothrips Priesner (Mound, 2002b). Members of these four genera are unusual in bearing a pair of setae on the dorsal apical margin of the first antennal segment. This character state is also shared by species in the following genera of Thripidae, although none involves fern-living species: Alathrips Bhatti, Bregmatothrips Hood, Ceratothripoides Bagnall, Craspedothrips Strassen, Diarthrothrips Williams, Furcithrips Bhatti, Megalurothrips Bagnall, Mycterothrips Trybom, Odontothrips Amyot and Serville, Odontothripiella Bagnall, Pezothrips Karny, Sorghothrips Priesner, Watanabeothrips Okajima, Yoshinothrips Kudo. Moreover, although the two species comprising the Oriental genus Bathrips Bhatti lack this pair of setae on the dorsal apical margin of 68 the first antennal segment, they share many other character states with Trichromothrips species, and these two genera are possibly closely related. Trichromothrips veversae sp.n. Holotype 2 macroptera, Australian Capital Territory, Woods Reserve, from young fronds of Pteridium esculentum, 6.xii.2002 (LAM 4244), in ANIC, CSIRO Entomology, Canberra. Paratypes: 2 males, 17 females, same host, date and locality as holotype (Masumoto, Mound and Wells); 3 females at same locality but 16.1.1999 (LAM 3664). Specimens excluded from the type series were collected widely in southern Australia, including Tasmania, Western Australia, New South Wales, and the Australian Capital Territory (see Distribution below). Female _macroptera Colour: body yellow with orange pterothorax, ocelli bright red, antennae brown, abdomen with transverse light brown markings, wings shaded; colour of cleared and mounted specimens yellow, tergites shaded anteromedially and along antecostal line, IX and X shaded, mesonotum and metanotum weakly shaded; head and antennal segment I pale, segments If] to VIL almost uniformly dark brown with extreme base of segments III to V slightly paler, I paler than segment III; all legs greyish brown; fore wing and scale greyish brown, but base of fore wing paler. Structure: Head slightly wider than long, not prolonged in front of eyes, with a few transverse striae posteriorly on vertex (Fig. 1); ocellar setae I absent, setae III no longer than length of an ocellus and arising between anterior margins of posterior ocelli; three pairs of postocular setae, pairs I and II close together behind ocelli; ventral surface of head with 5 pairs of setae between compound eyes anterior to anterior tentorial pits; mouth-cone rounded, maxillary palpi 3- segmented; compound eyes without pigmented facets. Antenna 8-segmented (Fig. 3); forked sense-cones on If] and IV exceptionally stout; segment I with 2 dorsal apical setae; II with weak microtrichia laterally only, If] to VI with a few large microtrichia on dorsal and ventral surfaces; III with 2 dorsal and 2 ventral setae. Pronotum medially with few or no lines of sculpture and 4 to 10 discal setae; posterior margin with five pairs of setae, none of which is longer than the discal setae. Mesonotum with weak transverse lines of sculpture, without campaniform sensilla near anterior margin, median pair of setae far ahead of posterior margin. Metanotum (Fig. 2) medially without sculpture and one pair of small setae far from anterior Proc. Linn. Soc. N.S.W., 125, 2004 L. MOUND AND M. MASUMOTO Figure 1. Trichromothrips veversae, head and pronotum. margin, without campaniform sensilla. Prosternal ferna not divided; mesothoracic sternopleural suture not developed; meso- and metasternum each with well- developed spinula. All tarsi 2-segmented. Forewing veinal setae short, less than half width of wing in length; first vein with about 8 setae near base and 2 (rarely 3) setae near apex; second vein with about 10 setae; posterior fringe cilia wavy; forewing scale with 4 marginal setae. Abdominal tergites without posteromarginal craspeda or lateral ctenidia; tergites II to VIII without sculpture medially, lateral to seta S2 with about 7 anastomosing transverse lines bearing tuberculate microtrichia; tergite VIII without posteromarginal comb; tergite IX with paired campaniform sensilla posteromedially; tergite X undivided; pleurotergites without discal seta, sculpture similar to lateral areas of tergites. Sternites without discal setae; sternite II with two pairs of posteromarginal setae, sternites [II to VII with three pairs, on VII all three pairs arise in front of sternal posterior margin. Measurements (holotype female in um with small paratype female in parentheses): Body length 1400 (1100). Head, length 90 (85); width 125 (105). Pronotum, length 105 (95); width 160 (130); posteromarginal setae 15 (12). Forewing, length 750 (650). Antennal segments 25, 32, 50, 57, 40, 43, 10, 17 (25, 30, 40, 47, 35, 37, 7, 15). Figure 2. Trichromothrips veversae, mesonotum and metanotum. Male aptera Colour paler than female. Structure similar to female except: forked sense-cones on antennal segments III and IV small and slender; one of three available males lacks ocellar setae II; mesonotum transverse with 4 or 5 setae near lateral margins; pleurotergal sutures weakly developed; tergite IX Figure 3. Trichromothrips veversae, antenna. Proc. Linn. Soc. N.S.W., 125, 2004 THRIPS AND APHIDS ON AUSTRAL BRACKEN FERN posterior margin with horn-like paired drepanae extending beyond segment X; sternites III to VIII each with about 50 small, irregularly arranged, glandular areas, marginal setae arising at margin on all sternites. Measurements (paratype male in um). Body length 1000. Head, length 83; width 100. Pronotum, length 85; width 130; posteromarginal setae 15. Tergite IX drepanae length 60. Antennal segments 25, 30, 37, 40, 32, 37, 7, 15. Larva Il. Colour pale yellow with red eyes, progressively developing extensive pale red hypodermal pigment in meso- and metathorax and anterior abdominal segments, body usually turning deep yellow progressively; major dorsal setae parallel- sided with bluntly square apices, 3 pairs on head, 6 pairs on pronotum, 3 pairs on abdominal tergites II — VIII, 2 pairs on IX, antennal II with 2 pairs of similar but smaller setae; setae on tergite X and abdominal sternites with apices acute; sternite [X posterior margin with row of about 30 small tooth-like tubercles. Systematic relationships Currently, this new species cannot be placed in any of the 10 species-groups distinguished by Bhatti (2000) within Trichromothrips, although it shares with the other 27 species the many character states listed by that author in his diagnosis of the genus. In contrast to those species, it lacks any long pronotal setae, the metasternal spinula is well developed not weak, and females have unusually stout antennal sense cones. In Australia, only one other species of Trichromothrips has been collected in good numbers: T. bilongilineatus (Girault) from ferns near Gosford (Mound, 2002a). Of the other two members of the genus listed from Australia, the record of T. xanthius (Williams) is based on one female taken in quarantine in North America but labelled as coming from Australia (Mound, 1996), and T. obscuriceps (Girault) is known from a single sample apparently taken on Crinum lilies near Brisbane. The genus is probably well established in northern Australia, but only a few specimens are available, representing two further unidentified species, swept from grasses near Darwin. All of these species have long pronotal posteroangular setae. The lack of long pronotal setae gives T. veversae the superficial appearance of an Anaphothrips species. This is another example of the ineffective supra-generic classification within the subfamily Thripinae, in which traditional subtribal names such as Aptinothripina do not refer to definable groups (Mound, 2002c), despite their continued use by various 70 authors (eg. Vasiliu-Oromulu et al. 2001). There are several unrelated Thripinae genera in which species usually have two pairs of long pronotal setae, but in which one or more species have these setae no longer than the discal setae and are thus “Anaphothripine” in appearance, eg. Dichromothrips Priesner, Pseudanaphothrips Karny and Thrips Linnaeus. The presence or absence of long setae on the pronotum was recognised as a poor indicator of phylogenetic relationships by Mound and Palmer (1981), who proposed a series of informal genus- groups within the Thripinae. These authors included Scolothrips Hinds, a genus of predatory thrips, in their Dorcadothrips genus-group (Mound and Palmer, 1981). Scolothrips species resemble some Trichromothrips species in general appearance, for example the pale slender body and bulging compound eyes, but they have very long ocellar setae and the pronotum bears six pairs of elongate setae. Moreover, the dorsal apical margin of the first antennal segment does not bear a pair of setae, and the mesosternal sternopleural sutures are weakly developed. The character state on the first antennal segment discussed above suggests that the genus-groups recognised by Mound and Palmer (1981) require reappraisal. Distribution and host records T. veversae has been found to be locally abundant in many parts of southern Australia, including Western Australia near Albany, Tasmania near Hobart, and various sites in South Australia (Adelaide Hills; Cox’s Scrub south of Adelaide; and Kangaroo Island). It is abundant in the mountains of the ACT, and is widely distributed in the eastern forests of New South Wales from near Eden to the Blue Mountains. It possibly occurs even further north, but a sample taken from Preridium at Beerwah, north of Brisbane, yielded only Scirtothrips dobroskyi Moulton (Hoddle and Mound, 2003). In a survey of the insects associated with bracken in New Guinea, Kirk (1977) does not mention thrips, but since thrips on ferns are associated only with very young fronds, or even with croziers that are not yet fully expanded, these minute insects are often difficult to detect. Similarly, the list given by Balick et al. (1978) of insects taken from ferns worldwide is based on a survey of published records, derived mainly from general collecting, and some of the thrips species listed are fungus-feeders, not fern-feeders. Mound (2002b) emphasised that _ several published records of thrips on ferns are based on single samples or even single specimens, and thus cannot be relied on to indicate a host relationship. In Japan, the common species of bracken fern is considered also to represent Pteridiium esculentum Proc. Linn. Soc. N.S.W., 125, 2004 L. MOUND AND M. MASUMOTO and, as indicated above, the aphid species Shinjia orientalis has been recorded from this plant in Japan as well as Australia. However, searches for thrips on substantial populations of bracken in Japan, particularly near Narita City, have failed to discover Trichromothrips veversae. At Crafers in the Adelaide Hills, South Australia, a substantial population of adults and larvae of Thrips imaginis Bagnall was found on bracken fronds in an open field during December 2002, together with a few larvae of Trichromothrips veversae. However, this seems to be a rare host association for the highly polyphagous Australian Plague Thrips. At several sites, near Adelaide and on Kangaroo Island, larvae of T. veversae were found bearing up to 12 larval Eucharitidae (Hymenoptera). This is presumably a phoretic association, but no observations were made on associated ants, the probable host of these small wasps. ACKNOWLEDGEMENTS The authors are grateful to the Chief, CSIRO Entomology, for providing research facilities at Canberra, to Victor Eastop of the Natural History Museum, London, for information on aphids associated with ferns, and to Alice Wells for field assistance and comments on the manuscript. John Thomson of the Royal Botanic Gardens Sydney kindly drew our attention to several references. Mr. Kenji Morita, Head of Yokohama Plant Protection Station, and Mr. Tetsuo Imamura, Chief of Identification Section, Yokohama Plant Protection Station, kindly facilitated a study visit to Canberra by M. Masumoto, and Mr. Shigeo Aochi in Narita City, and Mr. Yuji Yoshida in Chiba City provided help in searching for thrips on Pteridium in Japan. REFERENCES Balick, M., Furth, D.G. and Cooper-Driver, G. (1978). Biochemical and evolutionary aspects of arthropod predation on ferns. Oecologia 35, 55- 89. Bhatti, J.S. (2000). Revision of Trichromothrips and related genera (Terebrantia: Thripidae). Oriental Insects 34, 1-65. Brownsey, P.J. (1989). The taxonomy of Bracken (Pteridium: Denstaedtiaceae) in Australia. Australian Systematic Botany 2, 113-128. Brownsey, P.J. (1998). Denstaedtiaceae, pp. 214-228 in Flora of Australia volume 48. Ferns, Gymnosperms and Allied Groups. CSIRO, Melbourne. Hoddle, M.S. and Mound, L.A. 2003. The genus Scirtothrips in Australia (Insecta, Thysanoptera, Thripidae). Zootaxa 268, 1-40. Proc. Linn. Soc. N.S.W., 125, 2004 Kirk, A.A. (1977). The insect fauna of the weed Preridium aquilinum (L.) Kuhn (Polypodiaceae) in Papua New Guinea: A potential source of biological control agents. Journal of the Australian Entomological Society 16, 403-409 Mabberley, D.J. (1997). The Plant Book. Second edition. Cambridge University Press, Cambridge. Mound, L.A. (1996). Thysanoptera, pp 249-336, 397-414 (Index), in Wells, A., Zoological Catalogue of Australia. Volume 26. Psocoptera, Phthiraptera, Thysanoptera. Melbourne. CSIRO Australia. Mound, L.A. (2002a). Thrips and their host plants: new Australian records (Thysanoptera: Terebrantia). Australian Entomologist 29, 49-60. Mound, L.A. (2002b). Octothrips lygodii sp.n. (Thysanoptera, Thripidae) damaging weedy Lygodium ferns in south-eastern Asia, with notes on other Thripidae reported from ferns. Australian Journal of Entomology 41, 216-220. Mound, L.A. (2002c). The Thrips and Frankliniella genus-groups: the phylogenetic significance of ctenidia, pp. 379-386 in Mound L.A. and Marullo, R. [eds] Thrips and Tospoviruses. Proceedings of the 7" International Symposium on Thysanoptera. Australian National Insect Collection, Canberra. Mound, L.A. and Palmer, J.M. (1981). Phylogenetic relationships between some genera of Thripidae (Thysanoptera). Entomologica Scandinavica 15, 153-17. Priesner, H. (1930). Contribution towards a knowledge of the Thysanoptera of Egypt, Ill. Bulletin de la Société Royal Entomologique d’Egypte 14, 6- 15. Thomson, J.A. (2000). Morphological and genomic diversity in the genus Pteridium (Dennstaedtiaceae). Annals of Botany 85, 77-99. Vasiliu-Oromulu, L. zur Strassen, R. and Larsson, H. (2001). The systematic revision of Thysanoptera _ species from the Swedish fauna and their geographical distribution. Entomologia romana 6, 93-101. 71 ‘ _ a A Le t . mrs Nay " ath ; is, at ’ 8 i , ar’ et be ee i : _ oat: t ry - - . >" x ee ee a J na ms iy ; : ? p 7 { - { » + g , ih “ial os ‘ ) = Zi e “mai we pieces enema | Smid ¢ “es pele TS brian, 50) longitudinal series along length of spine. Remarks Lindley (2003b) described the spines of Phyllacanthus imperialis var. javana K. Martin 1885 and Phyllacanthus sp. from the Lower Pliocene Kairuku Formation, Yule Island. Unfortunately, the characters diagnostic of these species, including spine collar length and the number of ridges on the distal part of the spine, are not visible on ANU 60637. Subclass EUECHINOIDEA Bronn 1860 Superorder ECHINACEA Claus 1876 Order TEMNOPLEUROIDA Mortensen 1942 Family TOXOPNEUSTIDAE Troschel 1872 Genus TOXOPNEUSTES A. Agassiz 1841 Synonymy Boletia Desor 1846, p. 362. Proc. Linn. Soc. N.S.W., 125, 2004 | 121 FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA Type species Echinus pileolus Lamarck 1816, p. 45, by original designation. Toxopneustes pileolus (Lamarck 1816) Synonymy Echinus pileolus Lamarck 1816, p. 45. Toxopneustes pileolus, A. Agassiz 1841, p. 7; H.L. Clark 1925, p. 123; Mortensen 1943a, p. 472; A.M. Clark and Rowe 1971, p. 156; Rowe and Gates 1995, p. 258. Mortensen (1943a: 472) lists additional synonymies. Material and locality Single naked test, B20022, from the vicinity of Cape Gazelle, New Britain, East New Britain Province, PNG. Remarks Toxopneustes pileolus (Lamarck 1816) is widely distributed throughout the Indo-West Pacific (Mortensen 1943a; A.M. Clark and Rowe 1971; Miskelly 2002). Genus TRIPNEUSTES L. Agassiz 1841 Type species Echinus granularis Lamarck 1816, p. 44, by original designation. Tripneustes gratilla (Linnaeus 1758) Synonymy Echinus gratilla Linnaeus 1758, p. 664. Tripneustes gratilla, H.L. Clark 1925, p. 124; Mortensen 1943a, p. 500; A.M. Clark and Rowe 1971, p. 156; Rowe and Gates 1995, p. 259. Mortensen (1943a: 500) lists additional synonymies. Material and locality Single naked test, B20023, from the vicinity of Cape Gazelle, New Britain, East New Britain Province, PNG. Remarks Tripneustes gratilla (Linnaeus 1758) is widely distributed throughout the Indo-West Pacific (Mortensen 1943a; A.M. Clark and Rowe 1971). Previous records from the Pacific include the Marshall Islands, Norfolk Island, Hawaiian Islands, Kermadec Islands, Solomon Islands, Fiji and Hood Lagoon, south coast of Papua (H.L. Clark 1925; Mortensen 1943a; A.M. Clark and Rowe 1971; Miskelly 2002). Order ECHINOIDA Claus 1876 Family ECHINOMETRIDAE Gray 1825 Genus ECHINOMETRA Gray 1825 Synonymy Ellipsechinus Litken 1864, p. 165. Plagiechinus Pomel 1883, p. 78. Mortensenia Doderlein 1906, p. 233. Type species Echinus lucunter Linnaeus 1758, p. 665, by original designation. 122 Proc. Linn. Soc. N.S.W., 125, 2004 I.D. LINDLEY Echinometra mathaei (de Blainville 1825) Synonymy Echinus lucunter Lamarck 1816, p. 50 (non E. lucunter Linnaeus). Echinometra mathaei, H.L. Clark 1925, p. 143; H.L. Clark 1932, p. 216; Mortensen 1943b, p. 381; H.L. Clark 1946, p. 332; A.M. Clark and Rowe 1971, p. 157; Rowe and Gates 1995, p. 211. Mortensen (1943b: 381) lists additional synonymies. Material and localities Fourteen naked tests from Gargaris village, northern coast of Malendok Island, Tanga Group, New Ireland Province, PNG; one partly naked test from beach at Ralum, Blanche Bay, East New Britain Province, PNG; one naked test from Penlolo village, south coast of New Britain, West New Britain Province, PNG; one naked test, B 20016, from Cape Gazelle, New Britain, East New Britain Province, PNG. Remarks Echinometra mathaei (de Blainville 1825) is a long ranging species, recorded from late Lower Miocene- early Middle Miocene rocks in the western and eastern Mediterranean Sea (Negretti et al. 1990). Extant E. mathaei is one of the most widely distributed echinoids, occurring throughout tropical-subtropical waters of the Indo-West Pacific (Mortensen 1943b; A.M. Clark and Rowe 1971). H.L. Clark (1908) recorded the species from Sorong, west New Guinea and Miskelly (2002) recorded it from the Solomon Islands. This record indicates a wide distribution throughout the Bismarck Archipelago (Tanga Group, New Ireland; Blanche Bay, New Britain; and south coast New Britain). Genus HETEROCENTROTUS Brandt 1835 Synonymy Acroladia L. Agassiz and Desor 1846, p. 373. Type species Echinus mamillatus Linnaeus 1758, p. 664, by subsequent designation of Pomel 1883, p. 77. Heterocentrotus mammillatus (Linnaeus 1758) Synonymy Echinus mamillatus Linnaeus 1758, p. 664. Heterocentrotus mammillatus, H.L. Clark 1925, p. 147; Mortensen 1943b, p. 409; H.L. Clark 1946, p. 333; A.M. Clark and Rowe 1971, p. 158; Rowe and Gates 1995, p. 213. Mortensen (1943b: 409) lists additional synonymies. Material and locality A single naked test, B 20017, and unlabelled isolated spines (housed in the East New Britain Historical and Cultural Centre, Kokopo) from Cape Gazelle, New Britain, East New Britain Province, PNG; an isolated primary spine, ANU 60648, from Nosnos village, Boang Island, Tanga Group, New Ireland Provine, PNG. Remarks Heterocentrotus mammillatus (Linnaeus 1758) is widely distributed throughout the Indo-Pacific, from the Gulf of Suez and Madagascar to the Hawaiian Islands and Fiji (Mortensen 1943b). It is recorded from the Solomon Islands by Miskelly (2002). The largest test of H. mammillatus noted by Mortensen (1943b) has a long diameter of 82 mm, with most individuals having diameters of 72 mm or less. The long diameter of the Cape Gazelle test is 72 mm. The Tanga spine has a length of 74 mm and, given that the primary spines of H. mammillatus usually do not exceed the long diameter of the test (Mortensen 1943b), appears to have come from a relatively large individual. Proc. Linn. Soc. N.S.W., 125, 2004 j 123 FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA Genus HELIOCIDARIS L. Agassiz and Desor 1846. Synonymy Toxocidaris A. Agassiz 1863, p. 22. Type species Echinus tuberculatus Lamarck 1816, p. 50, by original designation. Diagnosis Low hemispherical echinoids, widest at circular ambitus. Ambulacral plates with 7 or more pore-pairs to each plate; arcs may be irregularly double; expanded poriferous tracts of the flattened adoral surface are petaloid. Oculars I and IV usually insert. Gill-slits are shallow (Philip 1965; Fell and Pawson 1966). Remarks Heliocidaris L. Agassiz and Desor 1846 is distributed along the southern coasts of Australia, northern New Zealand, Kermadec Islands and Lord Howe Island (Mortensen 1943a). Two species are included in the genus by Mortensen (1943a), viz: Heliocidaris tuberculata (Lamarck 1816) and Heliocidaris erythrogramma (Valenciennes 1846) and, given their similar morphologies, he has questioned whether they are really conspecific. Anthocidaris Liitken 1864 is a closely allied genus (only known species Anthocidaris crassispina [A. Agassiz 1863]) from the coasts of southern Japan and China, distinguished from Heliocidaris by the spicules of the tubefeet (Mortensen 1943a). On the status of Anthocidaris, Mortensen (1943a: 328) questioned whether the genus should be merged into Heliocidaris. Philip (1965) described the only known fossil representative of the genus, Heliocidaris ludbrookae Philip 1965 from the Lower-early Middle Miocene (Longfordian-Batesfordian) of southeastern Australia. Heliocidaris robertsi sp. nov. Figs 4, 5a-e Diagnosis Test low hemispherical, somewhat inflated above. Ambulacral plates with 12 pore-pairs per plate; ambital and aboral pore-arcs doubled. Ambulacral and interambulacral plates relatively large; each bearing a primary tubercle and numerous secondary tubercles; aureoles of primaries not in contact. Primary tubercles of ambital and aboral ambulacral plates with an aborally positioned secondary tubercle. Etymology Named for Mr Michael Roberts, amateur conchologist of Kokopo, East New Britain Province, PNG. Material and locality Single naked test, ANU 60654, from the vicinity of Cape Gazelle, New Britain, East New Britain Province, PNG. Description Test low hemispherical, somewhat inflated above, widest at circular ambitus. The oral side is flattened, scarcely sunken towards the peristome. Only specimen of 38 mm Figure 4. Heliocidaris robertsi sp. nov. Cape Gazelle area, East New Britain gjameter. Province. 4a-b, plating diagrams at ambitus for interambulacrum, The pore zones are ambulacrum. conspicuously petaloid on the 124 Proc. Linn. Soc. N.S.W., 125, 2004 I.D. LINDLEY Figure 5. Heliocidaris robertsi sp. nov. Cape Gazelle area, East New Britain Province. 5a-e, ANU 60654, aboral, oral, lateral views. Bar scale = 10 mm; ambulacral plating at ambitus (refer to Fig. 4b for plating diagram). Bar scale = 5 mm; apical disc. Bar scale = 2.5 mm. Proc. Linn. Soc. N.S.W., 125, 2004 FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA oral surface, about 1.5-2 times the width of interporiferous zone. The pore-series in this area are almost horizontal and are separated by secondary tubercles forming a single prominent vertical series; scattered miliary tubercles are also present. In the ambital region there are 12 pore-pairs arranged in double arcs (Fig. 4). Above the ambitus the pore-zones become much narrower. Primary tubercles in the ambital zone are large, almost as large as the interambulacral primaries; aureoles of adjacent primaries in each vertical series widely separated. Sutures between adjacent plates are seen very distinctly on the outer adoral side of the boss. Each ambulacral plate at and above the ambitus has a prominent secondary tubercle positioned aborally to the primary tubercle; 4-5 other secondary tubercles are also present. Miliaries tend to be arranged along the perradial sutures of ambital and superoambital ambulacrals; elsewhere on each plate only a sparse covering of miliaries is present. The interambulacral primaries are large, forming prominent series aborally; their aureoles are distinctly separated, leaving a broad space at the upper edge of each plate, occupied by several small tubercles and miliartes. Usually sutures between adjacent plates are close to, but do not cross, aureole of successive tubercle. In the median space there is on the oral surface and in the ambital region a conspicuous double series of secondary tubercles about half the size of the ambulacral primaries. Below the ambitus all the tubercles decrease rapidly in size, with the secondaries disappearing, and only the primaries continuing to the peristome. The apical system is small, only about 18 percent of the test diameter. There is typically one large tubercle on each genital plate, except the very large madreporite, and a scattering of small tubercles over the remainder (Fig. 5e). Ocular I and IV are broadly insert. The peristome is very small, about 29 percent of test diameter. Gill-slits shallow. Details of spines and pedicellarie unknown. Remarks Heliocidaris robertsi sp. nov. is readily distinguished from H. tuberculata and H. erythrogramma and the closely allied A. crassispina by its possession of double pore-arcs on the adoral surface. The double pore- arcs of H. robertsi are very similar to those of Heterocentrotus trigonarius (Lamarck 1816), figured by Mortensen (1943a: fig. 132c) and Fell and Pawson (1966: fig. 324, 7c). However, any resemblance between the new species and H. trigonarius is easily discounted because of the latter’s possession of a distinctly elongated test and a significantly larger peristome (51 percent of test diameter). The biogeographical position of H. robertsi is noteworthy in that it is the tropical representative of two closely allied temperate water genera, Heliocidaris, a very common form restricted to southern Australia and New Zealand, and Anthocidaris, an equally common form restricted to Japan and China. Pore-arc doubling is almost as strongly developed in other echinometrids including Colobocentrotus Brandt 1835 and Zenocentrotus A.H. Clark 1931, and incipient development may also been seen in Echinometra Gray 1825 (Mortensen 1943a: 281). All three genera possess an elliptical or oblong ambitus. The functional significance of doubling of pore-arcs in compound plates relates to (a) increasing the area over which tube-feet are spread, and thereby increasing respiratory and feeding efficency (Mortensen 1943a; Woods 1958; Durham 1966; A.M. Clark 1968) and (b) strengthening of the test (Durham 1966). The doubling of pore-arcs on the aboral surface of H. robertsi greatly increases the number of tube-feet in this area, not only aiding in improved respiration, but allowing it to catch food particles falling onto its upper surface. With such adaptations to its upper surface, the echinoid may have been a reef rock borer, inhabiting a hole perhaps several centimetres deep. Superorder GNATHOSTOMATA Zittel 1879 Order HOLECTYPOIDA Duncan 1889 Suborder ECHINONEINA HL. Clark 1925 Family ECHINONEIDAE Agassiz and Desor 1847 Genus ECHINONEUS Leske 1778 Synonymy Echinanaus Gray 1825, p. 7 (nom. van.). Pseudohaimea Pomel 1885, p. 118. Koehleraster Lambert and Thiéry 1921, p. 331. Type species Echinoneus cyclostomus Leske 1778, by subsequent designation of H.L. Clark 1917, p. 101. 126 Proc. Linn. Soc. N.S.W., 125, 2004 I.D. LINDLEY Remarks Echinoneus Leske 1778 is an Oligocene-Recent form, with some ten fossil species described from the Oligocene and Miocene of Europe (Mortensen 1948a; Wagner and Durham 1966). Two Recent species are known, viz. Echinoneus cyclostomus Leske 1778 and Echinoneus abnormalis de Loriol 1883, distinguished by the presence or absence of imperforate primary tubercles and well developed glassy tubercles. Recent forms are distributed throughout the West Indies, Indo-Pacific and Australia. Mortensen (1948a) considered that many of the fossil species are very difficult to distinguish and may in fact be Recent E. cyclostomus. Kchinoneus sp. Fig. 3d Material One poorly preserved test, ANU 60640. Locality and horizon Nosnos village, Boang Island, Tanga Group, New Ireland Province, PNG. Grid reference 296246 Tanga 1:100 000 Sheet 9591 (Edition 1). Unnamed poorly compacted bioclastic limestone, Pleistocene-Holocene (Wallace et al. 1983). Description Test ovoid, moderate size, measuring 23 x 17 x 11.5 mm; oral surface weakly concave. Ambulacra narrow, not petaloid. Other details of ambulacra unknown. Details of interambulacra unknown. Details of tubercles unknown. Apical and periproctal systems unknown. Remarks The lack of well preserved tubercles on this specimen makes it difficult to assign a species. Kchinoneus cyclostomus Leske 1778 Synonymy Echinoneus cyclostomus Leske 1778, p. 173; H.L. Clark 1925, p. 177; H.L. Clark 1946, p. 353; Mortensen 1948a, p. 75; A.M. Clark and Rowe 1971, p. 158; Rowe and Gates 1995, p. 215. Mortensen (1948a: 75) lists additional synonymies. Material and locality Twelve naked tests, including ANU 60641, from Gargaris village, northern coast of Malendok Island, Tanga Group, New Ireland Province, PNG; one naked test, B 20021, from Cape Gazelle, New Britain, East New Britain Province, PNG. Remarks Echinoneus cyclostomus Leske 1778 is the only known case of a (tropical) cosmopolitan echinoid, having been recorded from the West Indies, Ascension (but not the African west coast) and the Indo-Pacific-East Africa (Zanzibar, Natal), Madagascar to the Pacific islands (Funafuti, Palmyra, Hawaiian Islands), and from Japan to Queensland (Great Barrier Reef) and Lord Howe Island (Mortensen 1948a). Miskelly’s (2002) record of E. cyclostomus from the Solomon Islands represents the nearest previous record to that from the Tanga Group and Cape Gazelle. Echinoneus abnormalis de Loriol 1883 Synonymy Echinoneus abnormalis de Loriol 1883, p. 41; H.L. Clark 1917, p. 102; H.L. Clark 1925, p. 176; Mortensen 1948a, p. 80; A.M. Clark and Rowe 1971, p. 158. Koehleraster abnormalis Lambert and Thiéry 1921, p. 331. Proc. Linn. Soc. N.S.W., 125, 2004 | 127 FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA Material and locality One naked test, ANU 60641, from Gargaris village, northern coast of Malendok Island, Tanga Group, New Ireland Province, PNG. Remarks This species is represented by a single naked test measuring 30 x 22.5 x 15 mm. Echinoneus abnormalis de Loriol 1883 is distinguished from E. cyclostomus by possessing perforated, non-glassy spine tubercles. The apical system of the Tanga specimen is distinctly anterior to that of co-occurring specimens of the much more common E. cyclostomus. Echinoneus abnormalis has a restricted distribution, known from Mauritius (type locality), Kei Islands, Palmyra Island, Banda, Ellice Islands and the Hawaiian Islands (Mortensen 1948a; A.M. Clark and Rowe 1971). The recent record of E. abnormalis from the vicinity of Raine Island on the northern Great Barrier Reef (Gibbs et al. 1976) represents the first from Australasian waters. The record from the Tanga Group is the second from the East Indies. The species is observed to be sympatric with the much more common E. cyclostomus in many localities, a fact Gibbs et al. (1976) suggested may have resulted in it having gone unrecognised in samples. Mortensen (1948a: 81) considered that the two species probably didn’t live together at the same localities. Of the 15 specimens of Echinoneus collected from the Malendok Island locality, only one was an E. abnormalis, suggesting that in this case, the species’ apparent rarity may be related to different niches within the same locality. Order CLYPEASTEROIDA A. Agassiz 1872 Suborder CLYPEASTERINA A. Agassiz 1872 Family CLYPEASTERIDAE L. Agassiz 1835 Genus CLYPEASTER Lamarck 1801 Type species Clypeaster rosaceus (Linnaeus 1758), by subsequent designation of Desmoulins 1835. Clypeaster reticulatus (Linnaeus 1758) Synonymy Lindley (2003a) lists previous synonymies. Material Single naked test, B20020, from the vicinity of Cape Gazelle, New Britain, East New Britain Province, PNG. Remarks Clarification of Lindley’s (2003a) statement on the distribution of Clypeaster reticulatus (Linnaeus 1758) is needed. The species is a very common Indo-West Pacific echinoid, distributed in the western Indian Ocean and the Red Sea, throughout the East Indies and east into the Pacific Ocean to the Hawaiian Islands (A.M. Clark and Rowe 1971). Previous south Pacific records of the species have been made by A. Agassiz (1863), Mortensen (1948b) and A.H. Clark (1954) from the Gilbert Islands, New Caledonia and Marshall Islands, respectively. Mortensen’s (1948b) New Caledonian record has not been confirmed by De Ridder (1986: 29). McNamara and Kendrick (1994) have also recorded the species from Barrow Island, northwestern Australia. The species is known from fossil in Java (Lower Miocene), Yule Island, PNG (Lower Pliocene), East Africa (Pliocene- Pleistocene) and the New Hebrides (Pleistocene) (Mortensen 1948b; Lindley 2003a). Family ARACHNOIDAE Duncan 1889 Subfamily ARACHNOIDINAE Duncan 1889 Genus ARACHNOIDES Leske 1778 Synonymy Echinarchinus Leske 1778, p. 217. 128 Proc. Linn. Soc. N.S.W., 125, 2004 I.D. LINDLEY Type species Echinus placenta Linnaeus 1758, p. 666, ICZN 1954. Arachnoides placenta (Linnaeus 1758) Synonymy Echinus placenta Linnaeus 1758, p. 666. Arachnoides placenta (Linnaeus 1758): L. Agassiz 1841, p. 94; Bell 1899, p. 136; H.-L. Clark 1925, p. 154; H.L. Clark 1946, p. 340; A.M. Clark and Rowe 1971, p. 161; Rowe and Gates 1995, p. 176. Mortensen (1948b) lists additional synonymies. Material and locality Single naked test, B20018, from the vicinity of Cape Gazelle, New Britain, East New Britain Province, PNG. Remarks Arachnoides placenta (Linnaeus 1758) is a common littoral species throughout the East Indies and the south Pacific (Mortensen 1948b; A.M. Clark and Rowe 1971). The first record of the species from the Bismarck Archipelago is that of Bell (1899) from an unspecified locality in New Britain. Suborder LAGANINA Mortensen 1948 Family LAGANIDAE A. Agassiz 1873 Genus LAGANUM Link 1807 Synonymy Lagana Gray 1825, p. 427. Type species Laganum petalodes (= Echinodiscus laganum Leske 1778, p. 204), by original designation. Laganum laganum (Leske 1778) Synonymy Laganum Bonani Klein 1734, p. 25. Echinodiscus laganum Leske 1778, p. 204. Laganum laganum, Mortensen 1948b, p. 312. Laganum depressum, Lindley 2001, p. 130. Mortensen (1948b: 312) list previous synonymies. Material and locality Single test, ANU 60649, from Penlolo village, south coast of New Britain, West New Britain Province, PNG. Remarks Laganum laganum (Leske 1778) is distinct with its pentagonal test with thick, swollen edges, and an oblong-elongate periproct situated midway between the mouth and test edge. The species is common in the East Indies, and is also recorded from Port Jackson and Tasmania (Mortensen 1948b). Mortensen (1948b) also recorded it from the Bismarck Archipelago (Table 2). H.L. Clark (1908) recorded the species from Saonek, Waigiou Island, in west New Guinea (Fig. 1) Suborder SCUTELLINA Haeckel 1896 Family ASTRICLYPEIDAE Stefanini 1911 Proc. Linn. Soc. N.S.W., 125, 2004 . 129 FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA Genus ECHINODISCUS Leske 1778 Type species Echinodiscus bisperforatus Leske 1778, p. 196. Kchinodiscus tenuissimus (L. Agassiz in Agassiz and Desor 1847) Synonymy Lobophora tenuissima L. Agassiz and Desor 1847, p. 136. Echinodiscus tenuissimus, Gray 1855, p. 20; H.L. Clark 1914, p. 71; H.-L. Clark 1925, p. 171; Mortensen 1948b, p. 411; A.M. Clark and Rowe 1971, p. 144 162; Rowe and Gates 1995, p. 185. Mortensen (1948b: 411) lists additional synonymies. Material and locality Two tests, B 20024 (naked) and B 20025 (with spines), from the vicinity of Cape Gazelle, New Britain, East New Britain Province, PNG. Remarks Echinodiscus tenuissimus (L. Agassiz in Agassiz and Desor 1847) is a widely distributed Indo-West Pacific form, occurring throughout the East Indies, northern Australia, southern Japan and the south Pacific (Mortensen 1948b; A.M. Clark and Rowe 1971). In the south Pacific, the species is recorded from Tanna, Vanuatu, (H.L. Clark 1925) and from New Caledonia (A.M. Clark and Rowe 1971). However, De Ridder (1986) only noted the occurrence of Echinodiscus bisperforatus Leske 1778 from New Caledonia. H.L. Clark (1925) observed that New Caledonian specimens of E. tenuissimus in the British Museum (Natural History) have a form more like E. bisperforatus. The Cape Gazelle specimens have very short lunules, about one quarter the length of the radius taken through them, and there is no difference in the tuberculation and spines of the ambulacral and interambulacral areas of the oral surface, both diagnostic characters of E. tenuissimus (Mortensen 1948b; A.M. Clark and Rowe 1971). Superorder ATELOSTOMATA Zittel 1879 Order SPATANGOIDA Claus 1876 Suborder HEMIASTERINA Fischer 1966 Family SCHIZASTERIDAE Lambert 1906 Genus SCHIZASTER L. Agassiz 1836 Type species Schizaster studeri L. Agassiz 1836, p. 185, by subsequent designation ICZN 1948. Remarks : McNamara and Philip (1980a, b) questioned the familial classification of the spatangoids used by Mortensen (1951) and Fischer (1966) and, in particular, the Family Schizasteridae. Within the Schizasteridae McNamara and Philip recognized genera sharing the gross morphological test features of Schizaster, viz. a posteriorly located apical system, with the apex of the test posterior to this; a long, typically sunken, poriferous frontal ambulacrum; and sunken petals, of which the posterior pair are markedly shorter than the anterior ones. Within this group, McNamara and Philip (1980a, b) included the genus Schizaster L. Agassiz 1836 (with its subgenera Dipneutes Arnaud 1891; Paraster Pomel 1869 and Ova Gray 1825 [= Diploraster Mortensen 1951)); Brisaster Gray 1855; Kina Henderson 1975; Moira L. Agassiz 1872 (= Moiropsis L. Agassiz 1881); and Proraster Lambert 1895 (= Hypselaster Clark 1917). The author accepts their emended diagnosis for Schizaster. Subgenus PARASTER Pomel 1869 Type species Schizaster gibberulus L. Agassiz 1847, by original designation of Pomel 1869, p. 14. 130 Proc. Linn. Soc. N.S.W., 125, 2004 I.D. LINDLEY Diagnosis Species of Schizaster with a small to moderate sized test, with a shallow frontal sinus. Apical system slightly posterior of centre. Frontal ambulacrum shallow with pore pairs inclined at about 45° and arranged in single rows. Anterior petals almost straight, diverging at an angle up to 110° (McNamara and Philip 1980a). Remarks There is difficulty in placing the Cape Gazelle species firmly within McNamara and Philip’s (1980a) subgenus Paraster Pomel 1869. This is particularly in relation to details of the anterior petals, their flexed nature and 80° angle of divergence, both characters diagnostic of subgenus Schizaster L. Agassiz 1836. The frontal ambulacrum does not possess the steeper sided walls typical of species referred to Schizaster (Schizaster) (McNamara and Philip 1980a). Furthermore, McNamara and Philip (1980a) noted that species referred to Schizaster (Schizaster) possess a more elongate, narrower test than those assigned to Paraster. The Cape Gazelle species is assigned to Schizaster (Paraster) by its possession of a small test, shallow frontal sinus, apical system slightly posterior of centre and shallow frontal ambulacrum with pore pairs inclined at about 45°. The species is probably morphologically transitional between the Paraster and Schizaster morphotypes. Schizaster (Paraster) ovatus sp. nov. Figs 6a-d Diagnosis A small species of Schizaster (Paraster) with a moderately depressed, ovoid test; apical system is 55 percent of test length from anterior, with four genital pores. Anterior ambulacrum relatively narrow and shallow; pore pairs inclined at about 45° and arranged in single rows; outer pores elongate, with similarly sized inner pores comma-shaped. Frontal sinus shallow. Etymology Ovatus L. egg- shaped, in reference to the form of the test, distinctive amongst the Schizasteridae. Material and locality Holotype ANU 60653, a complete naked test from the vicinity of Cape Gazelle, New Britain, East New Britain Province, PNG. Description Test of small size, elongate ovoid, with length x width x height measuring 34 x 28 x 18 mm; test length:width = 1.21, width:height = 1.55. Test moderately depressed, with apical system located 55 percent of test length from anterior; test highest posterior to apical system, along keel of ambulacrum Figure 6. Schizaster (Paraster) ovatus sp. nov. Cape Gazelle, East New Britain. y_ Oral surface is gently 6a-d, ANU 60653, aboral, oral, lateral, posterior views. Bar scale = 2.5 mm. Proc. Linn. Soc. N.S.W., 125, 2004 131 FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA convex. Apical system ethmolytic, depressed, with four genital pores, posterior pair being larger than anterior pair. Frontal ambulacrum long, shallow and narrow (12 percent of test length); pore pairs inclined at about 45° and arranged in a single row. Outer pores elongate, with similarly sized inner pores comma-shaped. Frontal sinus broad and shallow. Interambulacra II and III form sharp, high keels. Anterior petals diverging at angle of 80°; flexed distally and shallow, bearing pore pairs which are elliptical, widely spaced and conjugate; 26 pairs are present. Posterior petals are moderately long (occupying 21 percent of test length), bearing 18 pore pairs. Peripetalous fasciole is distinct, passing transversely between posterior petals and thickening at petal ends; the fasciole describes a concave arc between the extremities of the posterior and anterior petals, with an outwards flexure, corresponding with a constriction, forward of the apical system. Fasciole reaches maximum thickness at the extremities of the anterior petals. Peripetalous fasciole passes forward from anterior petals at about 60° before curving strongly to close with frontal ambulacrum; constrictions occur on interambulacral keel and adjacent to the abrupt curvature. Lateroanal fasciole is narrower than peripetalous fasciole and of more constant width. Lateroanal fasciole extends abaxially posteriorly from peripetalous fasciole at constriction between posterior and anterior petals; at ambitus it runs far below periproct, close to adoral surface. Peristome oval and slightly sunken; situated anteriorly, anterior tip of labrum 15 percent of test length from anterior. Anteriorly labrum is strongly curved; bounded by thick rim that degenerates laterally. Labrum as long as broad; posterior extension triangular, about as long as broad. Labrum carries several small tubercles anteriorly. Plastron is pear-shaped and broad, maximum width being 3/4 length. Plastron tubercles are arranged in curving rows. Periproct at mid-level on sub-truncate end of test. Periproct longitudinally elliptical, with a prominent narrow slit extending a short distance axially and aborally towards interambulacrum V, nearly reaching apical surface (Fig. 6d). Remarks Schizaster (Paraster) ovatus sp. nov. can be distinguished from other Schizaster-like heart urchins by its small, distinctively narrower and less inflated test, and long, shallow and narrow frontal ambulacrum. The test L:W and L:H ratios of 1.21 and 1.88 are larger than for most other echinoids of this group. The presence of four ~ genital pores would suggest that the holotype is a mature specimen. McNamara and Philip (1980b) noted that in Schizaster (Ova) myorensis McNamara and Philip (1980b) the onset of maturity, occurring at a test length of about 25 mm, followed the sequential opening of the first, second, third and fouth genital pores. Morphological adaptations in Schizaster-like heart urchins are related to a need to produce a more efficient current flow over the aboral surface in sediment of low permeability (McNamara and Philip 1980a). The posterior migration of the apex meant more water would flow over over the frontal sinus to the peristome; the deepening of the frontal ambulacrum and the frontal sinus assisted in channelling water to the peristome; and the deep and long frontal ambulacrum further enabled more-funnel-building tube feet to be accommodated, presumably in response to finer-grained sediment (McNamara and Philip 1980a). The weakly vaulted test of S.(P.) ovatus with its shallow, open frontal ambulacrum and shallow frontal sinus suggests the species was a shallow-burrower in coarse (permeable) shell gravel. — Suborder MICRASTERINA Fischer 1966 Family BRISSIDAE Gray 1855 Genus BRISSUS Gray 1825 Synonymy Bryssus Martens 1869, p. 128 (nom. van.). Brissus (Allobrissus) Mortensen 1950, p. 162. Type species Spatangus brissus unicolour Leske 1778, p. 248 by subsequent designation of ICZN, Op. 290 1948. Brissus (Brissus) latecarinatus (Leske 1778) Synonymy Brissus carinatus Gray 1825, p. 431; A. Agassiz 1872-74, p. 96, 596. Brissus latecarinatus (Leske 1778): H.L. Clark 1921, p. 153; H.L. Clark 1925, p. 219; H.L. Clark 132 Proc. Linn. Soc. N.S.W., 125, 2004 I.D. LINDLEY 1946, p. 375; Mortensen 1951, p. 514; A.M. Clark and Rowe 1971, p. 165; Gibbs et al. 1976, p. 135. Brissus (Brissus) latecarinatus: Rowe and Gates 1995, p. 187. Spatangus Brissus latecarinatus Leske 1778, p. 249. Mortensen (1951: 514) lists additional synonymies. Material and locality Three naked tests, ANU 60643-5, from Nosnos village, Boang Island, Tanga Group, New Ireland Province, PNG; one naked test, B 20014, from Cape Gazelle, New Britain, East New Britain Province, PNG. Remarks Brissus (Brissus) latecarinatus (Leske 1778) is a widely distributed species throughout the Indo-Pacific (Mortensen 1951; A.M. Clark and Rowe 1971). It is present on Australian coasts, from Queensland to Port Jackson, and is also known from Lord Howe Island (H.L. Clark 1946). Miskelly’s (2002) record of the species from the Solomon Islands is nearest to the present record in the Tanga Group. The largest specimen, ANU 60644 from the Tanga Group, measures 70 x 60 x 39 mm, considerably smaller than the largest known specimen, from Hawaii, measuring 130 x 108 x 74 mm (HL. Clark 1946). The shape of the periproct of the Tanga Group and Cape Gazelle specimens, somewhat pointed above and below, differs from the rounded periproct evident in specimens figured by Mortensen (1951: Plate XX XIII, fig. 7) and Miskelly (2002). In this respect, the Bismarck Sea specimens closely resemble Brissus (Allobrissus) agassizii Doderlein 1885 (Mortensen 1951: Plate XXXII, fig. 7). Gibbs et al. (1976) noted the similarity of a Pelican Island, Great Barrier Reef, specimen of B. (B.) latecarinatus with B. (A.) agassizii. The posterior end of this particular specimen, like that of B. (A.) agassizii, is vertically truncated, with the posterior interambulacrum being only slightly carinate aborally (and not prolonged backwards to overhang the periproct and conceal it from dorsal view). Genus METALIA Gray 1855 Synonymy Xanthobrissus Agassiz 1863, p. 28. Prometalia Pomel 1883, p. 34. Eobrissus Bell 1904, p. 236. Metaliopsis Fourtau 1913, p. 68. Type species Spatangus sternalis Lamarck 1816, p. 326, by original designation. Metalia spatagus (Linnaeus 1758) Synonymy Echinus spatagus Linnaeus 1758, p. 665. Metalia spatagus (Linnaeus 1758): H.L. Clark 1925, p. 216; H.L. Clark 1932, p. 219; H.L. Clark 1946, p. 372; Mortensen 1951, p. 540; A.M. Clark and Rowe 1971, p. 166; Gibbs et al. 1976, p. 136; Rowe and Gates 1995, p. 190. Mortensen (1951: 540) lists additional synonymies. Material and locality Two naked tests, ANU 60646-7, from Nosnos village, Boang Island, Tanga Group, New Ireland Province, PNG. Remarks Metalia spatagus (Linnaeus 1758) is widely distributed through the Indo-Pacific (Mortensen 1951; A.M. Clark and Rowe 1971). H.L. Clark (1932) provided the first record of this species from Australasian waters (Low Isles, Great Barrier Reef), recording the largest known specimen, measuring 110 x 93 x 52 mm. By comparison, the largest Tanga specimen measures 54 x 40 x 29 mm. Miskelly (2002) records the species from the Solomon Islands. Proc. Linn. Soc. N.S.W., 125, 2004 138 FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA ACKNOWLEDGMENTS The author is grateful to Alistair Norrie and Richard Joycey, East New Britain Historical and Cultural Centre, for the loan of specimens from the Kokopo Museum, Kokopo, PNG. Prof. Ken Campbell kindly provided his thoughts on the erection of new species Schizaster (Paraster) ovatus sp. nov. and Heliocidaris robertsi sp. nov. and Loisette Marsh, Western Australian Museum, kindly provided an opinion on the identification of the juvenile Protoreaster nodosus from New Britain. Dr. Richard Barwick and Dr. Maité LeGleuher, both of the Department of Geology, Australian National University, kindly photographed all specimens, and provided translation from French of sections from Koehler (1910) and De Ridder (1986), respectively. The comments of Geoff Francis and an anonymous reviewer improved the manuscript. REFERENCES Agassiz, A. (1863). List of the echinoderms sent to different institutions in exchange for other specimens, with annotations. Bulletin of the Museum Comparative Zoology, Harvard 1(2) 17-28. Agassiz, A. (1879). Preliminary report on the Echini of the exploring expedition of H.M.S. Challenger. 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(Papier, Druck and Verlag: Braunschweig). 134 pp. Negretti, B., Philippe, M., Soudet, H.J., Thomassin, B.A. and Oggiano, G. (1990). Echinometra miocenica Loriol, echinide Miocéne, synonyme d’ Echinometra mathaei (Blainville), actuel: Biogéographie et paléoécologie. Geobios 23, 445- 456. Philip, G.M. (1965). The Tertiary echinoids of south- eastern Australia II Stirodonta, Aulodonta, and Camarodonta (1). Proceedings of the Royal Society of Victoria 78(2): 181-196. Read, J.R.L. (1965). Preliminary geological investigation of the lower Warangoi hydro-electric scheme. New Britain August-November 1964. Bureau of Mineral Resources, Geology and Geophysics, Record 1982/15. Rowe, F.W.E. and Gates, J. (1995). Echinodermata. In ‘Zoological Catalogue of Australia’, Vol. 33. 510 pp. (CSIRO Australia: Melbourne). Sladen, W.P. (1889). Report on the scientific results of the voyage of H.M.S. Challenger during the years 1873-76: Zoology - vol. 30, Report on the 135 FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA Asteroidea. 893pp. (H.M. Stationery Office: London). Spencer, W.K. and Wright, C.W. (1966). Asterozoans. In “Treatise on Invertebrate Paleontology, Part U, Echinodermata 3’ (Ed. R.C. Moore) pp. U4-107. (Geological Society of America and University of Kansas Press: Lawrence). Sluiter, C.P. (1895). Die Asteriden Sammlung des Museums zu Amsterdam. Die Echiniden Sammlung des Museums zu Amsterdam. Bijdragen tot de Dierkunde 17, 49-64, 65-74. Struder, T. (1876). Uber Echinodermen aus dem -antarktischen Meere und zwei neue Seeigel von den Papua-Inseln gesammelt auf der Reise S.M.S. ‘Gazelle’ um die Erde. Monatsberichte der Koniglich Preussischen Akademie der Wissenschaften zu Berlin 1876, 452-465. Struder, T. (1880). Ubersicht tiber die wahrend der Reise S.M.S. Corvette ‘Gazelle’ um die Erde 1874-76 gesammelten Echinoiden. Monatsberichte der Koniglich Preussischen Akademie der Wissenschaften zu Berlin 1880, 861-885. Wagner, C.D. and Durham, J.W. (1966). Holectypoids. In ‘Treatise on Invertebrate Paleontology, Part U, Echinodermata 3’ (Ed. R.C. Moore) pp. U440- 450. (Geological Society of America and University of Kansas Press: Lawrence). Wallace, D.A., Johnson, R.W., Chappell, B.W., Arculus, R.J., Perfit, M.R. and Crick, I-H. (1983). Cainozoic volcanism of the Tabar, Lihir, Tanga and Feni Islands, Papua New Guinea: Geology, whole-rock analyses, and rock-forming mineral compositions. Bureau of Mineral Resources, Geology and Geophysics, Report 243. Willey, A. (1902). Zoological results based on material from New Britain, New Guinea, Loyalty Islands and elsewhere, collected during the years 1895 1896 and 1897. Pt. 2. (Cambridge University Press: Cambridge). 253pp. Woods, H. (1958). “Palaeontology Invertebrate’. (Cambridge University Press; Cambridge). 477pp. 136 Proc. Linn. Soc. N.S.W., 125, 2004 I.D. LINDLEY Table 1. Reported starfishes from the Bismarck Archipelago, Papua New Guinea. ASTERIINAE Tarsaster stoichodes Sladen 1889: Fisher 1919, p. 491: north of the Admiralty Group (150 fathoms). ASTERINIDAE Asterina cephus (Miiller and Troschel 1842): A.H. Clark 1954, p. 258: Seleo Island, Aitape district. Patiriella exigua (Lamarck 1816): A.H. Clark 1954, p. 258: Admiralty Group; Seleo Island, Aitape district. ASTEROPSEIDAE Asteropsis carinifera (Lamarck 1816): A.H. Clark 1954, p. 258: Seleo Island, Aitape district. ASTROPECTINIIDAE Astropecten monacanthus Sladen 1883: Bell 1899, p. 136: New Britain. Astropecten polyacanthus Miiller and Troschel 1842: Fisher 1919, p. 64: Admiralty Group. ECHINASTERIDAE Echinaster luzonicus (Gray 1840): Rowe and Gates 1995, p. 59. (= Echinaster eridanella Miller and Troschel 1842, p. 24; Bell 1899, p. 138): New Ireland; New Britain. LUIDUDAE *Luida aspera Sladen 1889: Fisher 1919, p. 171: north of Admiralty Group (150 fathoms). OPHIDIASTERIDAE Linckia laevigata (Linnaeus 1758): Bouillon and Jangoux 1984, p. 249: Laing Island reef, Hansa Bay. Nardoa novaecaledoniae (Perrier 1875): Rowe and Gates 1995, p. 88. (= Nardoa mollis de Loriol, 1891, H.L. Clark 1946, p. 115; A-H. Clark 1954, p. 255): New Britain; Seleo Island, Aitape district. Nardoa tuberculata Gray 1840: Rowe and Gates 1995, p. 88. (= Nardoa finschi de Loriol 1891; Nardoa pauciforis von Martens 1866, H.L. Clark 1946, p. 115): New Britain. Ophidiaster granifer Litken 1871: A.H. Clark 1954, p. 256: Seleo Island, Aitape district. OREASTERIDAE +Anthenea sidneyensis Déderlein 1915: Rowe and Gates 1995, p. 98: Manus Island (Admiralty Group). Culcita novaeguineae Miiller and Troschel 1842: A.H. Clark 1954, p. 254: Seleo Island, Aitape district. Pentaster obtusatus (Bory de St. Vincent 1827). [= Pentaceropsis obtusata (Bory de St. Vincent 1827) Bell 1899, p. 136]: Blanche Bay, New Britain. Protoreaster lincki (de Blainville 1830): Oreaster lincki (= Pentaceros lincki, Bell 1899, p. 136): Blanche Bay, New Britain. Protoreaster nodosus (Linnaeus 1758): H.L. Clark 1946, p. 106; A.H. Clark 1954, p. 254. (= Pentaceros nodosus, Bell 1899: p. 136; Oreaster nodosus H.L. Clark 1908): Blanche Bay, New Britain; Seleo Island, Aitape district. PTERASTERIDAE Hymenaster pullatus Sladen 1889: Fisher 1919, p. 467: southwest of the Admiralty Group (1,070 fathoms). NOTES + the writer follows Spencer and Wright (1966) and Rowe and Gates (1995) in placing Anthenea in Family Oreasteridae. H.L. Clark (1946) and A.M. Clark and Rowe (1971) placed the taxon in Family Goniasteridae. * Denotes type locality in Bismarck Archipelago. Proc. Linn. Soc. N.S.W., 125, 2004 137 FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA Table 2. Reported shallow and deep-water sea-urchins from the Bismarck Archipelago, Papua New Guinea. ARACHNOIDIDAE Arachnoides placenta (Linnaeus 1758): Bell 1899, p. 136; H.L. Clark 1925, p. 154: New Britain. ARBACIIDAE *Pygmaeocidaris prionigera (A. Agassiz 1879): A. Agassiz 1881, pl. XXXIV, figs 14 and 15; H.L. Clark 1925, p. 73 (= Podocidaris prionigera A. Agassiz 1879, p. 199): between New Guinea and Admiralty Group (1,070 fathoms). *Coelopleurus elegans (Bell 1899): H.L. Clark 1925, p. 73. (= Salmacis elegans Bell 1899, p. 135): New Britain. CIDARIDAE Eucidaris metularia (Lamarck 1816): H.L. Clark 1925, p. 20. (= Cidaris metularia de Blainville, 1830, Bell 1899, p. 134): New Britain. Prionocidaris baculosa var. annulifera (Lamarck): Mortensen 1928a, p. 437, 446. (= Schleinitzia crenularis Struder 1876, p. 463; 1880, p. 865): west New Guinea. Stylocidaris reini (Déderlein): H.L. Clark 1925, p. 24; Mortensen 1928a, p. 342, 347, 474 (= Phyllacanthus annulifera Bell 1899, p. 134): New Britain; Milne Bay. DIADEMATIDAE Echinothrix calamaris (Pallas 1774): A.H. Clark 1954, p. 250: Bougainville Island. *Micropyga nigra H.L. Clark 1925: A. Agassiz 1879, p. 200; H.L. Clark 1925, p. 47. (= Astropyga elastica Struder, Bell 1899, p. 135): New Britain. Micropyga tuberculata A. Agassiz 1879, p. 200: A. Agassiz 1881, pl. VII; H.L. Clark 1925, p. 48: Blanche Bay, New Britain. ECHINOMETRIDAE Echinometra mathaei (de Blainville 1825): A.H. Clark 1954, p. 251: Bougainville Island; Seleo Island, Aitape district; Normanby Island. (= Echinometra lucunter Bell 1899, p. 136). ECHINOTHURIIDAE Araeosoma gracile (A. Agassiz 1881): A. Agassiz 1881, p. 89; H.L. Clark 1925, p. 61: Admiralty Group (150 fathoms). LAGANIDAE Laganum decagonale (de Blainville 1827): A. Agassiz 1881; H.L. Clark 1925, p. 156; Mortensen 1948b, p. 332, 336; Lindley 2003a, p. 133: near Admiralty Group (150 fathoms). Laganum depressum var. tonganense (Quoy and Gainard): Mortensen 1948b, p. 324: Admiralty Group. Laganum laganum (Leske): Mortensen (1948b), p. 312: Bismarck Archipelago. SPATANGIDAE Maretia ovata (Leske 1778): A. Agassiz 1881; H.L. Clark 1925, p. 226: Admiralty Group. TEMNOPLEURIDAE Prionechinus agassizii Wood-Mason and Alcock 1891: H.L. Clark 1925, p. 78. (= Echinus elegans, A. Agassiz 1881): near Admiralty Group. *Prionechinus sagittiger A. Agassiz 1879, p. 202: A. Agassiz 1881, pl. IVa, figs 11-14; H.L. Clark 1925, p. 79: between New Guinea and Admiralty Group (1,070 fathoms). Temnopleurus sp., Bell 1899, p. 135: New Britain. Temnopleurus reevesii (Gray 1855): A. Agassiz 1881; H.L. Clark 1925, p. 81: near Admiralty Group (150 fathoms). 138 Proc. Linn. Soc. N.S.W., 125, 2004 I.D. LINDLEY Temnotrema scillae (Mazetti 1894): Mortensen 1904, p. 86; H.L. Clark 1925, p. 91 (= Pleurechinus reticulatus in H.L. Clark 1925, p. 91): New Britain. TOXOPNEUSTIDAE Tripneustes gratilla (Linnaeus 1758): A.H. Clark 1954, p. 250: Bougainville Island; Seleo Island, Aitape district. INVALID RECORDS Astriclypeus manni Verrill, Sluiter 1895, p. 73, New Ireland; Mortensen 1948b, p. 416, 418. Colobocentrotus mertensi Brandt 1835, Sluiter 1895, p. 69, New Ireland; Mortensen 1943b, p. 433. Mellita longifissa Michelin 1858, Sluiter 1895, p. 73, New Ireland; Mortensen 1948b, p. 427, 428. Taxonomic reason: Erroneous labelling (Mortensen 1948b, p. 418; Mortensen 1943b, p. 433; Mortensen 1948b, p. 428, respectively). NOTES * Denotes type localities in Bismarck Archipelago Proc. Linn. Soc. N.S.W., 125, 2004 139) igh re it 7 ‘ sive we — th, +. i if 1 * ¥ 2 . Li / > t ae ic \ ey. ere) ; : ++ a eee: rt — 4, z , eS 7 et 4 toma ‘ = : < ~ Wa - & i _ = P \Vecrr ae - i : - _ : vale se F . “ a Pests sere Re, SRE Ni ppt ay) Woy erin Y ? NPC Seah py + ¥7 yi hn 4 rt | Fe » ee Bre . by mS N ae PP ith - 4 2 » i + ~ © ‘ Weel wy a Poe Oe a deb ja gtleg . | ‘7 : wa ey } ane tiie + RAMANA YS i, * ; 1 : ; ‘4 e yy Cy +) alcatel i ; i AG” \ , Qh igen aa T +4 : oly: why * rs “ ee . oF : 2 3 TOF wie) sei UTD AE ss : Z 2 ; el fa valer st “ Pork. a na . e ~ = af = ~ t : ae 4 ” ? i mene: ‘ ‘ S } . ‘ x = 9 ~ : +. E F = ee : a ) “ So . * A eh ele and cae Hi \ , : ; Late ah ie ’ ‘ ca : } 5 j x ; Z < i 4 xs 5 \ =a ie : BR Sta e th Sy r i ; ; : i t s Y \ ‘ A a 4 4 Beas = <9 TE . 2 a } 3 os t T H ts Soy - - ; : ¥ " we : y hire y ze = ¥ ‘ "e 4 4 & ' . Conodont Faunas from the Mid to Late Ordovician Boundary Interval of the Wahringa Limestone Member (Fairbridge Volcanics), Central New South Wales Y.Y. ZHEN", I.G. PERcIVAL?’ AND B.D. WEBBY? ‘Division of Earth and Environmental Sciences, The Australian Museum, 6 College Street, Sydney, N.S.W. 2010 (yongyi@austmus.gov.au); 7Geological Survey of New South Wales, Department of Mineral Resources, P.O. Box 76, Lidcombe, N.S.W. 2141 (ian.percival @minerals.nsw.gov.au); *Centre for Ecostratigraphy and Palaeobiology, Department of Earth and Planetary Sciences, Macquarie University, N.S.W. 2109, Australia (bwebby @laurel.ocs.mq.edu.au); Honorary Research Associate, Centre for Ecostratigraphy and Palaeobiology, Macquarie University, N.S.W. Zhen, BY: Percival, I.G. and Webby, B.D. (2004). Conodont faunas from the Mid to Late Ordovician boundary interval of the Wahringa Limestone Member (Fairbridge Volcanics), central New South Wales. Proceedings of the Linnean Society of New South Wales 125, 141-164. Twenty-nine conodont species are documented from the Wahringa Limestone Member and other isolated limestone pods of the Fairbridge Volcanics, in the Bakers Swamp area between Wellington and Orange, central New South Wales. Three conodont assemblages are recognised within the Wahringa Limestone Member. The oldest is characterised by the occurrence of Pygodus protoanserinus and Pygodus serra, indicative of a late Darriwilian age (Da3 to early Da4). The overlying assemblage B, bearing Belodina monitorensis, probably ranges across the Mid to Late Ordovician boundary. Assemblage C with abundant Belodina compressa in the upper part of the Wahringa Limestone Member is of late Gisbornian (Gi2) age. The conodont faunas are significant in being the first described from the Lachlan Orogen in New South Wales spanning the Mid to Late Ordovician interval, although resolution of the actual boundary level is limited in the section measured. Manuscript received 17 September 2003, accepted for publication 17 December 2003. KEYWORDS: Conodonts, Fairbridge Volcanics, Late Ordovician (Gisbornian), Mid Ordovician (Darriwilian), Wahringa Limestone Member. INTRODUCTION The Molong Volcanic Belt (MVB) is a meridionally-aligned tectonic feature of Ordovician age within the east Lachlan Orogen in central New South Wales (Glen et al. 1998). Ordovician strata in the northern sector of the MVB near Bakers Swamp between Orange and Wellington (Fig. 1) are represented by the Early Ordovician Mitchell Formation and Hensleigh Siltstone — the latter yielding conodonts of Bendigonian age (upper Prioniodus elegans Zone) from allochthonous limestones (Zhen et al. in press) — and the Fairbridge Volcanics of Mid to early Late Ordovician age. Two autochthonous limestone units occur in the Fairbridge Volcanics. The lower one, known as the Wahringa Limestone Member, is the subject of this paper. Representative conodonts were illustrated earlier in a preliminary report defining this unit, and their age connotations discussed (Percival et al. 1999). Subsequent detailed sampling has yielded many more elements and species, enabling broad confirmation of the original age determination and providing increased precision for the upper age limit of the limestone. The faunas are systematically described here for the first time. They are significant in spanning the Mid to Late Ordovician boundary, an interval which is otherwise poorly represented in shallow water settings of eastern parts of the Lachlan Orogen. STRATIGRAPHY Much of the Fairbridge Volcanics consists of andesitic lava flows, with subsidiary boulder conglomerates (Morgan, Scott & Percival, in Meakin & Morgan 1999). Allochthonous limestones are ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 00 mE /} CATOMBAL x GROUP c WE Bakers ©1474, C1483 ‘@ C1472 C1695-C1700 1471, C1693, C1694 C1429 FO WAHRINGA 145g PEE LIMESTONE MEMBER ©1456 C1450 C1486, C1487, C1488 C1463 4 C ©1463 1468.” A AMC (ARG A A FAIRBRIDGE VOLCANICS ae. At * g'Wohringa’ HENSLEIGH SILTSTONE MITCHELL FORMATION LOCATION MAPS REFERENCE SN z ft 5G Koei a Quaternary alluvium _*2 Dip, and strike “ | PY ? | q (pat sane Ns Undifferentiated Devonian mee Syncline, with plunge , poets \ i Ae sft \ i Undifferentiated Silurian —_t— Anticline 2 es C1458e Fossil locality o 5 S Fairbridge Volcanics mo} ak 2S. allochthonous —s«__ Thrust fault >So limestones YF ) £20 Wahringa Limestone Read NEW SOUTHWALES —/ : Member . Wellington 7 - = Type section Study Area—ia Pa f a - P 1 Orange y eine) § fat Hensleigh Siltstone c > Sam | 5 3 limestones 0 1 2km J | Di a ee ee a ee es Sl) . a) I | Re ed itchell Formation A rik Mitchell Formati 24578-B Figure 1. Locality maps. A. location of Wahringa area, between Wellington and Orange, central New South Wales; B. simplified geological map of the Wahringa area; C. generalised stratigraphic column for this area, showing spot sampled horizons within the Wahringa Limestone Member and the allochthonous limestone blocks within the Fairbridge Volcanics. For further details of the sampled section, see Figure 2. 142 Proc. Linn. Soc. N.S.W., 125, 2004 Y.Y. ZHEN, I-G. PERCIVAL AND B.D. WEBBY uncommon in the lower Fairbridge Volcanics below the Wahringa Limestone Member and conodont yields are disappointingly low. Numerous allochthonous limestone pods emplaced within the Fairbridge Volcanics, stratigraphically overlying the Wahringa Limestone Member, were also processed for conodonts. With the exception of six samples that contained Belodina compressa (indicating an early Late Ordovician or Gisbornian age), these were either barren or yielded only sparse, non-diagnostic elements. Wahringa Limestone Member of Fairbridge Volcanics (Fig. 2) The name derives from the “Wahringa” property, located approximately 28 km south of Wellington on the Mitchell Highway. Here the Wahringa Limestone Member is exposed along strike for approximately 400 m and attains a thickness of 88 m in its type section (situated just north of a bend in Bakers Swamp Creek). Invertebrate macrofossils described or illustrated from the Wahringa Limestone Member comprise brachiopods, gastropods, nautiloids, crinoids, demosponges, stromatoporoids, and a species of tabulate coral (Percival et al. 2001). The unit is subdivisible into three parts: lower beds rich in oncolites, ooids and volcaniclastic detritus, a middle C1687 -Se Se C1683 : to Pe C1678. fd — assemblage C eet as es = LE C1675 m, 8 C1673 s 8 » | s&s 3 | $6§& s % G6 AEN S| bss C1672 —Ra = £s ECS RES EES = 2 sb 2 c1668_E = s | =s2°0 61667 4a 7 |?at oe 3 ce eet 3 C1664 aoe g assemblage B aes eee & s a $2 g |sé & 6. a6 3 2 E cs g g2es"| 8 o&ca o fee e. 4 w” Ss0 8 10m VoLes Sees fey a assemblage A C1652 ss ew eee 0 (C1450) Figure 2. Stratigraphic section measured through the Wahringa Limestone Member showing sampled horizons and ranges of selected, age-significant conodont taxa. Proc. Linn. Soc. N.S.W., 125, 2004 part of muddy, thinly bedded limestones rich in brachiopods, and an upper section that is more massive. The variation in lithologies reflects an increase in water depth from shallow subtidal at the base, to below normal wave base in the middle and upper beds. However, this depth increase does not have a controlling influence on the three distinct conodont assemblages recognised, which represent age- significant rather than biofacies-distinct assemblages. Lithologies in the lower part of the member, which consist mainly of red ooidal grainstones, calcarenites and oncolitic grainstone-packstones, are particularly characteristic of very shallow deposition. Fauna present in these beds (not observed at other levels of the measured section) include the large gastropod Maclurites cf. M. florentinensis, the siphonotretoidean brachiopod Multispinula, and a Calathium-like receptaculitid. Demosponges, including Archaeoscyphia? sp. B, Malongullospongia? sp. and Hindia cf. H. sphaeroidalis, are more widely distributed but are especially common at this level. Fossil grains are subangular to rounded, frequently algal-coated and include dasycladacean and solenoporid algae. Large oncolites with well-preserved cyanobacterial Girvanella filaments and ooids are also abundant. The middle part of the Wahringa Limestone Member is characterised by fine to coarse grained skeletal grainstone (interbedded with silty layers) that is dominated by remains of echinoderms, brachiopods, dasycladacean algae, molluscs, trilobites, and ostracods. Brachiopods, including Sowerbyites?, Leptellina and rare Sowerbyella are concentrated in thin-bedded packstones in the middle part of the unit. Stromatoporoids (Labechia, Labechiella), mostly preserved in growth position, occur slightly above this level and range into the uppermost limestone beds of the unit. : The most common lithologies in the upper part of the Wahringa Limestone Member are fine to coarse grained skeletal, oncolitic grainstone and lesser packstone to wackestone lacking internal lamination. Grains include ostracods, dasycladacean algae, rare ooids, and oncolites with associated Girvanella. CONODONT BIOSTRATIGRAPHY Twenty-nine conodont species based on 897 individual specimens were recovered from 44 samples (Fig. 3), collected from the Wahringa Limestone Member and various limestone pods within the enclosing Fairbridge Volcanics. The faunas range in age from late Darriwilian (Da3, lower Eoplacognathus suecicus Zone) to late Gisbornian (Gi2, Belodina 143 ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. Ist pods below Wahringa Lst a Qa;a alalala ele SIR EIRE a ) alla e| =| = = Rj PEP Eesehas Protopanderodus nogamu \Protopanderodus robustus Figure 3. Distribution chart of conodont species from 44 samples through the Wahringa Limestone Member, and allochthonous limestone blocks in the sole sample Fairbridge Volcanics, from above and below the Wahringa Limestone Member. Column with asterisk depicts occurrence of species in (C1429: middle or upper beds) from southwestern extremity of outcrop of the Wahringa Limestone Member. Wahringa Limestone Member type section Wahringa Lst Mbr Isolated limestone pods in Fairbridge NE of type eae Voics above Wahringa Lst type section Proc. Linn. Soc. N.S.W., 125, 2004 144 Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY compressa Zone). The oldest fauna is represented by a small assemblage, including Appalachignathus delicatulus, Protopanderodus nogamii, ?Periodon aculeatus, Ansella sp., Erraticodon sp., and Stiptognathus sp. A from a single sample C1463, which was obtained from several small limestone clasts within the Fairbridge Volcanics at a stratigraphic level some 120 m below the Wahringa Limestone Member (Percival et al. 1999). This fauna is comparable with that recently described from allochthonous limestones of Da3 age in the Oakdale Formation of the Bell River valley (Zhen and Percival in press), situated approximately 23 km southeast of the “Wahringa” area. Of the six species recognised in sample C1463, four also occur in the Bell River valley fauna. Stiptognathus sp. A is rare, but the other three species (Appalachignathus delicatulus, Protopanderodus nogamii, and Erraticodon sp.) dominate in all five samples from the Oakdale Formation (Zhen and Percival in press). On this basis, the age of sample C1463 can now be revised downwards to the E. suecicus Zone from the previously-interpreted level near the Mid/Late Ordovician boundary (Zhen et al. 2001). In its type section, the Wahringa Limestone Member consists of three laterally continuous outcrops separated by two intervals of poor or negligible exposure (Fig. 2). Initially, one spot sample was collected from each of these three major outcrops, representing the lower, middle, and upper beds of the unit. Subsequent more intensive collecting during section measuring produced 17 samples that yielded conodonts. Although most of these samples have low yields and diversity, three conodont assemblages (herein referred to as A, B, and C from oldest to youngest) can be distinguished. Assemblage A was recovered from sample C1652 at the base of the Wahringa Limestone Member and spot sample C1450 within the lower part of this unit (essentially an equivalent stratigraphic level to C1652) in the type section. The fauna consists of 15 species including Acodus sp., Ansella nevadensis, Ansella biserrata, Belodina sp. B, Dapsilodus variabilis, Drepanoistodus sp., Erraticodon balticus?, Oistodus? sp. cf. venustus, Panderodus gracilis, Periodon aculeatus, Phragmodus flexuosus, Protopanderodus nogamii, Protopanderodus varicostatus, Pygodus serra, and Pygodus protoanserinus. Most of these species are widely distributed and relatively long ranging, but the two species of Pygodus are important biostratigraphically. Pygodus protoanserinus has a range from the upper E. robustus Subzone to the E. lindstroemi Subzone (of the upper Pygodus serra Zone). One specimen (see Fig. 9K, L) referrable to the middle form of the Pa Proc. Linn. Soc. N.S.W., 125, 2004 element of Pygodus serra (Zhang 1998a) was also recovered in the sample C1652. Co-occurrence of P. serra and P. protoanserinus places the base of the Wahringa Limestone Member precisely within the upper E. robustus Subzone (upper Da3 to lowest Da4) of the P. serra Zone. Closest correlations are with successions in China. In the top Guniutan Formation (upper P. serra Zone) of Hunan Province, Zhang (1998b) recorded the co-occurrence of Pygodus protoanserinus with Erraticodon balticus?, Protopanderodus varicostatus, and Periodon aculeatus. Pygodus serra, Periodon aculeatus, Protopanderodus varicostatus, Protopanderodus nogamii, and Panderodus gracilis also occur in the lower part (P. serra Zone) of the Pingliang Formation of the Ordos Basin (An and Zheng 1990). Only five samples from the middle section of the Wahringa Limestone Member have yielded conodonts. These faunas are of very low diversity and productivity, and are referred to herein as Assemblage B. They include Ansella nevadensis, Ansella sp., Belodina monitorensis, Panderodus gracilis, and Periodon aculeatus. Of these, only B. monitorensis is significant for age determination, occurring widely within the late Darriwilian to Gisbornian interval (Sweet in Ziegler 1981). Assemblage B is likely very close to the Mid/Late Ordovician boundary, probably within the Cahabagnathus sweeti Zone, although the precise recognition of the boundary within the middle Wahringa Limestone Member is not determinable on current evidence. The upper part of the type section of the Wahringa Limestone Member is relatively more productive, with 12 samples yielding an assemblage (designated as Assemblage C) of 15 species including Acodus sp., Ansella nevadensis, Belodina compressa, Belodina monitorensis, Besselodus sp., Dapsilodus variabilis, Dapsilodus viruensis, Drepanoistodus sp., Oistodus? sp. cf. venustus, Panderodus gracilis, Periodon aculeatus, Protopanderodus cooperi, Protopanderodus varicostatus, Protopanderodus liripipus and Stiptognathus sp. B. As a zonal index species of late Gisbornian equivalents in the North American Midcontinent zonal scheme, the occurrence of Belodina compressa in the upper part of the Wahringa Limestone Member indicates that top of this unit may be as young as Gi2. This species also occurs in six samples from limestone pods within the Fairbridge Volcanics above the Wahringa Limestone Member. The presence of two elements confidently identified as this species in limestone pods in the Fairbridge Volcanics slightly below the base of the Wahringa Limestone Member (samples C1487 and C1488) cannot be explained at present, as this 145 ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. Figure 4. A-G, Acodus sp.; A, B, P element, MMMC2639, C1450, A, inner lateral view, B, outer lateral view; C, D, P element, MMMC2640, C1652, C, outer lateral view, D, anterior view; E-G, Sa element, MMM(C2641, C1680, lateral views. H-P, Ansella nevadensis (Ethington and Schumacher 1969); H-J, Pa element, MMMC2642, C1450, H, inner lateral view, I, outer lateral view, J, showing surface striation; K, L, Pb element, MMMC2643, C1450, K, outer lateral view, L, inner lateral view; M, N, Sa element, MMMC2644, C1683, lateral views; O, P, Sc element, MMMC2645, C1450, O, outer lateral view, P, inner lateral view. Q, Ansella biserrata Lehnert and Bergstrém in Lehnert et al. 1999; Pa element, MMMC2646, C1652, outer lateral view. R, S, Ansella sp.; R, Pb element, MMMC2647, C1486, inner lateral view; S, Pb element, MMMC2648, C1672, outer lateral view. T, Appalachignathus delicatulus Bergstrom et al. 1974; Pb element, MMMC2649, C1463, inner lateral view. Unless otherwise indicated scale bars are 100 um. 146 Proc. Linn. Soc. N.S.W., 125, 2004 Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY contradicts the species succession in all known global occurrences. LOCALITIES AND SAMPLES Details of the localities and measured section are shown in Figures 1 and 2, and summarised in the Appendix. Distribution of conodont species is presented in Figure 3. All illustrations of conodont elements are presented as SEM photomicrographs (Figs 4-9). Figured specimens bearing the prefix MMMC (“Mining Museum microfossil catalogue’) are deposited in the collections of the Geological Survey of New South Wales, Sydney. Individual samples are referred to by the prefix “C”. Although all species recorded are documented by illustration, only those where adequate material was recovered, or which are of biostratigraphic significance, are described. Unless otherwise mentioned, all specimens are from the Wahringa Limestone Member. SYSTEMATIC PALAEONTOLOGY Class CONODONTATA Pander 1856 Genus ANSELLA Fahraeus and Hunter 1985a Type species Belodella jemtlandica Lofgren 1978. Ansella nevadensis (Ethington and Schumacher 1969) Fig. 4H-P Synonymy . Roundya sp. Sweet and Bergstrom 1962, p. 1244, 1245, text-fig. 5. New Genus A Ethington and Schumacher 1969, p. 478, 479, pl. 68, fig. 12, text-fig. 4J. Oepikodus copenhagenensis Ethington and Schumacher 1969, p. 465, pl.68, figs 5, 9, text-fig. 4L. Oistodus nevadensis Ethington and Schumacher 1969, p. 467, 468, pl. 68, figs 1-4, text-fig. 5C; Tipnis et al. 1978, pl. 6, fig. 7. Belodella nevadensis (Ethington and Schumacher); Bergstrom 1978, pl. 79, figs 9, 10; Bauer 1987, text-fig. 5D. Ansella nevadensis (Ethington and Schumacher); Fahraeus and Hunter 1985a, p. 1175, 1176, pl. 1, figs 7, 10, pl. 2, figs 11a, b, 13a, b, 14, text-fig. 2A-C; Bergstrom 1990, pl. 1, figs 11-14; McCracken 1991, p. 47-49, pl. 3, figs 3, 4, 8, 9, 13, 14, 19-31 (cum syn.); Proc. Linn. Soc. N.S.W., 125, 2004 ?Bauer 1990, pl. 1, fig. 1; 7Bauer 1994, fig. 3.4, 3.5. Material Ten specimens (1 Pa, 4 Pb, 4 Sa, 1 Sc). Description The P elements are characterised by a prominent median costa on each side, and display a sharply inner laterally curved anterior margin. The Pa element has a row of denticles along the posterior edge (Fig. 4H-J). The denticle next to the cusp is the largest, and the others become gradually smaller towards the base. A sharp costa on each side extends from the tip of the cusp and disappears a short distance away from the basal margin. The Pb element has a sharp posterior margin without any denticles, a weaker and broader costa on the inner lateral side, and a sharp, strong costa on the outer lateral side (Fig. 4K, L). The Sa element is symmetrical with a row of closely spaced small denticles along the posterior margin, and bears a sharp antero-lateral costa on each side (Fig. 4M, N). The asymmetrical Sc element has an inner laterally curved anterior margin, and a row of small closely spaced denticles along the posterior margin (Fig. 40, P). Specimens are ornamented with fine striation. Discussion Originally proposed as the form species Oepikodus copenhagenensis Ethington and Schumacher 1969 and New Genus A Ethington and Schumacher 1969 (found in association with Oistodus nevadensis in the Copenhagen Formation of Nevada), these elements were considered as part of the species apparatus of A. nevadensis by Fahraeus and Hunter (1985a), and are herein assigned to the Pa and Sb positions. Fahraeus and Hunter (1985a) also illustrated a symmetrical Sa element (Fahraeus and Hunter 1985a, pl. 2, fig. 14) from the Cobbs Arm Limestone of Newfoundland. Specimens from the Wahringa Limestone Member permit differentiation of denticulate Pa and adenticulate Pb elements. The latter has not been recognised previously, but its assignment to the Pb position is consistent with the apparatus composition of comparable species such as A. jemtlandica and A. crassa Bauer 1994, from central New South Wales (Zhen and Percival in press). Ansella biserrata Lehnert and Bergstrom in Lehnert et al. 1999 Fig. 4Q Synonymy Ansella biserrata Lehnert and Bergstro6m in Lehnert et al. 1999, p. 210, 212, pl. 1, figs 4, 7, pl. 3, figs 1-3, 5 (cum syn.). 147 ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. Material One specimen (Pa). Discussion Lehnert and Bergstr6m in Lehnert et al. (1999) recognised a quadrimembrate apparatus for A. biserrata including biserrate, planoconvex, oistodiform, and triangular elements. We refer the biserrate and planoconvex elements to the Pa and Pb positions respectively, whereas the triangular element is regarded as taking either the Sa (symmetrical) or Sb (asymmetrical) position. The sole specimen from the lower Wahringa Limestone Member (C1652), with smooth lateral faces and fine denticles along its anterior and posterior margins, strongly resembles the holotype (a biserrate element) of A. biserrata from the basal Lindero Formation (Pygodus serra and P. anserinus zones) of west central Argentina (Lehnert et al. 1999). Ansella sp. Fig. 4R, S Synonymy Serraculodus? sp. Zhen and Webby 1995, p. 286, only pl. 5, figs 1, 3, 4. Ansella sp. Zhen et al. 2003a, p. 38, fig. 4A, 4B. Material Three specimens (Pb). Discussion These specimens are similar to the Pb element of A. nevadensis, but they lack the prominent lateral median costae of that species. They are identical with some specimens from the Fossil Hill Limestone of Eastonian age at Cliefden Caves previously assigned to Serraculodus? sp. (Zhen and Webby 1995). They can be distinguished from the Pb elements of both A. jemtlandica (Lofgren 1978) and A. crassa Bauer 1994 in lacking the posteriorly more expanded base displayed in the latter two species (Zhen and Percival in press). Genus BELODINA Ethington 1959 Type species Belodus compressus Branson and Mehl 1933. Belodina compressa (Branson and Mehl 1933) Fig. 5A-I Synonymy Belodus compressus Branson and Mehl 1933, p. 114, pl. 9, figs 15, 16. Belodus grandis Stauffer 1935, p. 603-604, pl. 72, figs 46, 47, 49, 53, 54, 57. 148 Belodus wykoffensis Stauffer 1935, p. 604, pl. 72, figs 51, 52, 55, 58, 59. Oistodus fornicalus Stauffer 1935, p. 610, pl. 75, figs 3-6. Belodina dispansa (Glenister); Schopf 1966, p. 43, plimle ties 7. Belodina compressa (Branson and Mehl); Bergstr6m and Sweet 1966, p. 321-315, pl. 31, figs 12-19; Webers 1966, p. 24, pl. 1, figs 2, 6, 7, 13, 15; Sweet in Ziegler 1981, p. 65-69, Belodina - plate 2, figs 1-4; An et al. 1983, only pl. 25, figs 13, 14; Moskalenko 1983, fig. 3Q-S; Leslie 1997, p. 921-926, figs 2.1-2.20, 3.1-3.4 (cum syn.). Belodina confluens Sweet; Percival et al. 1999, p. 13, fig. 8.21. Material 255 specimens, including eobelodiniform, compressiform, grandiform and dispansiform elements, mainly from the upper part of the Wahringa Limestone Member; some specimens from allochthonous limestones in the Fairbridge Volcanics above the Wahringa Limestone Member, and two elements from limestone pods (C1487, C1488) in the Fairbridge Volcanics which apparently underlie the Wahringa Limestone Member. Discussion Of the known species of Belodina, three including B. compressa, B. confluens Sweet 1979, and B. monitorensis Ethington and Schumacher 1969, are morphologically very similar to each other, reflecting their close phylogenetic relationship. Well- documented successions in the U.S.A. (Sweet 1979) show that the oldest species, B. monitorensis, preceded B. compressa which was succeeded by B. confluens. Sweet (in Ziegler 1981, p. 65) revised all three species as consisting of trimembrate apparatuses, and emphasised that the type species of the genus, B. compressa, was characterised by having a distinct flattening (in lateral view) of the anterior margin near the antero-basal corner. This feature is more prominent in the compressiform element, as shown by the types (Branson and Mehl 1933) and also the specimens figured by Webers (1966); also see Sweet in Ziegler (1981). Both B. confluens and B. compressa are commonly found in association with a more slender, rastrate element bearing smaller denticles. Bergstrom (1990) suggested that these dispansiform elements might represent juveniles of the rastrate elements. Many other workers included these dispansiform elements in a separate species (dispansa) assigned either to Pseudobelodina (Sweet in Ziegler 1981, Nowlan and McCracken in Nowlan et al. 1988, Proc. Linn. Soc. N.S.W., 125, 2004 Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY Figure 5. A-I, Belodina compressa (Branson and Mehl 1933); A, inner lateral view, B, outer lateral view, grandiform element, MMMC2650, C1472; C, dispansiform element, MMMC2651, C1429, inner lateral view; D, compressiform element, MMMC2652, C1472, inner lateral view; E, compressiform element, MMMC2653, C1458, outer lateral view; F, compressiform element, MMMC2654, C1683, outer lateral view; G, dispansiform element, MMMC2655, C1429, outer lateral view; H, inner lateral view, I, outer lateral view, eobelodiniform element, MMMC2656, C1683. J-N, Belodina monitorensis Ethington and Schumacher 1969; J, outer lateral view, K, inner lateral view, grandiform element, MMMC2657, C1687; L, inner lateral view, M, outer lateral view, compressiform element, MMMC2658, C1456; N, eobelodiniform element, MMMC2659, C1456, outer lateral view. O, P, Belodina sp. B; O, eobelodiniform element, MMMC2660, C1450, inner lateral view; P, eobelodiniform element, MMMC2661, C1652, outer lateral view. Q, Belodina sp. A; grandiform element, MMMC2662, C1429, outer lateral view. R-T, Besselodus sp.; R, S, Sa element, MMMC2663, C1676, lateral views; T, M element, MMMC2664, C1675, anterior view. Scale bars are 100 um. Proc. Linn. Soc. N.S.W., 125, 2004 149 ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. McCracken and Nowlan 1989, Trotter and Webby 1995, Leslie 1997, McCracken 2000) or to Belodina (Schopf 1966, Barnes 1977, Nowlan and Barnes 1981, Sansom et al. 1995). In comparing the apparatus architecture of Panderodus, Sansom et al. (1995) suggested that these slender dispansiform elements, which were previously included in Belodina dispansa (Glenister 1957) and Belodina arca Sweet 1979, might belong to the species apparatus of co-occurring B. confluens. Based on twelve well preserved, fused clusters of B. compressa recovered from the Plattin Limestone of Missouri and Iowa, Leslie (1997) demonstrated that B. compressa consisted of a quadrimembrate apparatus including the M (eobelodiniform), Sl (compressiform), S2 (grandiform) and S2 (dispansiform) elements. He further suggested that the dispansiform element — although superficially similar in morphology to Pseudobelodina dispansa — was apparently not conspecific. Leslie also rejected the possibility that such elements represented the juveniles of compressiform and grandiform elements in consideration of the range of sizes and growth series preserved in the dispansiform elements. The material of B. compressa and B. confluens from central New South Wales shows recognisable differences between the two species. Compressiform elements of B. compressa display in lateral view a straight segment of the anterior margin near the antero-basal corner. In comparison, the anterior margin of the compressiform element of B. confluens is regularly curved near the antero-basal corer. Hence specimens of B. compressa previously reported from the Fork Lagoons Beds of central Queensland (Palmieri 1978), and from the Trelawney Beds of the New England Fold Belt (Philip 1966) were subsequently reassigned to B. confluens (Zhen and Webby 1995). Specimens from the Wahringa Limestone Member and various limestone pods within the Fairbridge Volcanics are the first confirmed records of B. compressa from eastern Australia. Belodina monitorensis Ethington and Schumacher 1969 Fig. 5J-N Synonymy Belodina monitorensis monitorensis Ethington and Schumacher 1969, p. 456, pl. 67, figs 3, 5, 8, 9, text-fig. 5D. Belodina monitorensis marginata Ethington and Schumacher 1969, p. 456, pl. 67, figs 1, 2, 4, 6, text-fig. SE. Eobelodina occidentalis Ethington and Schumacher 1969, p. 456, pl. 67, figs 16, 20, text-fig. SH. 150 Belodina monitorensis Ethington and Schumacher 1969, p. 455, 456; Sweet in Ziegler 1981, p. 79-81, Belodina - plate 1, figs 10, 11; Belodina - plate 2, figs 5-7; Bauer 1987, p. 12, pl. 1, figs 10, 13, 14; Bauer 1990, pl. 1, fig. 9; Bauer 1994, fig. 3.16, 3.17, 3.20, BAe Material 17 specimens including eobelodiniform, compressiform and grandiform elements. Discussion Belodina monitorensis was originally defined as having prominent antero-lateral costae on either side of the grandiform element and generally four or five denticles on both grandiform and compressiform elements. A similar antero-lateral costa is also commonly found in the grandiform elements of B. compressa (Fig. 5B; also see Leslie 1997, fig. 2.3), and in the grandiform elements of B. confluens (McCracken 1987, pl. 1, fig. 1; Zhen and Webby 1995, pl. 1, figs 17, 20; Zhen et al. 1999, fig. 5.8). Therefore, this character appears to be unreliable in characterising B. monitorensis. Although B. confluens and B. compressa typically have a greater number of denticles (five to nine), it seems rather arbitrary to split B. monitorensis (typically four or five denticles) from B. confluens based solely on the former having fewer denticles. Though the species status of B. monitorensis is uncertain in our view, stratigraphically it occurs much earlier than typical B. confluens. In the type section of the Wahringa Limestone Member, B. monitorensis occurs lower than B. compressa, but it is also found in association with the latter species in several samples in the upper part of the type section. The Wahringa Limestone Member specimens are comparable with the type material of B. monitorensis in having only three or four denticles, and in having an antero-lateral costa on the furrowed side of the grandiform element (Fig. 5J), also shown by the holotype (Ethington and Schumacher 1969, pl. 67, fig. 5). Therefore, the species is tentatively retained here pending further detailed studies on B. monitorensis and other related species. Belodina sp. A Fig. 5Q Material One specimen from sample C1429 (upper beds of the Wahringa Limestone Member at the southwestern extremity of its outcrop). Discussion This compressiform element has a squat cusp and two Proc. Linn. Soc. N.S.W., 125, 2004 Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY short and stout denticles along the posterior margin. The specimen is not as strongly compressed laterally as other known species of Belodina. Belodina sp. B Fig. 5O, P Material Three eobelodiniform specimens. Discussion From its association with Pygodus protoanserinus in two samples (C1450 and C1652) from the lower part of the Wahringa Limestone Member this species has a late Darriwilian age (upper Da3 to lowest Da4, upper P. serra Zone), making it one of the earliest representatives of the genus Belodina. With a less extended heel it shows some morphological resemblance to the eobelodiniform element of Belodina beiyanhuaensis Qiu in Lin, Qiu and Xu 1984, but no rastrate elements have been recovered to confirm such an assignment. Genus ERRATICODON Dzik 1978 Type species Erraticodon balticus Dzik 1978. Erraticodon balticus? Dzik 1978 Fig. 6P, Q Synonymy Erraticodon balticus Dzik 1978, p. 66, pl. 15, figs 1- 3, 5, 6, text-fig. 6; ?Stouge 1984, p. 84, pl. 17, figs 9-19; Watson 1988, p. 113, pl. 5, figs 2-10, pl. 8, figs 1, 2, 5, 6, 8-13 (cum syn.); Dzik 1991, p. 299, fig..12A; Ding et al. in Wang 1993, p. 176, pl. 37, only figs 19-28, non fig. 18; ?Pohler 1994, pl. 3, figs 3-5; Lehnert 1995, p. 87, pl. 10, figs 13, 16, pl. 12, figs 3-5; ?Zhang 1998b, p. 71, pl. 9, figs 6-13; ?Albanesi in Albanesi et al. 1998, p. 176, pl. 4, figs 16-18; ?Johnston and Barnes 2000, p. 19, pl. 4, figs 18, 20, 23, 24, 29; Zhao et al. 2000, p. 203, pl. 36, figs 1-16; ?Pyle and Barnes 2002, p. 111, pl.20, figs 8, 9. Material One specimen (M). Discussion Dzik (1978) originally defined the species as consisting of a seximembrate apparatus, but later (Dzik 1991) determined a septimembrate apparatus with digyrate Pa and Pb elements as typical of the species (Zhen et al. 2003b). Erraticodon balticus is characterised by Proc. Linn. Soc. N.S.W., 125, 2004 having an accentuated denticle on the posterior process of the Sa, Sb and Sc elements (Dzik 1991, fig. 12A). The specimen from the Wahringa Limestone Member is broadly comparable with the M element of the illustrated type material (Dzik 1978, pl. 15, fig. 5), except that the latter has a reclined cusp; as our specimen has an erect cusp, it is only questionably referred to this species. Specimens ascribed to Erraticodon balticus from the Guniutan Formation of South China (Zhang 1998b), the San Juan Formation of the Precordillera in Argentina (Albanesi in Albanesi et al. 1998), the Ospika Formation of British Columbia (Pyle and Barnes 2001), and the Cow Head Group of western Newfoundland (Johnston and Barnes 2000), all apparently lack the distinctive larger denticle on the posterior process of the S elements, and therefore should only be doubtfully assigned to the species. Erraticodon sp. Fig. 60 Material One specimen (Sa) from sample C1463, a limestone pod in the Fairbridge Volcanics stratigraphically below the Wahringa Limestone Member. Discussion This alate element is identical with the Sa element of a new species of Erraticodon under description from allochthonous limestones within the Oakdale Formation of central New South Wales (Zhen and Percival in press). It has a prominent cusp with a flange-like costa on each side, which extends basally to define the upper margin of the lateral processes. The lateral processes bear four widely spaced, peg- like denticles. Comparison with other species of Erraticodon are discussed elsewhere (Zhen and Percival in press). Genus PERIODON Hadding 1913 Type species Periodon aculeatus Hadding 1913. Periodon aculeatus Hadding 1913 Figs 6R, S, 7A-K Synonymy Periodon aculeatus Hadding 1913, p. 33, pl. 1, fig. 14; Lindstrom 1955b, p. 110, pl. 22, figs 10, 11, 14-16, 35; Lofgren 1978, p. 74, pl. 10, fig. 1; pl. 11, figs 12-26, Fig. 29 (cum syn.); Sweet in Ziegler 1981, p. 237, Periodon - plate 1, figs 1-6; Nowlan 1981, pl. 4, figs 1-9; An 1987, p. 167, pl. 24, figs 7-17; Bergstrom 1990, pl. 1, figs 15, 16, pl. 151 ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. Figure 6. A-D, Dapsilodus variabilis (Webers 1966); A, B, symmetrical distacodontiform element, MMMC2665, C1675, lateral views; C, symmetrical distacodontiform element, MMMC2666, C1450, lateral view; D, acodontiform element, MMMC2667, C1652, outer lateral view. E-J, Dapsilodus viruensis (Fahraeus 1966). E, F, Sa element, MMMC2668, C1675, lateral views; G, outer lateral view, H, inner lateral view, Sb element, MMMC2669, C1675; I, outer lateral view, J, inner lateral view, Sc element, MMMC2670, C1675. K-M, Drepanoistodus sp.; K, outer lateral view, L, inner lateral view, Sc element, MMM(C2671, C1652; M, P element, MMMC2672, C1450, inner lateral view. N, Oistodus? sp. cf. venustus Stauffer 1935; anterior view, MMMC2673, C1450. O, Erraticodon sp.; Sa element, MMMC2674, C1463, postero-lateral view. P, Q, Erraticodon balticus? Dzik 1978; M element, MMM(C2675, C1450, P, posterior view, Q, anterior view. R, S, Periodon aculeatus Hadding 1913; Pb element, MMMC2676, C1450, R, inner lateral view, S, outer lateral view. Scale bars are 100 pm. 152 Proc. Linn. Soc. N.S.W., 125, 2004 Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 2, fig. 15; An and Zheng 1990, pl. 12, figs 12-17; McCracken 1991, p. 50, pl. 1, figs 13, 20, 22, 25-28, pl. 2, figs 24-27, 31, 34, 35 (cum syn.); Pohler 1994, pl. 4, figs 30- 32; Dzik 1994, p. 111, pl. 24, figs 10-13, text-fig. 31b; Lehnert 1995, p. 110, pl. 10; fig. 2, pl. 11, figs 10, 11, pl. 13, figs 9, 11, 12, pl. 16, figs 8, 9, 11-13; Armstrong 1997, p. 774, pl. 2, figs 13-21; text-fig. 3; Albanesi in Albanesi et al. 1998, p. 170, pl. 15, figs 16-17, pl. 16, figs 19, 20 (cum syn.); Zhang 1998b, p. 80, 81, pl. 14, figs 1-8 (cum syn.); Johnston and Barnes 2000, p. 32-35, pl. 13, figs 12, 13, 17, 18, 20-31, pl. 14, figs 1-7, text-figs 4, 5 (cum syn.); Rasmussen 2001, p. 110, pl. 13, figs 8-11 (cum syn.); Pyle and Barnes 2002, p. 107, pl. 21, figs 7-9. Material 97 specimens. Description Both Pa and Pb elements are angulate with a prominent cusp which is laterally compressed with a median costa on each side. The Pb element differs fram the Pa element in having a twisted posterior process and a strongly inner laterally curved and downwardly extended anterior process (Fig. 6R, S). The M element is makellate with an adenticulate outer lateral process, and with 4-6 closely spaced denticles along the inner lateral margin. The alate Sa element has a long posterior process bearing closely spaced denticles. The sixth denticle away from the cusp is much larger and more robust (Fig. 7E, F). The lateral process on each side is blade-like, bearing small closely spaced rudimentary denticles along the edge. The basal cavity is Shallow with a recessive basal margin zone. The Sb element is tertiopedate and asymmetrical, and bears a long denticulate posterior process with closely spaced, strongly reclined denticles, a long inner lateral process with more than seven small confluent denticles, and a short outer lateral process with only two small denticles (Fig. 7H, G). The Sc element is bipennate with a long, denticulate posterior process, bearing closely spaced, strongly reclined denticles, and with an inner laterally curved anterior process bearing small confluent denticles (Fig. 7I-K). Discussion Following the Treatise definition of the genus (Clark et al. 1981, p. W128), Sweet (1988) proposed a seximembrate apparatus for P. aculeatus, consisting of angulate Pa and Pb, makellate M, and ramiform Proc. Linn. Soc. N.S.W., 125, 2004 alate Sa, tertiopedate Sb and bipennate Sc, elements. Albanesi (in Albanesi et al. 1998, text-fig 31, pl. 9, fig. 10) suggested a septimembrate apparatus for the species by recognizing a lozognathiform Sd element, which bears a long denticulate, twisted posterior process, a short, denticulate outer lateral process and an inner laterally curved, sharp, blade-like anterior costa. Rasmussen (2001, pl.13, fig.11) also recognised a modified tertiopedate Sd element, and described it as characterised by a multidenticulate, twisted, posterior process and weakly denticulated anterior process, and process-like extension of the outer-lateral costa or carina, but only a poorly preserved specimen was illustrated. In the Wahringa collections no Sd elements have been recognised. Lofgren (1978, p. 75) suggested that the number of small denticles between the cusp and the biggest denticle increases from a mean of 4.7 to 5.6 in successively younger samples. Specimens from the Wahringa Limestone Member may therefore represent more advanced forms of the species within its stratigraphic range, as shown by the M elements, which are strongly geniculate with a sinuous basal margin and bear 4-6 denticles (mean 5.5) along the inner lateral margin. Genus PHRAGMODUS Branson and Mehl 1933 Type species Phragmodus primus Branson and Mehl 1933. Phragmodus flexuosus Moskalenko 1973 Fig. 7L, M Synonymy Phragmodus sp. Moskalenko 1972, p. 48-50, text- fig. 1, table 2. Phragmodus flexuosus Moskalenko 1973, p. 73, 74, pl. 11, figs 4-6; Sweet in Ziegler 1981, p. 255-258, Phragmodus - plate 2, figs 1-6 (cum syn.); Bauer 1987, p. 24, 25, pl. 3, figs 10, 14, 15, 17, 18, 20, 24, text-fig. 8; Ethington and Clark 1982, p. 79-82, pl. 9, figs 2-7 (cum syn.); Bauer 1994, p. 367, 368, fig. 5.25, 5.26, 5.28, 5.30-5.33; Percival et al. 1999, fig. 8.15. Material One specimen (Sa). Discussion This specimen is alate with a suberect cusp, a long denticulated posterior process, and with a flange-like costa on each lateral side. The straight posterior process supports more than seven widely spaced denticles, ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. Figure 7. A-K, Periodon aculeatus Hadding 1913; A, outer lateral view, B, inner lateral view, Pa element, MMM(C2677, C1450; C, posterior view, D, anterior view, M element, MMMC2678, C1450; E, F, Sa element, MMMC2679, C1450, lateral views; G, outer lateral view, H, inner lateral view, Sb element, MMMC2680, C1450; I, outer lateral view, J, inner lateral view, Sc element, MMMC2681, C1450; K, Sc element, MMMC2682, C1652, outer lateral view. L, M, Phragmodus flexuosus Moskalenko 1973; Sa element, MMMC2683, C1450, lateral views. N-U, Panderodus gracilis (Branson and Mehl 1933); N, posterior view, O, inner lateral view, P, outer lateral view, graciliform element, MMMC2684, C1450; Q, posterior view, R, basal view, S, lateral view, aequaliform element, MMMC2685, C1458; T, falciform element, MMMC2686, C1697, outer lateral view; U, falciform element, MMMC2687, C1458, inner lateral view. Scale bars are 100 um. which are reclined, similar in size, with V- or U-shaped _ or less equal-sized denticles, and a few small, spaces between. It is comparable to the type material rudimentary denticles on the lower edge of the lateral from Siberia, except that the Wahringa specimen __ processes. Although Moskalenko (1972) initially exhibits a rather straight posterior process with more recognised nine morphotypes for the species, she later 154 Proc. Linn. Soc. N.S.W., 125, 2004 Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY (Moskalenko 1973) formally described the species as a form species. Based on a detailed revision, Ethington and Clark (1982) redefined P. flexuosus as consisting of a seximembrate apparatus. However, the illustrated S elements from Utah display more pronounced undulations and twisting of the posterior process and size variation of the denticles on the posterior process (Ethington and Clark 1982, pl. 9, figs 3, 6) than the type material from Siberia (Moskalenko 1973). Genus PROTOPANDERODUS Lindstr6m 1971 Type species Acontiodus rectus Lindstr6m 1955a. Protopanderodus cooperi (Sweet and Bergstr6m 1962) Fig. 8A-E Synonymy Scandodus rectus Lindstr6m 1955a, p. 593, only pl. 4, figs 22, 23. ~Acontiodus cooperi Sweet and Bergstrom 1962, p. 1221, pl. 168, figs 2, 3, text-fig. 1G. Scandodus sp. Sweet and Bergstr6m 1962, p. 1246, pl. 168, figs 13, 16. Protopanderodus cooperi (Sweet and Bergstrém); Zhang 1998b, p. 81, 82, pl. 14, figs 13-17 (cum syn.). Material Seven specimens (6 Sa, 1 Sb). Discussion This species is rare in the Wahringa collections. Two morphotypes are recognised as representing the Sa and Sb elements, all with sharp anterior and posterior margins, and a suberect cusp and one costa on each lateral side. The Sa element is symmetrical with a sharp median costa (Fig. 8A, B). The Sb element resembles the Sa, but is slightly asymmetrical with a more strongly developed costa on the inner side (Fig. 8C- E). No scandodiform P elements and no laterally compressed Sc elements, as characterising P. cooperi of previous authors, were recovered. Based on the original definition of the species given by Sweet and Bergstrom (1962) and more recent revision (Zhang 1998b), elements of P. cooperi exhibit sharp anterior and posterior margins, a well developed anticusp-like extension at the antero-basal corner, deep antero-lateral recesses in the basal margin, and no more than one costa on each lateral face. Protopanderodus cooperi can be differentiated from P. rectus (Lindstr6m) in having an anticusp-like extension at the anterobasal - cormer,and from P. varicostatus in displaying no more Proc. Linn. Soc. N.S.W., 125, 2004 than one costa on each lateral side. Zhang (1998b) provided a comprehensive synonymy list, and illustrated what she recognised as P, M, Sa and other undifferentiated S elements; however, Zhang provided neither diagnosis nor descriptions of the constituent elements of the species apparatus of P. cooperi. This species was originally proposed as a form species from the Ferry Formation of Alabama. The holotype (Sweet and Bergstrom 1962, pl. 168, figs 2, 3) is slightly asymmetrical, defined here as taking the Sb position. Zhang (1998b) included the form species Scandodus sp. Sweet and Bergstr6m 1962 in the P position of P. cooperi. Based on their illustrations and brief discussion, the P element is inferred to be a scandodiform element with broad costa on the inner lateral face and with a smooth outer lateral face. Zhang (1998b) also included the holotype of Scandodus rectus Lindstrom 1955a as occupying the M position in P. cooperi. This scandodiform element is similar to the P element previously defined, except that only a broad carina is developed on the inner lateral face. For consistency, these two scandodiform elements are tentatively taken to represent the Pa and Pb positions in Protopanderodus. The symmetrical Sa of P. cooperi was illustrated from the Guniutan Formation of South China (Zhang 1998b, pl. 14, fig. 13), and was also recovered from the Wahringa samples (Fig. 8A, B). Protopanderodus robustus (Hadding 1913) Fig. 8J-M Synonymy Drepanodus robustus Hadding 1913, p. 31, pl. 1, fig.5. Protopanderodus robustus (Hadding); Lofgren 1978, p. 94, 95, pl. 3, figs 32-35, text-fig. 31G-J (cum syn.); An 1987, p. 173, pl. 11, figs 7-10 (cum syn.); McCracken 1989, p. 20-22, pl. 1, figs 1-10, text-fig. 3E (cum syn.); Albanesi in Albanesi et al. 1998, p. 129, 130, pl. 11, figs 17-20, text-fig. 14A (cum syn.). Material Two specimens (Sa, Sc). Discussion One specimen in the Wahringa collection (Fig. 8L, M) which has sharp anterior and posterior margins and is laterally compressed with a postero-lateral costa on each side, is regarded as representing the Sa element of Protopanderodus robustus. The other specimen with a single costa on the outer lateral face is referred to the 155 ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. Figure 8. A-E, Protopanderodus cooperi (Sweet and Bergstrém 1962); A, B, Sa element, MMMC2688, C1683; lateral views; C, upper view, D, outer lateral view, E, inner lateral view, Sb element, MMMC2689, C1682. F-I, Protopanderodus nogamii (Lee 1975); F-H, Sa element, MMMC2690, C1450, lateral views; I, Sa element, MMMC2691, C1463, lateral view. J-M, Protopanderodus robustus (Hadding 1913); J, outer lateral view, K, inner lateral view, Sc element, MMMC2692, C1680; L, M, Sa element, MMMC2693, C1458, lateral views. N-X, Protopanderodus varicostatus (Sweet and Bergstrém 1962); N, Pa element, MMMC2694, C1652, inner lateral view; O, outer lateral view, P, inner lateral view, Pb element, MMMC2695, C1675; Q, R, Sd element, MMMC2696, C1675, lateral views; S, T, Sd element, MMMC2697, C1682, lateral views; U, outer lateral view, V, inner lateral view, Sb element, MMMC2698, C1675; W, inner lateral view, X, outer lateral view, Sc? element, MMMC2699, C1675. Y, Z, Protopanderodus liripipus Kennedy et al. 1979; Sa element, MMMC2700, C1458, lateral views. Scale bars are 100 um. 156 Proc. Linn. Soc. N.S.W., 125, 2004 Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY Sc element of the same species (Fig. 8J, K). Although Lindstr6m (1971), and Johnston and Barnes (2000) maintained the original generic assignment of the form species Drepanodus robustus Hadding 1913, most other authors followed the revision of Lofgren (1978) who regarded it as a species of Protopanderodus. L6éfgren (1978) recognised three morphotypes: symmetrical acontiodiform (an Sa element) with a postero-lateral costa on each side (Fig. 8L, M), asymmetrical acontiodiform (an Sc element) with a strong costa on the outer lateral face and a non costate inner lateral face (Fig. 8J, K), and scandodiform (interpreted here as a P element). Based on this multielement species definition, P. robustus is morphologically very close to P. cooperi. Specimens referable to D. robustus were also recorded from the Pratt Ferry Formation of Alabama, where the type material of P. cooperi was described (Sweet and Bergstrom 1962). Zhang (1998b) suggested that the holotype of D. robustus might be an element of an uncertain species of Protopanderodus, but she included all the material described from Sweden by Lofgren (1978) as P. robustus (Hadding) in her synonymy of P. cooperi. This implies that P. cooperi may be a junior synonym of P. robustus. Given that the base of the holotype of Drepanodus robustus is apparently broken (see also Lindstr6m 1955b), it remains difficult to separate these two species. Protopanderodus varicostatus (Sweet and Bergstrom 1962) Fig. 8N-X Synonymy Scolopodus varicostatus Sweet and Bergstrom 1962, p. 1247, pl. 168, figs 4-9, text-fig. 1A, C, K. Scandodus unistriatus Sweet and Bergstr6m 1962, p. 1245, pl. 168, fig. 12, text-fig. 1E. Protopanderodus varicostatus (Sweet and Bergstrom); Dzik 1976, only text-fig. 16f, g; Fahraeus and Hunter 1985b, p. 183, text- fig. 2; Bauer 1987, p. 27, pl. 3, figs 19, 21- 23; An 1987, p. 173, pl. 11, figs 2, 3; Dzik 1994, p. 74, pl. 14, figs 1-5, text-fig. 11b; Zhang 1998b, p. 83, 84, pl. 15, figs 14-19 (cum syn.). Material Seven specimens from sample C1675, and one specimen from C1652. Discussion Sweet and Bergstrom (1962) originally recognised three form-groups for the species, namely symmetrical, slightly asymmetrical, and markedly asymmetrical. Proc. Linn. Soc. N.S.W., 125, 2004 Fahraeus and Hunter (1985b) proposed a quinquimembrate apparatus for this species, with’ elements referred to as groups A to E. Group A is a symmetrical multi-costate element with two costae on each lateral face. Group B is an asymmetrical tri- costate element (= the markedly asymmetrical form group of Sweet and Bergstr6m 1962) with two costae on the inner lateral face and a postero-lateral costa on the outer lateral face. Group C is an asymmetrical multi-costate element (= slightly asymmetrical form group of Sweet and Bergstrom 1962) with a twisted cusp and two costae on each side. Group D is a tri- costate element, similar to group B but less laterally compressed with costa on the outer lateral face situated more towards the middle. Group E is a scandodiform element represented by the form species Scandodus unistriatus Sweet and Bergstrom 1962 (here assigned to the Pb position). Zhang (1998b) illustrated one of the multi- costate specimens as the P element, and two morphologically different scandodiform specimens as the M element. One of the latter specimens (Zhang 1998b, pl. 15, fig. 19) is comparable with the form species S. unistriatus Sweet and Bergstrom 1962, and is regarded here as representing the Pb element of P. varicostatus (Fig. 8O, P). The other specimen illustrated as the M element (Zhang 1998b, pl. 15, fig. 14), which was recovered from the same sample with other illustrated specimens of P. varicostatus, has a multi-costate inner lateral face with three costae bordering two grooves and a few, shorter and weaker secondary costae near the base. It is designated here as occupying the Pa position of the species. Similar specimens (arcuatiform) referrable to the Pa element of P. varicostatus were also reported from allochthonous limestone clasts within the Shinnel Formation of Scotland (Armstrong 1997, pl. 3, figs 3, 4). Morphologically it resembles the Pa element of Protopanderodus cf. calceatus Bagnoli and Stouge 1996, recovered from the allochthonous limestones in the Oakdale Formation of central New South Wales (Zhen and Percival in press, pl. 17, figs A, C). However, this latter element has one larger, open groove on the inner lateral face, while the Pa element of P. varicostatus from South China (Zhang 1998b, pl. 15, fig. 14) and from the Wahringa area (Fig. 8N) has two equally developed, narrower grooves with a sharp costa between them. Protopanderodus liripipus Kennedy et al. 1979 is also multi-costate, but with a more posteriorly extended base (Fig. 8Y, Z). Three morphotypes of multicostate (S) elements with two costae on each side are recognised from sample C1675 and possibly should be assigned to the Sd, Sb and Sc? positions, as no tri-costate elements have been recovered. The Sd element is 157 ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. symmetrical with a reclined cusp and a short base. The Sb element is weakly asymmetrical with suberect cusp and a longer base. The Sc? element is asymmetrical and laterally strongly compressed with a suberect cusp and a short base. Genus PYGODUS Lamont and Lindstr6m 1957 Type species Pygodus anserinus Lamont and Lindstrom 1957. Pygodus protoanserinus Zhang 1998a Fig. 9B-J Synonymy Pygodus anserinus Lamont and Lindstr6m 1957, p. 68, only fig. 1d. Pygodus serrus (Hadding); Bergstr6m 1971, p. 149, pl. 2, figs 22, 23; An 1981, pl. 4, figs 1-3; An 1987, pl. 24, fig. 25, pl. 26, figs 3, 6, 13, pl. 29, figs 2, 3; Nicoll 1980, fig. 3H-L; Ding et al. in Wang, 1993, p. 198, pl. 30, figs 10, 13, 15-18, 20-22, 24, pl. 35, 24, 26. Pygodus protoanserinus Zhang 1998a, p. 96, Fig. 2D, pl. 3, figs 9-18 (cum syn.); Zhang 1998b, p. 86, 87, pl. 16, figs 6-8 (cum syn.). Pygodus serra (Hadding); Percival et al. 1999, fig. 8.18; Pickett and Percival 2001, fig. 4C. Material Four Pa (stelliscaphate), five Pb (pastinate), and one Sb (tertiopedate) elements. Discussion Five species were assigned to Pygodus in the recent study of the genus by Zhang (1998a). They have short stratigraphic ranges and hence are very useful biostratigraphic index fossils. Sweet and Bergstrom (1962) and Bergstr6m (1971) initially suggested that the apparatus of Pygodus anserinus, the type species of the genus, included elements represented by the form species Pygodus anserinus Lamont and Lindstr6m 1957, and Haddingodus serrus (Hadding). Bergstrom (1971) also raised the possibility that the Pygodus apparatus might include elements represented by the form species Tetraprioniodus lindstroemi Sweet and Bergstrom 1962 and Roundya pyramidalis Sweet and Bergstrom 1962. This quadrimembrate Pygodus apparatus composition has been widely accepted (Léfgren 1978, Clark et al. 1981, Sweet 1988). Subsequently, Armstrong (1997) has implied a septimembrate apparatus for Pygodus, but with only confirmed elements occupying the Pa, Pb, Pc, M and Sc positions. By including two pygodiform elements in the apparatus, Armstrong (1997) suggested that the 158 P. anserinus apparatus consisted of the stelliscaphate Pa, pastiniscaphate Pb, bipennate Pc (= pastinate Pb of other authors’ usage - see Zhang 1998a, b), tertiopedate M (termed an S element by other authors - see Zhang 1998a, b, and herein), and the ramiform Sc element. More recently Zhang (1998a, 1998b) proposed a quinquimembrate apparatus for Pygodus, including stelliscaphate Pa, pastinate Pb, and three ramiform S elements (alate, tertiopedate and quadriramate). Distinctions between P. serra and P. protoanserinus were discussed in detail by Zhang (1998a). Pygodus protoanserinus ranges from the E. robustus Subzone to the E. lindstroemi Subzone of the upper serra Zone in Baltoscandia, Scotland, North America, China, and Australia. The stelliscaphate Pa element from the lower part of the Wahringa Limestone Member is identical with the type material of P. protoanserinus, being characterised by having the middle denticle row situated more towards the outer denticle row on the upper surface. Specimens illustrated by Nicoll (1980) as P. serrus from the Pittman Formation at Black Mountain, Canberra, ACT, are here reassigned to P. protoanserinus on this same basis. A single specimen from the lower part of the Wahringa Limestone Member has the middle row of the denticles positioned centrally, and is therefore referred to P. serra (Fig. 9K, L), being more comparable with the middle form of the Pa element of that species as defined by Zhang (1998a). Genus STIPTOGNATHUS Ethington, Lehnert, and Repetski 2000 Type species Reutterodus borealis Repetski 1982. Stiptognathus sp. A Fig. 90, P Synonymy Stiptognathus sp. Zhen and Percival in press, fig. 21L-O. Material Two specimens from sample C1463 from an allochthonous limestone within the Fairbridge Volcanics, stratigraphically below the Wahringa Limestone Member. Discussion The symmetrical Sa and geniculate M elements recovered from sample C1463 are identical with those from the allochthonous limestones of the Oakdale Formation (Zhen and Percival in press). Denticles on these specimens are small, closely spaced and blunt. Proc. Linn. Soc. N.S.W., 125, 2004 Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY Figure 9. A, Pseudooneotodus mitratus (Moskalenko 1973); upper view, MMMC2701, C1474. B-J, Pygodus protoanserinus Zhang 1998a; B, Pa element, upper view, MMMC2702, C1450; C, Pa element, upper view, MMMC2703, C1652; D, upper view, and EF, enlargement showing surface structure, Pa element, MMMC2704, C1652; F, inner lateral view, G, outer lateral view, H, anterior view, Pb element, MMMC2705, C1652; I, outer lateral view, J, showing surface structure, Sb element, MMMC2706, C1652. K, L, Pygodus serra (Hadding 1913); K, upper view, L, basal view, Pa element, MMMC2707, C1652. M, N, Stiptognathus sp. B; M, antero-lateral view, N, posterior view, Sa element, MMMC2708, C1675. O, P, Stiptognathus sp. A; O, M element, MMMC2709, C1463, posterior view; P, Sa element MMMC2710, C1463, antero-lateral view. Unless otherwise indicated scale bars are 100 um. Proc. Linn. Soc. N.S.W., 125, 2004 ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. Stiptognathus sp. B Fig. 9M, N Material Two specimens (Sa) from sample C1675. Discussion The cusp of this symmetrical element is triangular in cross section with a gently curved, wide anterior face, posterior costa, and an antero-lateral costa on each side. Three sharp costae extend basally into three blade-like processes, which bear small, upward-pointed denticles along the edges. They are easily distinguishable from the blunt denticles of Stiptognathus sp. A. ACKNOWLEDGMENTS This study was supported by a Science Fellowship provided by the Sydney Grammar School to Zhen. Initial field collecting was undertaken with the support of the Australian Research Council during 1996 to 1999 (grant A39600788 to B.D. Webby). Gary Dargan from the Geological Survey of N.S.W. assisted in acid leaching, separation and other laboratory work. A grant to Y.Y. Zhen from the Betty Mayne Scientific Research Fund of the Linnean Society of New South Wales defrayed costs of some of the SEM work. The scanning electron microscope illustrations were prepared in the Electron Microscope Unit of the Division of Life and Environmental Sciences, Macquarie University and in the Electron Microscope Unit of the Australian Museum. I.G. Percival publishes with the permission of the Director General, New South Wales Department of Mineral Resources. REFERENCES Albanesi, G.L., Hiinicken, M.A. and Barnes, C.R. 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APPENDIX LOCALITY DATA All grid references are AMG66 co-ordinates and relate to the Cumnock 8632-S 1:50,000 topographic sheet (first ed., 1978). Allochthonous limestones stratigraphically below Wahringa Limestone Member C1463: GR 679150 mE 6371900 mN C1486: GR 678750 mE 6371800 mN C1487: GR 678720 mE 6371800 mN C1488: GR 678700 mE 6371800 mN Wahringa Limestone Member, type section C1450, C1652 (basal beds): GR 678650 mE 6372000 mN C1456, C1664, C1667-C1668, C1672 (middle beds): GR 678700 mE 6372050 mN C1458, C1464, C1673-1683, C1687 (upper beds): GR 678700 mE 6372100 mN Wahringa Limestone Member, northeast extremity of outcrop C1707, C1709-1713: centred on GR 679330 mE 6372620 mN Wahringa Limestone Member, southwest extremity of outcrop (middle or upper beds) C1429: GR 677570 mE 6370800 mN Allochthonous limestones stratigraphically above Wahringa Limestone Member C1693, C1694: GR 678000 mE 6371310 mN C1695, C1696: GR 678100 mE 6372000 mN C1697, C1698: GR 678050 mE 6372000 mN C1699: GR 678025 mE 6372000 mN C1700: GR 678010 mE 6372000 mN C1471: GR 678500 mE 6372200 mN C1472: GR 679250 mE 6373550 mN C1474: GR 678100 mE 6372600 mN C1483: GR 678500 mE 6372800 mN 164 Proc. Linn. Soc. N.S.W., 125, 2004 Wenlock (Early Silurian) Brachiopods from the Orange District of New South Wales A.J. Wricut! and D.L. Strusz? ‘School of Earth and Environmental Sciences, University of Wollongong, Wollongong NSW 2522, tony_wright@uow.edu.au; *Department of Earth and Marine Sciences, Australian National University, Canberra ACT 0200; dstrusz@ems.anu.edu.au Wright, A.J. and Strusz, D.L. (2003). Wenlock (Early Silurian) brachiopods from the Orange District of New South Wales. Proceedings of the Linnean Society of New South Wales 125, 165-172. Two late Wenlock (Early Silurian) brachiopod species from the Ulah Formation near Orange, New South Wales, are closely associated with graptolite faunas. Visbyella cumnockensis occurs in the testis Biozone on Wallace Creek in the Four Mile Creek area, and Strophochonetes melbournensis is recorded from the Judensis Biozone on Spring Creek. Poorly preserved but similar Visbyella? and Strophochonetes? From the Pridoli Wallace Shale at Cheesemans Creek are also illustrated. These occurrences provide significant new stratigraphic and distributional data for the species. Manuscript received 19 March 2003, accepted for publication 22 October 2003. KEYWORDS: Brachiopods, New South Wales, Pgidoli, Silurian, Strophochonetes melbournensis, Ulah Formation, Visbyella cumnockensis, Wallace Shale, Wenlock. INTRODUCTION The Silurian strata of the area west and southwest of Orange, NSW, in the valleys of Spring Creek and Four Mile Creek (Fig. 1), have yielded a diversity of fossils, but very few shelly fossils have ever been described, apart from corals described by authors including Etheridge and McLean (full references to these works can be found in Pickett 1982). The most abundant and important fossils in the region are graptolites, which have been known for more than 50 years and were reported by Packham and Stevens (1955) and Jenkins (1978, 1986). Jenkins recorded (but did not describe) brachiopod faunas from limestones in the Four Mile Creek area, but few brachiopods have been reported from clastic strata common in the area. Rickards and Wright (1997) described two brachiopod species from late Wenlock strata (Judensis Biozone) in Cobblers Creek (Fig. 1), and in the section at “Mirrabooka Park’ brachiopods were noted in Wenlock strata during field work by L. Muir, R.B. Rickards, G.H. Packham and A.J. Wright. A diverse and abundant shelly fauna occurring with the late Wenlock graptolite Testograptus testis on the Cadia gold mine access road, several kilometres to the east of Four Mile Creek, was illustrated by Rickards et al. (2001). The two species described here are recorded for the first time from the region near Orange. One, Visbyella cumnockensis Walmsley et al., 1968, was originally described from near Cumnock, 55 km northwest of Orange, where it occurs with T. testis (Walmsley et al. 1968:315). Visbyella has been reported also, but not illustrated, by Pickett (1982) and Pogson and Watkins (1998). The other species, Strophochonetes melbournensis (Chapman 1903), was previously known only from Wenlock and Ludlow strata in the Melbourne Trough, Victoria. Pickett’s report was based on the record of Visbyella cf. cumnockensis by Sherwin (1971). Sherwin’s locality is younger, and contains a sparse and poorly preserved fauna including also a chonetoide similar to Strophochonetes? savagei Strusz, 2000 from Cumnock. These taxa are illustrated but not described. Documented brachiopod occurrences in the Orange region are still insufficient, however, to permit any notion of a regional brachiopod zonation. EARLY SILURIAN BRACHIOPODS FROM ORANGE NSW to Wellington Borenore a Mirrabooka Park’ ia Spring Creek - penne Be Mt. Canobolas metas, Creek area A Cargo_ 148° 45° Figure 1. Map of the area southwest to west of Orange, central ORANGE Wenlock to Pridoli; the age of the strata at this locality is late Wenlock. w940. The somewhat more abundant specimens of Strophochonetes melbournensis were collected from dark siltstones of the Ulah Formation on the southern side of Spring Creek at ‘Mirrabooka Park’, directly opposite One Tree Hill. There are also occasional poorly preserved brachiopods, including pentamerides, in beds at about the same level on One Tree Hill itself. The shells at W940 occur with a graptolite fauna that includes Monograptus ludensis (R.B. Rickards, pers. comm.). Only disarticulated valves are known at this locality; small phosphatic brachiopods are quite common, and there are rare specimens of other brachiopods including strophomenides and atrypides. Most specimens of Strophochonetes melbournensis at this locality retain shelly material and the spines on the pedicle valve hinge line are often preserved. The environment was most probably a low-energy one. to Bathurst New South Wales, showing the geographic context of the two localities, LM3 on Wallace Creek east of Cargo and W940 near ‘Mirrabooka Park’ east-southeast of Cudal. Inset: the location of Orange within Australia. LOCALITIES LMS. Visbyella cumnockensis was collected from Wallace Creek in the Four Mile Creek area, in grey- brown siltstones assigned by Jenkins (1978) to the Wenlock-Ludlow Ulah Formation. These beds have also yielded the graptolites Cyrtograptus and a new species of Monograptus (L. Muir, pers. comm.), and overlie beds containing T. testis. The brachiopod specimens are moulds of a single pedicle and a single brachial valve on the same bedding surface, which could represent the disarticulated valves of a single shell. No other macrofossils have been found at this locality. In contrast, the type material of V. cumnockensis is entirely of specimens in the ‘butterfly’ position, with the shell opened so that the conjoined valves lie on the bedding surface. The age assigned to the Ulah Formation by Chapman et al. (2003) is late 166 MO/1/27. A few poorly preserved orthide and chonetoide specimens have been collected from this outcrop of fine thin- bedded siltstone low in the Wallace Shale, about 600 m east of ‘Mirrabooka’ homestead. The fauna also includes occasional trilobites. The lo- cality lies within the Monograptus transgrediens Biozone. SYSTEMATIC PALAEONTOLOGY Suprageneric taxonomy follows that in Kaesler (2000); references to authorship of suprageneric taxa are therefore not repeated here. Specific diagnoses have been rephrased to accord with currently accepted terminology (Kaesler 1997). Details of localities are given in the descriptive section below. Abbreviations. Ls - shell length Ld - dorsal valve length. Ws - shell width Wh - hinge line width Proc. Linn. Soc. N.S.W., 125, 2004 A.J. WRIGHT AND D.J. STRUSZ Figure 2. a-g, Visbyella cumnockensis Walmsley et al., 1968. a-c, ventral valve counterparts; a, latex cast of exterior, AM F124331. b-c, internal mould and latex cast, AM F124332. d-g, dorsal valve counterparts; d, latex cast of exterior, AM F124333. e-g, internal mould and latex cast (in ventral and postero-ventral views), AM F124334. h-k, cf. Visbyella cumnockensis, Pridoli, Wallace Shale. h, latex cast of ventral valve, MM F37431. i, latex cast of shell in ‘butterfly’ position, MM F21132. j, external mould of dorsal valve plus internal mould of ventral valve, MM F21125. k, latex cast of incomplete interior of shell in ‘butterfly’ position, MM F37428. Scale bar 2 mm. AM - Australian Museum MM -— Mining Museum Collection, Geological Survey of NSW CPC - Commonwealth Palaeontological Collection NMV - Museum of Victoria SU - Sydney University (Geology Department) Suborder DALMANELLIDINA Moore 1952 Superfamily DALMANELLOIDEA Schuchert 1913 Family DALMANELLIDAE Schuchert 1913 Subfamily RESSERELLINAE Walmsley and Boucot 1971 Genus VISBYELLA Walmsley, Boucot, Harper and Savage 1968 Type species Orthis visbyensis Lindstr6m 1861, by original designation; late Llandovery, Gotland. Diagnosis Subcircular, small valves with apical deltidium and hypercline dorsal interarea; ventral interior with recessive dental plates and cordate muscle scar; dorsal interior with trilobed, dorsally-facing cardinal process and median septum (Harper p. 797 in Kaesler 2000). Visbyella cumnockensis Walmsley, Boucot, Harper and Savage 1968 Fig. 2 (a-g) Synonymy 1968 Visbyella cumnockensis sp. nov.; Walmsley et al., pp. 313-315, pl. 61 figs 6-12. Proc. Linn. Soc. N.S.W., 125, 2004 Type material Holotype AM F67781; paratypes AM F67782- 67788 (formerly SU P19511, 19512-19518; all renumbered when collections were transferred from the University of Sydney to the Australian Museum). New material External and internal moulds of a ventral valve (AM F124331, F124332) and a dorsal valve (AM 167 EARLY SILURIAN BRACHIOPODS FROM ORANGE NSW F124333, F124334) from one bedding plane at locality LM3 (Grid reference 782 988, Cudal 8631 II and II 50 000 topographic sheet, Wallace Creek, Four Mile Creek area south of Orange, N.S.W.); Ulah Formation, Testograptus testis Biozone; late Wenlock (Early Silurian). Diagnosis Relatively small, weakly sulcate, coarsely multicostellate Visbyella with semicircular outline. Dorsal median ridge broad and low posteriorly, becoming narrower and higher to form an anterior median septum (after Walmsley et al. 1968) Description Shell small, almost plano-convex. Ventral valve broadly naviculate, with low suberect beak; dorsal valve weakly convex with shallow but distinct sulcus. Outline suboval, moderately transverse, with straight hinge, obtuse slightly rounded cardinal angles; greatest width at about 0.4Ls. Ventral interarea strongly apsacline, almost flat, apical angle about 120°; delthyrium open, apical angle about 70°, rimmed by narrow crescentic deltidium. Dorsal interarea low, concave, catacline, apical angle about 150°; notothyrium filled by cardinal process, apical angle about 80°. Ribs rather angular, stronger medially than laterally, increasing by bifurcation on the ventral valve, intercalation on the dorsal valve; about 30 counted at ventral valve margin. Ventral interior with prominent subtriangular muscle field, impressed posteriorly but slightly raised anteriorly, length 1/3Ls and width 1/4Ws. Diductor scars elongate oval, divergent, depressed a little below slightly shorter flat adductor field. Raised anterior margin to adductor field distinctly denticulate, extends forward to about 3/4Ls as low ridge. Vascula media flank this ridge as broad, shallow furrows extending from the diductor scars. Muscle field flanked by stout dental plates, divergent forward at about 100° and slightly divergent ventrally, not extending beyond muscle field. Teeth strong, wide, triangular, with distinct crural fossettes on antero-median faces. Valve floor faintly radially furrowed, marginally strongly crenulated. Dorsal interior with prominent oval muscle field extending to 2/3Ld, width 1/3Ws, defined by strong ridges arising just in front of brachiophores and increasingly raised anteriorly, which converge to abut on median septum. Diductor scars impressed, elongate oval, subequal, posterior scars subparallel, anterior scars convergent forward; scars separated by tapering ridge from which rises the stout median septum. Septum highest a little in front of muscle field, and extends to valve margin. Cardinal process large, 168 directed posterodorsally, continuous with well developed notothyrial platform; no shaft. Brachiophores stout, blade-like, divergent ventrally, supported by low, thick plates. Sockets oval, diverging from valve axis at about 75°, deeply excavated into thick triangular socket pads. Valve floor radially grooved, marginally strongly crenulated. Dimensions AM F124332 AM F124334 valve ventral dorsal Ls, Ld est 2.85 2.59 Ws 3.90* 3.73 Wh 3.60* 3.32 Ls/Ws est. 0.73 0.69 Wh/Ws est. 0.92 0.89 * values obtained by doubling exposed half-width, assuming a symmetrical shell. Remarks The Wallace Creek occurrence of this species is almost exactly the same age as the original occurrence at Cumnock, and our admittedly limited new material corresponds closely in all specific characters to the type material. The specimens are slightly larger than shells of the type series (the maximum length and width of any specimens of the type series are 2.1 mm and 3.1 mm respectively), but the ratio Ls:Ws is close to the 2:3 cited for the type material; while the marginal crenulations in the ventral valve are less extensive. The internal moulds of the disarticulated valves are somewhat better than the types, and features of the hinge line can be seen more clearly. The species was also tentatively recorded by Sherwin (1971, p. 223) from the Pridoli Wallace Shale at locality MO/I/27 in the Cheesemans Creek area north of Quarry Creek; his report was the basis for subsequent reports by Pickett (1982, pp. 154-155) and Pogson and Watkins (1998, p. 131). This occurrence is in significantly younger strata than the two other occurrences noted herein. Sherwin’s report was based on several specimens from one locality; we were recently guided to this locality by Dr Sherwin, and collected a further seven specimens of the ‘orthid’ species, which is very rare at the locality (also collected were a few poor specimens of a chonetide, identified as Strophochonetes? cf. savagei Strusz, 2000, and illustrated in Fig. 4 for comparison with Strophochonetes melbournensis). Unfortunately the only internal mould of a dorsal valve of the Wallace Shale orthide (Fig. 2k) is incomplete, and appears to lack a median septum, although its presence anteriorly cannot be completely ruled out. It was initially thought that the absence of a Proc. Linn. Soc. N.S.W., 125, 2004 A.J. WRIGHT AND D.J. STRUSZ septum would rule out the presence of Visbyella. However, one specimen (AM F125552) of Visbyella cumnockensis on one of the type slabs is close in size to the Wallace Shale material and, unlike all the other type specimens, lacks a median septum, so this is not an infallible character of this species. Other morphological features of the Wallace Shale material are not well preserved; there appear to be more than 30 costellae, and the internals of both valves, in so far as they are preserved, are similar to those of the Wallace Creek material (compare Figs 2h-i with Fig. 2a, and Fig. 2} with Fig. 2b). Hence no conclusive argument can be presented to refute the presence of Visbyella at this locality, unlikely as it might seem. This opinion is slightly supported by the presence of a similar orthide (probably Resserella), but definitely lacking a median septum, in the late Ludlow Cardinal View Shale (Bauer 1994) at Bungonia, NSW. Unfortunately, our experience gives us no reason to expect more definitive material at this very unproductive Wallace Shale locality. Suborder CHONETIDINA Muir-Wood 1955 Superfamily CHONETOIDEA Bronn 1862 Family STROPHOCHONETIDAE Muir-Wood 1962 Subfamily STROPHOCHONETINAE Muir-Wood 1962 Genus STROPHOCHONETES Muir-Wood 1962 Type species Chonetes cingulatus Lindstrom 1861, by original designation; Wenlock, Gotland. Diagnosis Shell small, plano- to moderately concavo- convex; well developed median enlarged costa; long, symmetrically arranged high-angled spines varying from intraverse cyrtomorph proximally to orthomorph vertical distally; cardinal process strongly bilobed internally, anteriorly bounded by cardinal process pit; no median septum; anderidia long, narrow, anteriorly divergent at about 60° and isolated on valve floor; inner socket ridges short, thin, as two rounded ridges almost parallel to hinge (after Racheboeuf p. 369 in Kaesler 2000). Strophochonetes melbournensis (Chapman 1903) Fig. 3 Synonymy 1903 Chonetes melbournensis sp. nov.; Chapman, pp. 74-76, pl. XI, fig. 2 only. Proc. Linn. Soc. N.S.W., 125, 2004 1945 Chonetes (Chonetes) melbournensis Chapman; Gill, pp. 132-133. 1953 Chonetes infantilis n. sp.; Opik; p. 15, pl. Ill, figs 19-22. 2000 Strophochonetes melbournensis (Chapman, 1903); Strusz, pp. 249-251, figs 2-3. Type material Lectotype NMV P1419, paralectotypes NMV P615-6, 619, 623, 625-7, 630-633, 637-43 designated by Strusz (2000); Melbourne Formation, Melbourne and South Yarra, Victoria; Ludlow (Late Silurian). Type material of Chonetes infantilis Opik, 1953: holotype CPC 661, paratypes CPC 662-663, Illaenus Band, Wapentake Formation, Heathcote, Victoria; late Wenlock (Early Silurian). New material AM F124306 - 124330, locality W940 (grid reference 743 123, Cudal 8631-II and III 50 000 topographic sheet; south bank of Spring Creek, “Mirrabooka Park’, southwest of Orange, central N.S.W.); Ulah Formation, with Monograptus ludensis; Late Wenlock (Early Silurian). Diagnosis Small, weakly concavo-convex, subquadrate Strophochonetes with up to 5 pairs of gently intraverse- cyrtomorph hinge spines, and finely multicostellate Ornament with median rib on ventral valve usually strongly enlarged. Valve floors heavily papillose, ventral muscle field distinct, anderidia short and diverging at 60-80° (after Strusz 2000). Description Shell small, plano-convex, ventral valve of very low convexity. Outline subquadrate, lateral margins gently sigmoid, with shallow re-entrants in front of small triangular ears; hinge width usually slightly less than greatest width (mean Wh/Ws 0.93). Ventral protegulum posteromedially furrowed, variably raised above remaining shell surface; distinct protegular lobe, weaker lateral lobes on dorsal valve. Maximum observed width 9.8 mm, length 6.5 mm, most specimens being much smaller; mean Ls/Ws 0.75, ratio decreasing with increasing shell size. Interareas mostly obscure; ventral interarea apparently low, apsacline, flat, delthyrium wide, beak very low; pseudodeltidium not seen; dorsal interarea linear, attitude unclear. Myophore small, projecting posteroventrally, bifid, each lobe less strongly bifid, flanked by small but distinct cardinal crests. Chilidium obscure, might be present as very narrow ridge wrapped around base of myophore. Hinge spines fine, relatively long, upright 169 EARLY SILURIAN BRACHIOPODS FROM ORANGE NSW Figure 3. Strophochonetes melbournensis (Chapman, 1903). a-g, ventral valves; some hinge spines are visible in b-e, only spine bases in f-g. a, juvenile valve AM F 124320. b, juvenile with particularly prominent protegulum, AM F124317. c, juvenile AM F124322. d, AM F124324. e-f, internal mould and latex cast, AM F124312. g, AM F124326. h-j, dorsal valves; h-i, incomplete external mould and latex cast showing well developed protegular and lateral nodes, AM F 124318. j, latex cast of incomplete interior, AM F124307. Scale bar 3 mm. ? or nearly so (initial angle with hinge line about 60- 80°), straight (particularly in small specimens) to gently cyrtomorph intraverse, symmetrically placed; up to 4 each side of beak (AM F124324). Ornament of fine, rounded radial ribs, 29-34 counted in 5 mm at 5 mm radius, separated by narrower furrows; increase is by bifurcation only. Median rib on ventral valve prominent, arises within protegulum; remaining ribs arise at or in front of margins of concentrically wrinkled protegular regions. Ventral interior with low, narrow median . septum, reaching forward to about 0.2Ls; septum posteriorly raised and slightly widened. Teeth small, widely divergent. Muscle field generally obscure other than for weak or absent endospines; in one specimen (AM F124312) the field is weakly impressed, with 170 small, elongate subtriangular, slightly divergent adductor scars further impressed posteriorly. Remainder of valve floor densely covered by fine endospines radially arranged beneath ribs, weakest towards cardinal margin and ears. Dorsal interior still not well known. Cardinal process small, internally bifid, fused to short but strong inner socket ridges which are curved parallel to hinge margin. Short, shallow furrow in front of cardinal process, but no median ridge developed. Anderidia visible in only one specimen (AM F124307); they are short (0.2Ld), fine, low, diverging at about 60°. Muscle field obscure. Distal two-thirds of valve floor with numerous small radially arrayed endospines, as in ventral valve. Proc. Linn. Soc. N.S.W., 125, 2004 A.J. WRIGHT AND D.J. STRUSZ Dimensions valve Ls, Ld Ws AM F124307 dorsal 49 = F124312 ventral est. 4.8 5.6* F124318 dorsal 55) - F124322 ventral 3.6 47 F124324 ventral 5.3 TD, F124326 ventral 5.6 8.4 Wh = Ls/Ws Wh/Ws Si = 5.4* est. 0.86 0.96 AG Os: 0.94 7.2 0.74 1.00 7.2 0.67 0.86 * values obtained by doubling exposed half-width, assuming a symmetrical shell. Discussion Although preservation is not particularly good, the Wenlock specimens from Spring Creek conform in all important aspects (very low ventral convexity, rib increase only by bifurcation, and less prominent protegular and lateral lobes on the dorsal valve) with S. melbournensis rather than S. kemezysi Strusz, 2000. Some of the minor differences could be related to the small size of most of the specimens (several are clearly juvenile, none approaches the maximum size recorded for the Victorian material). Some could be of age Significance, but without better and more abundant material from older levels in Victoria this remains unclear. Thus no ventral valves show the anterior sulcus seen in some Victorian Late Silurian shells, and no more than 4 spines have been seen to either side of the ventral beak. The NSW specimens tend also to be more elongate (Ls/Ws very variable, mean 0.76; for Victorian specimens the mean is 0.61). Internally, the ventral muscle field is less obvious, and there are no coarser endospines near the hinge. In this last respect, and in a greater tendency for spines on small specimens to be straight, the Late Wenlock Spring Creek specimens are more like the few poor specimens from the Early Wenlock of Heathcote than the Ludlow material from Melbourne. Dorsal interiors, while still few and inadequate, do add some information, 3mm Figure 4. Strophochonetes? cf. savagei Strusz, 2000. Latex cast of ventral valve, MM F21133. Scale bar 3 mm. Proc. Linn. Soc. N.S.W., 125, 2004 particularly the form of the cardinal process and its flanking cardinal crests. The presence internally of a weak posteromedian dorsal furrow instead of a low ridge places these specimens closer to typical Strophochonetes than are the type specimens. Three similar chonetoide specimens (MM F21133, 37435, 37436) are available from the Wallace Shale locality - the best of them is figured (Fig. 4). All are small and weakly convex. In the absence of internal data, particularly of the dorsal valve, generic identity must remain uncertain. The long more or less upright hinge spines, low ventral valve convexity, fine ribbing and accentuated median rib all indicate Strophochonetes, however, and of the Australian taxa described by Strusz (2000) the closest is undoubtedly S? savagei from the Early Lochkovian of Manildra, northwest of Orange. S. melbournensis and S. kemezysi Strusz, 2000, while superficially similar, are both larger and more coarsely ribbed; the latter has very prominent protegulae. In only one respect these specimens appear unlike typical Strophochonetes, and that is in the alternating pattern of hinge spine insertion described for instance by Strusz (2000, p. 259) for the strongly convex and fairly coarsely ribbed Australian species of Johnsonetes Racheboeuf, 1987 (all of which lack spine 1’). However it is not clear that spine 1' is undeveloped in the Wallace Shale specimens. Moreover, the Manildra species show considerable variation in spine form, and some asymmetry cannot be ruled out. ACKNOWLEDGMENTS We gratefully acknowledge access graciously made available by Ian Street to ‘Mirrabooka Park’ and Ken Williams to ‘Ashburnia’, and thank Dr L. Sherwin for drawing our attention to the report of Visbyella cf. cumnockensis from the Wallace Shale and subsequently guiding us to the locality. Prof. Barrie Rickards and Dr Lucy Muir kindly allowed us to cite identifications of the graptolites. Robert Jones readily made the type material of Visbyella cumnockensis available for study. Strusz wishes to thank Dr Patrick DeDeckker for providing facilities at the 171 EARLY SILURIAN BRACHIOPODS FROM ORANGE NSW Australian National University; Wright’s research has been supported by the University of Wollongong and the Linnean Society of NSW. REFERENCES Bauer, J.A. (1994). Siluro-Devonian Bungonia Group, Southern Highlands, N.S.W. Helictite 32(2), 25-34. Chapman, A.J., Rickards, R.B., Wright, A.J., and Packham, G.H. (2003). Dendroid and tuboid graptolites from the Llandovery (Silurian) of the Four Mile Creek area, New South Wales. Records of the Australian Museum. Chapman, F. (1903). New or little-known Victorian fossils in the National Museum. Part II - some Silurian Molluscoidea. Proceedings of the Royal Society of Victoria 16, 60-82. Gill, E.D. (1945). Chonetidae from the Palaeozoic rocks of Victoria and their stratigraphical significance. Proceedings of the Royal Society of Victoria 57, 125- 150. Jenkins, C.J. (1978). Llandovery and Wenlock stratigraphy of the Panuara area, central New South Wales. Proceedings of the Linnean Society of New South Wales 102, 109-130. Jenkins, C.J. (1986). The Silurian of mainland Australia: a field guide. (UGS Silurian Subcommission and University of Sydney: Sydney). Kaesler, R.L. (ed.) (1997). Treatise on Invertebrate Paleontology, Part H, Brachiopoda, revised, volume 1: Introduction. (Geological Society of America and University of Kansas Press: Lawrence, Kansas). Kaesler, R.L. (ed.) (2000). Treatise on Invertebrate Paleontology, Part H, Brachiopoda, revised, volumes 2-3: Linguliformea, Craniiformea, and Rhynchonelliformea (part). (Geological Society of America and University of Kansas Press: Lawrence, Kansas). Lindstrém, G. (1861). Bidrag till kinnedomen om Gotlands brachiopoder. Ofversigt af kungliga Vetenskaps- Akademiens Férhandlingar, Stockholm for 1860, 17, 337-382. Muir-Wood, H. (1962). On the morphology and classification of the brachiopod suborder Chonetoidea. (British Museum of Natural History: London). Opik, A.A. (1953), Lower Silurian fossils from the “I//aenus Band”, Heathcote, Victoria. Geological Survey of Victoria, Memoir 19, 1-42. Packham, G.H., and Stevens, N.C. (1955). The Palaeozoic stratigraphy of Spring and Quarry Creeks, west of Orange, N.S.W. Journal and Proceedings of the Royal Society of New South Wales 88, 55-60. Pickett, J.W. (ed.) 1982. The Silurian System in New South Wales. Geological Survey of New South Wales, Bulletin 29,1- 264. Pogson, D.J., and Watkins, J.J. (compilers) 1998. “Bathurst 1: 250 000 Geological Sheet $1/55-8: Explanatory Notes’. (Geological Survey of New South Wales: Sydney). Racheboeuf, P.R. (1987). Upper Lower and Lower Middle Devonian chonetacean brachiopods from Bathurst, Devon and Ellesmere Islands, Canadian Arctic Archipelago. Geological Survey of Canada, Bulletin 375, 1-29. Rickards, R.B., Percival, I1.G., Simpson, A.J. and Wright, A.J. (2001). Silurian biostratigraphy of the Cadia area, near Orange, New South Wales. Proceedings of the Linnean Society of New South Wales, 123, 173-191. Rickards, R.B., and Wright, A.J. (1997). Graptolite zonation of the late Wenlock, with a new graptolite- brachiopod fauna from New South Wales. Records of the Australian Museum 49, 229-248. Sherwin, L. (1971). Stratigraphy of the Cheesemans Creek district, New South Wales. Records of the Geological Survey of New South Wales 13, 199-237. Strusz, D.L. (2000). Revision of the Silurian and Early Devonian chonetoidean brachiopods of southeastern Australia. Records of the Australian Museum 52, 245-287. Walmsley, V.G., Boucot, A.J., Harper, C.W. and Savage, N.M. (1968). Visbyella - a new genus of resserellid brachiopod. Palaeontology 11, 306-316. Proc. Linn. Soc. N.S.W., 125, 2004 Early Silurian Graptolites from Cadia, New South Wales R.B. Rickarps! AND A.J. WricHT” ‘Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge CB2 3EQ, England and *School of Earth and Environmental Sciences, University of Wollongong, Wollongong NSW 2522. Rickards, R.B. and Wright, A.J. (2004). Early Silurian graptolites from Cadia, New South Wales. Proceedings of the Linnean Society of New South Wales 125, 173-175. A low-diversity graptolite fauna is reported from the Ulah Formation at Cadia, central western New South Wales. The assemblage includes Testograptus testis, Monoclimacis flumendosae, fragments of Monograptus flemingii, possible Cyrtograptus and unidentifiable retiolitid meshworks, and is correlated with the Jungreni- testis Biozone, of late Wenlock (Early Silurian) age. Manuscript received 3 May 2003, accepted for publication 23 July 2003. KEY WORDS: Cadia, graptolites, Lower Silurian, Wenlock. INTRODUCTION Three Silurian faunas were documented by Rickards et al. (2000) from the vicinity of Cadia open cut, south of Orange, New South Wales. One of these faunas, of late Wenlock-early Ludlow aspect, consisted of shelly fossils and graptolites collected by Dr Ian Percival from a slumped mudstone at a locality on the access road to the Cadia open cut. This fauna was discussed and illustrated by Rickards et al. (2000), who figured but could not determine the poor graptolite material to genus or species because of the poor preservation of the fragmentary material. The locality (grid reference 687240E, 6295047 N, Canowindra 8360-N 1:50 000 topographic sheet) is on the eastern face of the access road to the Cadia open cut, about 1 km from the entrance gates; a map of the region showing the location of this and other fossil localities was provided by Rickards et al. (2000, Fig. 1). The fossiliferous strata are considered to correlate with the Ulah Formation, at Four Mile Creek west of Cadia (see Rickards et al. 2000, Fig. 1), in which the Testograptus testis fauna occurs. NOTES ON THE GRAPTOLITE FAUNA Since the publication of Rickards et al. (2000), we have made a further but small graptolite collection from the Cadia mine shelly fossil locality which permits fuller identification of the low-diversity fauna and determination of its age. The Cadia graptolite fauna consists of Testograptus testis (Barrande), Monoclimacis flumendosae (Gortani), fragments of Monograptus flemingii (Salter), fragmentary stipes possibly belonging to Cyrtograptus, and fragmentary retiolitid meshworks which cannot be assigned, even approximately, to a genus. In discussing this as ‘the Cadia graptolite fauna’ we are mindful of the presence of other graptolites in Silurian strata in the vicinity of the Cadia mine. Full documentation of any such graptolite faunas as that documented here is important as graptolite localities in the vicinity of Cadia mine (such as the Pridoli ‘borrow pit’ locality, W910 of Rickards et al. 2000) are very much less common than at Four Mile Creek, and are under threat. A brief review of graptolites previously reported from Cadia by Offenberg (1963) was given by Rickards et al. (2000). We have not provided here any systematic descriptions of the fauna, but limited comments on the morphological detail are included in the explanatory text for Figure 1. The Cadia specimens have undergone soft sediment deformation, with a considerable amount of twisting and breakage, in contrast to the Rodds Creek black shale specimens (Rickards et al. 2000) which were undeformed other than by diagenetic flattening. AGE OF THE CADIA GRAPTOLITE FAUNA The dominant species is Testograptus testis (Barrande), which normally indicates the late Wenlock (Early Silurian) /undgreni-testis Biozone. Testograptus testis has been recorded, very rarely, from the /udensis SILURIAN GRAPTOLITES FROM NEW SOUTH WALES Figure 1. (A) Monoclimacis flumendosae (Gortani), AM F 114926, distal thecae, undeformed, low relief. (B-E) Testograptus testis (Barrande). (B) proximal end, AM F114928, showing some soft sediment deformation distally; (C) AM F114925, a proximal end with spines visible on thl1; (D) AM F114930, spines on several thecae; (KE) AM F114929, distal thecae with a growing end visible. (F) Monograptus flemingii (Salter), AM F114927, subscalariform view of mesial thecae. All figures x10, scale bar 1mm; heavy bar indicates deformation stretching direction, possibly not tectonic. All specimens from locality W 937, grid reference 687240E, 6295047 N, Canowindra 8360-N 1:50 000 topographic sheet. Unfigured specimens are AMF 114931-940. Biozone (Rickards et al. 1995) but, as the Cacia specimens are abundant and occur with Monoclimacis flumendosae (Gortani), a pre-ludensis Biozone is indicated for this fauna. The Cadia fauna is probably slightly younger than the Rodds Creek fauna (Rickards et al. 2000). Although this latter assemblage included some lundgreni-testis Biozone indicators, the presence of Cyrtograptus ex gr. rigidus Tullberg indicated a. probable middle rather than late Wenlock for the Rodds Creek fauna. The Cadia fauna is thus significantly older than the Pridoli fauna from the “borrow pit’ locality (W910) 2 km to the southeast (Rickards et al. 2000). 174 Correlation with the Four Mile Creek sequence is probably with testis-bearing beds of the Ulah Formation in Wallace Creek; in Spring and Quarry Creeks, the testis-bearing beds of the same formation are largely green and black mudstones (Packham, Rickards and Wright, unpublished data). SHELLY FAUNAS The disarticulated and fragmental shelly fauna in this slump unit is unusually abundant and diverse for the region, in contrast with clastic units of Proc. Linn. Soc. N.S.W., 125, 2004 R.B. RICKARDS AND A.J. WRIGHT this age in the Four Mile Creek area and the Spring- Quarry Creek areas which are singularly poor in shelly fossils. The faunas at Cadia have undergone soft- sediment deformation and are clearly transported. Described shelly faunas (other than corals) from the Four Mile Creek area and the Spring Creek areas are limited to two species of Judensis Biozone brachiopods described by Rickards and Wright (1997) from Cobblers Creek (see Fig. 1 of Rickards et al. 2000) and by Wright and Strusz (2004) from Spring Creek and Wallace Creek (see Fig. 1 of Rickards et al. 2000: ludensis Biozone and lundgreni-testis Biozone respectively). Other brachiopod faunas from the region were listed by Jenkins (1978, 1986), but the only rich faunas cited by him are from Llandovery (Early Silurian) limestones. CONCLUSIONS Graptolites identified from the Cadia Mine access road locality are Testograptus testis, Monoclimacis flumendosae, fragments of Monograptus flemingii, ?Cyrtograptus and retiolitids. The fauna is late Wenlock (Early Silurian) and is probably best correlated with a level high in the lundgreni-testis Biozone. It appears to be slightly younger than the probably middle Wenlock Rodds Creek black shale fauna (Rickards et al. 2000), and is assumed to correlate with the testis fauna of the Ulah Formation in the Four Mile Creek area to the west of Cadia. ACKNOWLEDGEMENTS We are grateful to Ian Tedder (Newcrest Mining Limited) for allowing access to, and guiding us to, this locality in November 2001. The universities of Cambridge and Wollongong have provided financial support for participation by RBR and AJW in this study, and funds were provided to AJW by the Linnean Society of New South Wales through the Betty Mayne Fund. Proc. Linn. Soc. N.S.W., 125, 2004 REFERENCES Jenkins, C.J. (1978). Llandovery and Wenlock stratigraphy of the Panuara area, central New South Wales. Proceedings of the Linnean Society of New South Wales 102, 109-130. Jenkins, C.J. (1986). The Silurian of mainland Australia: a field guide. 82 p. {UGS Silurian Subcommission, Sydney. Offenberg, A.C. (1963). Geology of the Panuara-Cadia- Errowanbong area, south of Orange, New South Wales. BSc (Hons) thesis, University of Sydney (unpublished), 112 p. Rickards, R.B., Packham, G.H., Wright, A.J. and Williamson, P.L. (1995). Wenlock and Ludlow graptolite faunas and biostratigraphy of the Quarry Creek district, New South Wales. Association of Australasian Palaeontologists, Memoir 17, 1-68. Rickards, R.B., Percival, 1.G., Simpson, A.J. and Wright, A.J. (2000). Silurian biostratigraphy of the Cadia area, near Orange, New South Wales. Proceedings of the Linnean Society of New South Wales 123, 173-191. Rickards, R.B. and Wright, A.J. (1997). Graptolite zonation in the late Wenlock (Early Silurian), with a new graptolite-brachiopod fauna from New South Wales. Records of the Australian Museum 49, 229-248. Wright, A.J. and Strusz, D.L. (2004). Wenlock (Early Silurian) brachiopods from the Orange district of New South Wales. Proceedings of the Linnean Society of New South Wales 125, 165-172. 175 yelaas wat wey: thy wie ace WE ye ee Anis a a fetes i Salter 5, 4 ae EE a om? Ccsiias od i be 2 ‘TRI wate er AEH RT WA ats ra yn oe “praensy ; : ay es bo aie fond wiper at ae rive "sy; oie Nhe iy se : fe ge bods Tt wer y tuclegsia ou: noid paste ogy’) ar) sBeiginbse ataokaa bith wy frat. F ree. Es SS atti dist “Thad: nog rab, wad pibeD ory pense} ont _netge’ Tayo wr Sit fansipes em seston tx ee ae bstioqeass yttnsi. 298 bak’ sipvmirlOteb " Mbetryrais We aniog nigaputts GAL Yo Ps ee _ fal cing Kelaped nied’ ssdiI0) ewan ‘lls | oy mn Pl : ai oa ee a er on at di sl ies og Po) gange, ob bre, hs oti + aqlont ooh widieds RT Wy saint his 1 AItS pe j i recht AC: : as em ext eM ae Ay se Tlary Fala Sr ebqqaipioin) ano penttxiynsiul Weedioage. od wisi : % “Jeaming a, Agr LATeRt). JAgiiW bape | ene) Og: Ye) ‘pri THe ety ix "We eecini if nga yf a6 aaah, be sy ant 3 = Ne eipiaitvanz A Lanta itisiod ‘Noe ity dea. aa ae a ho th AN oo avi peyobatisendGoraiealinilihe tee 5, ede He A : z hae : nigtt Aw od tA \ aaa a eer aaal 148°50' Rylstone Figure 1. Location of trilobite collection at Brogans Creek. Map of NSW indicates Taemas, where the same species have been described (Chatterton 1971). Shading in inset map (after Colquhoun 1995) shows distribution of Lower Devonian platform sediments. 178 Proc. Linn. Soc. N.S.W., 125, 2004 G.D. EDGECOMBE AND A.J. WRIGHT area of NSW indicate that the Receptaculites and Warroo limestones at Taemas, which overlie the Cavan Formation with its Polygnathus pireneae to P. dehiscens fauna, are probably early Emsian. Lindley (2002) recorded Polygnathus nothoperbonus from the Warroo Limestone Member, further confirming the assignment of this limestone to the medial Emsian Polygnathus perbonus Conodont Zone. However, Basden et al. (2002) concluded that the Warroo Limestone should be correlated with the Polygnathus inversus to P. serotinus Conodont Zones. On balance the co-occurrence of P. nothoperbonus in both areas of NSW seems to indicate unequivocally a medial Emsian age for the macrofaunas. This supports the conclusions of Garratt and Wright (1988:Fig. 3), who correlated their Malurostrophia-Taemostrophia- Howittia fauna (essentially the shelly fauna discussed here) with the Polygnathus gronbergi (=P. perbonus) Conodont Zone. Faunal characters and affinities The fossiliferous limestones have yielded very rich and well-preserved invertebrate faunas, dominated by brachiopods, tabulate corals and tetracorals, trilobites, gastropods, ostracodes, cephalopods, tentaculitids, crinoid debris and sponges; bivalves are subordinate at this locality. Most of the trilobites and brachiopods at Brogans Creek are conspecific with those described from Emsian limestones in the Lake Burrinjuck sequence at Taemas and ‘Bloomfield’ by Chatterton (1971, 1973). With respect to the trilobites, the faunal composition of the Brogans Creek assemblage is best matched in the lower half of the Receptaculites Limestone at Locality I of Chatterton (1971). The three species identified here, Proetus nemus, Dentaloscutellum hudsoni and Acanthopyge bifida, are represented in the lower Receptaculites Limestone at Locality I and at that locality as well as Brogans Creek they occur with Sthenarocalymene. Silicified residues from Brogans Creek yield the following for minimal number of individuals per species, based on the most abundant skeletal element: Proetus nemus (N=54), Dentaloscutellum hudsoni (N=16), Acanthopyge bifida (N=7), and Sthenarocalymene sp. (N=2). About 120 kilograms of limestone have been etched to produce our fauna. In terms of diversity, the silicified assemblage consists additionally of more than 15 brachiopod species (Malurostrophia flabellicauda reverta Chatterton; Salopina kemezysi Chatterton and other dalmanellids; Schuchertella murphyi Chatterton; Coelospira dayi Chatterton; Howellella sp.; Ambothyris runnegari Chatterton; Howittia sp.; Proc. Linn. Soc. N.S.W., 125, 2004 ?Buchanathyris sp.; reticulariid indet.; Cydimia parva Chatterton; Parachonetes flemingi Chatterton; P. sp. cf. P. konincki Chatterton; rhynchonellids). Some 30 gastropod species are under study by Dr A.G. Cook. Tetracoral species are dominated numerically by an abundant solitary Plasmophyllum, as well as other solitary corals (?acanthophyllids) and rare fragments of ?Calceola. The sponge Amphipora is locally abundant, and presumably represents lagoonal phases of deposition or influx of lagoonal debris; several biofacies are evident. Colquhoun (1998) indicated that the Brogans Creek limestone was deposited in a well- oxygenated, normal salinity environment. The trilobite material is represented by disarticulated sclerites, but many brachiopods shells are articulated. Scolecodonts are at least as common as conodonts in residues; this is also a feature of limestones in the Capertee Valley (S of Brogans Creek) where the strata are highly deformed and preservation is poor. Despite the disarticulated nature of parts of the Brogans Creek shelly fauna, their excellent preservation indicates that postmortem transportation was minimal. SYSTEMATIC PALAEONTOLOGY Figured material is in the Palaeontology collection, Australian Museum, Sydney (prefix AMF). Order PROETIDA Fortey and Owens, 1975 Family PROETIDAE Salter, 1864 Subfamily PROETINAE Salter, 1864 Genus PROETUS Steininger, 1831 Type species Calymmene concinna Dalman, 1827; by original designation. Proetus nemus Chatterton, 1971 Fig. 2a-p, Fig. 3a-t Proetus nemus Chatterton, 1971:65-67, Pl. 16, Figs 18-32. Ryckholtia? nemus (Chatterton). Liitke, 1990:21. Material 39 cranidia, 103 librigenae, 3 hypostomes, 62 thoracic segments, 50 pygidia. Diagnosis Proetus with relatively elongate, tapering glabella, its posterior two thirds with dense, mostly moderate sized tubercles, its anterior third granulate. Facial suture divergent between y and . Genal ridge strong along 179 EARLY DEVONIAN TRILOBITES FROM N.S.W. Figure 2. Proetus nemus Chatterton, 1971. Carwell Creek Formation (medial Emsian), Brogans Creek, NSW. Scale bars 1 mm. a-c, AMF 124700, cranidium, dorsal, anterior and lateral views; d-f, AMF 124701, cranidium, dorsal, anterior and lateral views; g, AMF 124702, cranidium, dorsal view; h, AMF 124703, cranidium, dorsal view; i, AMF 124704, cranidium, lateral view; j, AMF 124705, cranidium, anterior view; k, AMF 124706, cranidium, dorsal view; Il-m, AMF 124707, cranidium, dorsal and lateral views; n, AMF 124708, librigena, dorsal view; 0, AMF 124709, librigena, dorsal view; p, AMF 125485, cranidium, dorsal view. Proc. Linn. Soc. N.S.W., 125, 2004 G.D. EDGECOMBE AND A.J. WRIGHT Figure 3. Proetus nemus Chatterton, 1971. Carwell Creek Formation (medial Emsian), Brogans Creek, NSW. Scale bars 1 mm. a, AMF 124710, librigena, internal view; b, AMF 124711, librigena, dorsal view; c, AMF 124712, librigena, dorsal view; d-e, AMF 124713, hypostome, ventral and lateral views; f, AMF 124714, thoracic segment, dorsal view; g, AMF 124715, thoracic segment, dorsal view; h-j, AMF 124716, pygidium, posterior, lateral and dorsal views; k, AMF 124717, thoracic segment, dorsal view; 1, AMF 124718, thoracic segment, anterior view; m, AMF 124719, thoracic segment, anterior view; n, AMF 124720, pygidium, dorsal view; o-p, AMF 124721, pygidium, lateral and dorsal views; q-r, AMF 124722, pygidium, posterior and dorsal views; s, AMF 124723, pygidium, ventral view; t, AMF 124724, pygidium, dorsal view. Proc. Linn. Soc. N.S.W., 125, 2004 | 181 EARLY DEVONIAN TRILOBITES FROM N.S.W. all but posteriormost part of librigenal field, distinct but less prominent on preocular fixigena; small caecal pits abundant on librigenal field; genal spine relatively long. Pygidium with seven axial rings and lunate terminal piece (7+1); anterior three or four pleural furrows well impressed, fifth and sixth faint. Description Cranidial length about equal to maximum width at o; width at 6 slightly more than 80% width at w; width at B 85-95% width at 6. Axial furrow narrow, moderately, evenly deep. Glabella widest basally, length (excluding LO) 1.1-1.2 times basal width, with moderate taper anteriorly; slightly constricted at S2, gently convex (sag., tr.); frontal lobe rounded; terminating at but not overhanging anterior border furrow. S1 originating opposite midlength of palpebral lobe, shallow, directed posteromedially, distally birfucate, with posterior branch terminating well in front of SO; S2 parallel with S1, more weakly incised, originating just behind anterior edge of palpebral lobe; S3 obscure. Posterior two thirds of glabella with mostly moderate sized tubercles, some small tubercles, densely packed so as to nearly touch; anterior third of glabella granulate, non-tuberculate. SO transverse medially, narrow (sag., exsag.), deep, flexed forwards abaxially against lateral occipital lobes. LO distinctly wider than basal part of glabella, length about 20% its width; lateral occipital lobes large, drop-shaped, isolated from remainder of LO by deep furrows; LO, including lateral lobes, covered with tubercles as on posterior part of glabella, including moderately large median tubercle behind midlength. Preglabellar region 13-15% of cranidial length; in large specimens, composed of an inclined, medially flat posterior half and moderately convex (sag.) anterior half bearing 5-6 terrace lines in dorsal view; in small specimens, posterior half forms a wide (sag., tr.) depressed field with a broad (tr.), gently inflated transverse median swelling. Genal ridge well developed on preocular fixigena, anteromedially directed, terminating at juncture of preglabellar and anterior border furrows, stronger in small specimens. Postocular fixigena 25-35% width (tr.) and about 60% length (exsag.) of LO. Palpebral lobe arcuate, 35-45% length of glabella; palpebral furrow faint or indistinct. Anterior sections of facial suture diverging from each other at 45-62° between y and B, running subparallel against anterior border furrow, then strongly converging between B and a. Posterior sections of facial suture running subparallel or gently diverging between e€ and €, close to axial furrow, then sharply turned outwards to @. Librigenal field moderately wide, gently convex (tr.); genal ridge strong along all but 182 posteriormost part of field, closer to eye socle than to lateral border furrow; most of field with abundant, small caecal pits, least distinct at posterolateral corner of field. Eye socle narrow, separated from visual surface and librigenal field by shallow furrows. Posterior border furrow narrow, deep; lateral border furrow wider, the two merging at genal angle, extending along a variable extent of the genal spine, usually along about half its length. Lateral border 70- 80% as wide as narrowest part of librigenal field in dorsal view, strongly convex (tr.); terrace lines well defined along entire length and width of lateral border and along genal spine. Genal spine relatively long, its inner margin straight or faintly concave. Panderian notch large, semicircular. Connective suture with straight, diagonal course along most of its length, its extent relative to cranidium indicating that rostral plate is trapezoidal or triangular, fairly wide anteriorly (cf. P. concinnus: Owens 1973:Text-fig. 1B). Hypostomal width across shoulders about 65% sagittal length. Anterior margin weakly convex medially, flexed backward abaxially. Anterior lobe of middle body strongly inflated (tr.), anteromedial part raised but not forming discrete rhynchos; middle body gently convex (sag.) along most of length, fairly steeply turned up anteromedially; anterior lobe bearing many sinuous terrace lines. Middle furrow moderately deep, ~ directed posterolaterally across abaxial third of middle body then abruptly effacing. Border furrow narrow, distinctly impressed around entire middle body, shallowest against anterior wing. Anterior border uniformly narrow (sag., exsag.); lateral border gently converging between anterior wing and shoulder; shoulder rounded; posterolateral margin straight between shoulder and pair of blunt spines at lateral edge of posterior border; posterior border narrow (sag., exsag.), about 10% length of hypostome, with gently convex posteromedian margin. Number of thoracic segments unknown. Axial furrow narrow, shallow. Axis strongly convex (tr.), 32-41% width of thorax. Articulating half ring varying from equal in width (sag.) to 1.6 times as wide as preannulus along length of thorax, 70-90% length of ring; preannular furrow transverse to gently concave medially, sharply impressed but much shallower than articulating furrow; ring covered with small, dense tubercles or coarse granules. Pleural furrow narrow, about as deep as articulating furrow, gently flexed forward at fulcrum, abruptly shallowing then effacing on inner part of articulating facet; anterior and posterior pleural bands equal in width (exsag.) proximal to fulcrum; pleurae moderately declined abaxial to fulcrum, at midwidth (tr.) of rib. Pleural tips with curved anterolateral margin, blunt rounded posterior Proc. Linn. Soc. N.S.W., 125, 2004 G.D. EDGECOMBE AND A.J. WRIGHT projection. Panderian notch deep, U-shaped. Pygidium subsemicircular, length (excluding articulating half ring) 55-60% width. Axial width about 35% pygidial width anteriorly; axial furrows narrow, uniformly impressed along most of length. Seven axial rings and short, lunate terminal piece (7+1); first one or two ring furrows lengthened medially as short preannulus; more posterior ring furrows shallower but with moderately deep incision across axis, posterior few gently convex backwards; axis raised strongly above pleurae, gently convex in sagittal profile, moderately arched (tr.); rings with dense small tubercles or coarse granules. Postaxial region about 20% length of pygidium. Pleural furrows narrow (exsag.), anteriorly convex, anterior three or four well impressed, fifth and variably sixth faintly discernible; first pleural furrow terminates near pygidial lateral margin, others terminate at shallow posterior border furrow; interpleural furrows narrower and shallower than pleural furrows; pleural ribs with sculpture of dense, medium sized granules. Border widening back to its intersection with third pleural furrow, then maintaining even width, occupying most of postaxial region, weakly convex. Doublure extending in nearly as far as border furrow, bearing several terrace lines. Discussion The sample from Brogans Creek resembles that from Taemas in that the largest cranidia (Fig. 2a-c, h, i, p; Chatterton 1971:Pl. 16, fig. 28) have the anterior end of the glabella abutting the inclined posterior part of the anterior border, whereas small specimens have a broad depression between the frontal lobe and the convex, terraced part of the anterior border (Fig. 2d-f, j, k; Chatterton 1971:Pl. 16, fig. 25). The latter morphology, associated with a more pronounced fixigenal ridge (Fig. 2d, k versus 2a, h, p) is confined to small specimens. This difference in the preglabellar region is bridged by intermediate sized specimens, and is ascribed to ontogenetic variation. The transverse median swelling in the depression of small specimens (Fig. 2e, j) retains a faint expression in large cranidia. No bimodality can be detected in the strength of the librigenal ridge (Figs. 2n, 0, 3b, c), which is consistently pronounced. In assigning this species to Proetus, Chatterton (1971) acknowledged its distance from the type species, the Wenlock P. concinnus (Dalman). However, several other Australian Emsian and Eifelian Proetinae are validly assigned to that genus. These include Proetus talenti Chatterton, 1971 (type of Devonoproetus Liitke, 1990), P. sparsinodosus Feist and Talent, 2000, and P. latimargo Feist and Talent, . 2000, the latter two originally assigned to Proc. Linn. Soc. N.S.W., 125, 2004 Devonoproetus at the subgeneric level. Devonoproetus iS ajunior synonym of Proetus s.s. (Adrain 1997; Zhou et al. 2000). Proetus nemus was reassigned, with question, to the otherwise Ludlow-Lochkovian Ryckholtia Snajdr, 1980 (type Proetus ryckholti Barrande, 1846) by Liitke (1990). The new material described herein conflicts with this reassignment. Membership in Ryckholtia is precluded by the pronounced tuberculate sculpture on the glabella and axial rings of P. nemus, the strongly defined lateral occipital lobes, and sagittal elimination of the preglabellar field. This species displays characters that suggest alternative assignments. The elongate, tapering glabella of Proetus nemus and its pattern of sculpture (strong tuberculation posteriorly, becoming subdued anteriorly), together with the profile of the preglabellar region, including the wide (sag., exsag.) anterior cranidial border furrow, and the divergence of the facial suture between y and B resemble Longiproetus tenuimargo (Richter, 1909) (type of Longiproetus Cavet and Pillet, 1958). Longiproetus has been regarded as a synonym of Gerastos Goldfuss, 1843 (Owens 1973), a valid subgenus of Gerastos (Snajdr 1980), restricted to its type species on the basis of a distinctive shape of the rostral plate (Liitke 1990), or slightly expanded to include a small group of Rhenohercynian mid Eifelian to early Givetian species (Basse 1996, 2002). Liitke (1990) reassigned the Bohemian species that had been referred to Longiproetus (e.g., Snajdr 1980) to Coniproetus Alberti, 1966, and other genera, whilst the inadequately known Emsian species referred to Longiproetus by Pillet (1972) defy classification. Despite the similarities in the glabella and preglabellar region, several characters conflict with an alliance between P. nemus and Longiproetus. Notably, the strong genal ridge of P. nemus is lacking in L. tenuimargo and other certain congeners (sensu Basse 2002), the prominent lateral occipital lobes contrast with the inconspicuous lobes in Longiproetus s.s., LO is wider than the basal part of the glabella, the cephalon is much less vaulted, the palpebral lobe is situated more posteriorly, and the pygidium is relatively paucisegmented (7+1 rings versus 8+1). The course of well preserved connective sutures on librigenae suggests that the rostral plate of P. nemus is more regularly trapezoidal or triangular than is that of L. tenuimargo (Liitke 1990:Text-fig. 8). Affinities to species that have been assigned to Devonoproetus by recent workers better account for the large occipital lobes, width of LO relative to the glabella, and 7+1 pygidial segmentation. Among these, Proetus latimargo Feist and Talent, 2000 (Eifelian, Queensland) and P. zhusilengensis Zhou et al., 2000 183 EARLY DEVONIAN TRILOBITES FROM N.S.W. (Emsian, Inner Mongolia) resemble P. nemus in having a tongue-shaped glabella (narrowest in P. nemus) with dense, pronounced tuberculation, and P. latimargo shares the divergence of the facial suture between o and . Among those species that have been referred to Devonoproetus, the strong genal ridge of Proetus nemus is developed in a group recognised by Basse (2002) as a separate genus, Rhenocynproetus, from which the Australian “Devonoproetus” species were explicitly excluded. The presence of a genal ridge in other genera of Proetinae [e.g. Gerastos: Snajdr 1980:Pl. 3, Fig. 13, Pl. 4, Fig. 17; Coniproetus (Bohemiproetus): Snajdr 1980:Pl. 6, Figs 5, 6, 14; Lieberman 1994:Fig. 9.3) demonstrates that this feature is not an infallible indicator of relationships. Characters cited by Basse (2002) as excluding Australian species of Proetus from Rhenocynproetus also distinguish P. nemus; these include the large size of the lateral occipital lobes and weaker outer edge of the eye socle. Proetus nemus possesses (plesiomorphic) features considered by Basse (2002) to more generally distingish Proetus from Rhenocynproetus, such as a less inflated glabella, the lateral occipital lobes wider than the base of the glabella, terrace lines developed on the dorsal as well as lateral extent of the cranidial border, and the well developed librigenal spine. The presence of a pair of posterior border spines on the hypostome (Fig. 3d) is shared with Proetus (e.g. Whittington and Campbell 1967:PI. 1, Fig. 17; Schrank 1972:Pl. 4, Fig. 7), including P. talenti, but is likely symplesiomorphic (Adrain 1997). Order CORYNEXOCHIDA Kobayashi, 1935 Suborder SCUTELLUINA Hupé, 1953 Family STYGINIDAE Vogdes, 1890 Genus DENTALOSCUTELLUM Chatterton, 1971 Type species Dentaloscutellum hudsoni Chatterton, 1971; by original designation. Dentaloscutellum hudsoni Chatterton, 1971 Fig. 4a-1 Dentaloscutellum hudsoni Chatterton, 1971:12-22, Pl. 1, Figs 1-24, Pl. 2, Figs 1-24, Pl. 3, Figs 1-12, Pl. 24, Fig. 15, Text-figs 4-5. Material 4 cranidia, 29 librigenae, 1 hypostome, | thoracic segment, 5 fragmentary pygidial margins. 184 Discussion This species was fully described based on specimens from the Receptaculites Limestone near Taemas (Chatterton 1971). The Brogans Creek material is considered to be conspecific, the only possible difference being slightly more numerous cranidial tubercles (Fig. 4b, c) than in the type material. Order LICHIDA Moore, 1959 Family LICHIDAE Hawle and Corda, 1847 Subfamily TROCHURINAE Phleger, 1936 Genus ACANTHOPYGE Hawle and Corda, 1847 Type species Acanthopyge leuchtenbergii Hawle and Corda, 1847; by subsequent designation of Reed (1902). Subgenus JASPERIA Thomas and Holloway, 1988 Type species Acanthopyge (Mephiarges) bifida Edgell, 1955; by original designation. Acanthopyge (Jasperia) bifida Edgell, 1955 Fig. 4j-t Acanthopyge (Mephiarges) bifida Edgell, 1955:138; Chatterton, 1971:30-41, Pl. 6, Figs 1-24, Pl. 7, Figs 1-27, Pl. 8, Figs 1-17, Text-figs 8-10. Material 7 cranidia, 1 rostral plate, 7 librigenae, 3 hypostomes, | thoracic segment, 2 pygidia. Discussion The Brogans Creek specimens are indistinguishable from those described from Wee Jasper (Edgell 1955) and Taemas (Chatterton 1971). The species was fully described by Chatterton (1971), rendering description of the Brogans Creek material unnecessary. A few specimens are illustrated (Fig. 4j-t) in support of the conspecificity of the collections. Order PHACOPIDA Salter, 1864 Suborder CALYMENINA Swinnerton, 1915 - Family CALYMENIDAE Milne Edwards, 1840 Genus STHENAROCALYMENE Siveter, 1977 Type species Sthenarocalymene lirella Siveter, 1977; by original designation. Proc. Linn. Soc. N.S.W., 125, 2004 G.D. EDGECOMBE AND A.J. WRIGHT Figure 4. a-i, Dentaloscutellum hudsoni Chatterton, 1971. Scale bars Imm. a, AMF 124725, librigena, dorsal view; b-d, AMF 124726, cranidium, dorsal, anterior and lateral views; e, AMF 124727, librigena, dorsal view; f, AMF 124728, librigena, ventral view; g, AMF 124729, fixigena, dorsal view; h, AMF 124730, incomplete pygidium, ventral view; i, AMF 124731, incomplete pygidium, ventral view. j-t, Acanthopyge (Jasperia) bifida Edgell, 1955. Scale bars 1 mm. j, AMF 124732, rostral plate, ventral view; k, AMF 124733, cranidium, dorsal view; I-m, AMF 124734, cranidium, dorsal and anterior views; n, AME 124735, librigena, dorsal view; o-q, AMF 124736, pygidium, lateral, dorsal and ventral views; r, AMF 124737, librigena, ventral view; s-t, AMF 124738, hypostome, ventral and dorsal views. Proc. Linn. Soc. N.S.W., 125, 2004 | 185 EARLY DEVONIAN TRILOBITES FROM N.S.W. Sthenarocalymene sp. Material Two cranidial fragments, one fragmentary librigena. Discussion A few calymenid cephalic fragments indicate the presence of a species lacking a buttress between the fixigena and L2. On this basis the material is assigned to Sthenarocalymene, the non-buttressed calymenid in many Australian Lower Devonian faunas [see Sandford (2000) for discussion of this genus, its synonym Apocalymene Chatterton and Campbell, 1980, and Gravicalymene Shirley, 1936]. The Brogans Creek material may be identical with S. quadrilobata (Chatterton, 1971), which co-occurs with the other taxa described herein in the lower Receptaculites Limestone at Locality T’ of Chatterton (1971), but specific identity requires better specimens. ACKNOWLEDGEMENTS Alex Cook (Queensland Museum) assisted AJW in the field, and processed and picked much of the material studied here. Yongyi Zhen (Australian Museum) photographed the specimens and assembled the plates, and processed several blocks of limestone. 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Suggestions for a revised classification of trilobites. Geological Magazine, New Series 2, 407-496, 538-545. Talent, J.A., Mawson, R. and Simpson, A.J. (2000). Silurian to Early Carboniferous (Tournaisian) platform-slope sequences in eastern Australia: recent advances in stratigraphic alignments. Geological Society of Australia, Abstracts 61, 114-120. Thomas, A.T. and Holloway, D.J. (1988). Classification and phylogeny of the trilobite order Lichida. Philosophical Transactions of the Royal Society of London 32, 179-262. Vogdes, A.W. (1890). A bibliography of Paleozoic Crustacea from 1698 to 1890 including a list of North American species and a systematic arrangement of genera. Bulletin of the United States Geological Survey 63, 1-177. 187 EARLY DEVONIAN TRILOBITES FROM N.S.W. Whittington H.B. and Campbell, K.S.W. (1967). Silicified Silurian trilobites from Maine. Bulletin of the Museum of Comparative Zoology 135, 447-483. Zhou Z.-q., Siveter, D.J. and Owens, R.M. (2000). Devonian proetid trilobites from Inner Mongolia. Senckenbergiana lethaea 79, 459- 499. 188 Proc. Linn. Soc. N.S.W., 125, 2004 A New Species of the Henicopid Centipede Dichelobius (Chilopoda: Lithobiomorpha) from Southeastern Australia and Lord Howe Island GREGORY D. EDGECOMBE Australian Museum, 6 College Street, Sydney, NSW 2010 (greged @ austmus.gov.au) Edgecombe, G.D. (2004) A new species of the henicopid centipede Dichelobius (Chilopoda: Lithobiomorpha) from southeastern Australia and Lord Howe Island. Proceedings of the Linnean Society of New South Wales 125, 189-203. The genus Dichelobius Attems, 1911, based on D. flavens Attems, 1911, from the southwest of Western Australia, has its only other previously assigned species in New Caledonia and Chile. The Tasmanian type species of the monotypic Tasmanobius Chamberlin, 1920, is regarded as a member of Dichelobius. Dichelobius giribeti n. sp. represents the genus in eastern mainland Australia (southeastern New South Wales, the Australian Capital Territory, and northeastern Victoria) and on Lord Howe Island. Dichelobius bicuspis Ribaut, 1923, is widely distributed in New Caledonia. Manuscript received 1 March 2003, accepted for publication 8 January 2004. KEYWORDS: Anopsobiinae, Chilopoda, Dichelobius, Henicopidae, Lithobiomorpha. INTRODUCTION The subfamily Anopsobiinae is a group of minute centipedes (Chilopoda) in the predominantly southern temperate family Henicopidae. Anopsobiinae is distributed chiefly in the Southern Hemisphere, with species described from Patagonian Argentina and Chile (Silvestri 1899, 1909a-b; Verhoeff 1939; Chamberlin 1962), the Falkland Islands (Eason 1993), New Zealand (Silvestri 1909a; Archey 1917, 1937), New Caledonia (Ribaut 1923), Tasmania (Chamberlin 1920), New South Wales (Edgecombe 2003), southwest Western Australia (Attems 1911), and the Cape region of South Africa (Attems 1928). Four Gondwanan genera have been named: Anopsobius Silvestri, 1899, Catanopsobius Silvestri, 1909b, Dichelobius Attems, 1911, and Tasmanobius Chamberlin, 1920. Four additional anopsobiine genera, all monotypic, occur in the Northern Hemisphere, namely Anopsobiella Attems, 1938, Ghilaroviella Zalesskaja, 1975, Shikokuobius Shinohara, 1982, and Rhodobius Silvestri, 1933. In total, 17 species and subspecies of Anopsobiinae have been described. Silvestri (1909a) cited the occurrence of an anopsobiine from Sydney, but formal descriptions of Anopsobiinae in eastern Australia are limited to Tasmanobius relictus Chamberlin, 1920, based upon a single specimen from Tasmania, and Anopsobius wrighti Edgecombe, 2003, from northern New South Wales. Mesibov (1986) indicated the presence of two species of Anopsobiinae in Tasmania. The present study continues a systematic treatment of Anopsobiinae of Australia by documenting a new species of Dichelobius from New South Wales, the Australian Capital Territory, Victoria, and Lord Howe Island (Fig. 1). For electron microscopy, specimens were photographed on a Leo 435VP using a Robinson backscatter detector. Digital images were assembled into plates with Photoshop. Morphological terminology is as summarised by Edgecombe (2001:203), with terminology for the mandible as in Edgecombe et al. (2002:40, Fig. 4). The following abbreviations are used for repositories of specimens examined: AM - Australian Museum, Sydney ANIC — Australian National Insect Collection, Canberra MCZ — Museum of Comparative Zoology, Harvard University, Cambridge, MA MNHN — Museum National d’ Histoire Naturelle, Paris NMW - Naturhistorisches Museum Wien QM — Queensland Museum, Brisbane WAM — Western Australian Museum, Perth. Other abbreviations: Berl., ANIC Berlesate; CBCR, Australian Museum Centre for Biodiversity and Conservation Research; Ck, Creek; Mt, Mountain; NP, National Park; rf, rainforest; SF, State Forest. A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS } Brisbane Lord Howe Island © Canberra ,¢ Melbourne Figure 1. a, southeastern Australia and Lord Howe Island. Inset shows location of map in b, indicating records of Dichelobius giribeti n. sp. (open dots) in New South Wales, the Australian Capital Territory, and Victoria. Collectors: GBM — G.B. Monteith; JFL — J.F. Lawrence; RJB — R.J. Brooks; RWT — R.W. Taylor. Order LITHOBIOMORPHA Pocock, 1902 Family HENICOPIDAE Pocock, 1901 Subfamily ANOPSOBIINAE Verhoeff, 1907 Genus DICHELOBIUS Attems, 1911 Tasmanobius Chamberlin, 1920 n. syn. Type species Dichelobius flavens Attems, 1911; by monotypy. 190 Assigned species Dichelobius relictus (Chamberlin, 1920) n. comb.; D. bicuspis Ribaut, 1923; D. schwabei Verhoeff, 1939; D. giribeti n. sp. Diagnosis Anopsobiinae with spiracle on segments 3, 10 and 12, variably present on segment 14. Discussion The Gondwanan genera Dichelobius, Tasmanobius and Anopsobius share several apomorphic characters relative to Northern Hemisphere Anopsobiinae. These include coxal pores confined to legs 14 and 15, a ventrodistal spur on the prefemur of legs 14 and 15, an elongate longitudinal median furrow on the head shield, the basal article of the female gonopod extended as a short process bearing the spurs, and indistinct scutes on the proximodorsal part of the pretarsal claws (Edgecombe and Giribet 2003). Considering previous concepts of Dichelobius (Attems 1928; Verhoeff 1939; Shinohara 1982), reduced spiracles are the only morphological character that unites its members to the exclusion of Anopsobius as delimited by Chamberlin (1962) and Edgecombe (2003). The Dichelobius distribution of spiracles is shared by the eastern Australian species D. giribeti. The cladistic reliability of a diminished number of segments with spiracles can be questioned because other genera of Anopsobiinae have been diagnosed based on having spiracles confined to segments 3, 10 and 12 (Tasmanobius), 3, 12 and 14 (Rhodobius) or 3 and 10 only (Catanopsobius). However, molecular sequence data provide independent support for a close relationship between D. flavens and D. giribeti, with the implication that their shared spiracle distribution can be considered a synapomorphy (Fig. 2a). Parsimony analysis of five molecular loci as well as combination of the molecular data and morphology unite D. flavens and D. giribeti to the exclusion of Anopsobius species under many explored gap costs and transversion:transition ratios (Edgecombe and Giribet 2003) (Fig. 2c). An alternative relationship between D. giribeti and Anopsobius (Fig. 2b) is discussed below. Verhoeff (1925) cited the presence of a median suture in the maxillipede pleural band as an additional character by which Dichelobius is distinguished from Anopsobius. The presence of a median suture (see Fig. 6j) is a plesiomorphic character, shared with Henicopinae, and is thus not useful for defining Dichelobius as a clade. Tasmanobius relictus Chamberlin, 1920, is considered to be a member of Dichelobius as grouped Proc. Linn. Soc. N.S.W., 125, 2004 G.D. EDGECOMBE Shikokuobius Anopsobius TAS Anopsobius wrighti japonicus Anopsobius neozelanicus Dichelobius Dichelobius giribeti flavens Shikokuobius Anopsobius TAS Anopsobius wrighti japonicus Anopsobius neozelanicus Dichelobius flavens Dichelobius giribeti Gap cost c Tv : Ts cost Figure 2. a, b, alternative cladograms for Anopsobiinae based on combined morphological and molecular data (Edgecombe and Giribet 2003). Character 1, absence of spiracles on segment 8; character 2, short posteroventral spine on pretarsal claw; c, summary of 12 analyses for combined morphological and molecular data with different gap costs (gap:substitution = 1:1, 2:1, 4:1) and transversion:transition costs (1:1, 2:1, 4:1, infinity). Black squares, parameters that resolve cladogram a (Dichelobius monophyletic); white squares, parameters that resolve cladogram b (Dichelobius paraphyletic); grey square, cladograms a and b of equal length. Proc. Linn. Soc. N.S.W., 125, 2004 herein (with Tasmanobius consequently being a junior subjective synonym of Dichelobius). Tasmanobius relictus was described as having spiracles on segments 3, 10, and 12, as in Dichelobius. Mesibov (1986) suggested that a widespread Tasmanian anopsobiine species (Anopsobiine sp. 2 of Mesibov 1986) may be Tasmanobius relictus, and that species closely resembles Dichelobius giribeti. The holotype and sole type specimen of T. relictus (MCZ 14533) is in poor condition, and lacks locality data more specific than Tasmania, making the identification of any other specimen as this species problematical. The description by Chamberlin did not note a spiracle on segment 14 which is present in the Tasmanian Dichelobius, though this is not obvious in contracted specimens, as noted by Mesibov (1986). A spiracle being absent on segment eight in T. relictus and the colour being “nearly chestnut’ (Chamberlin 1920) make it probable that this species is identical with the Tasmanian Dichelobius (=Anopsobiinae sp. 2 of Mesibov 1986) rather than the northwestern Tasmanian Anopsobius (=Anopsobiinae sp. 1 of Mesibov 1986), which has a spiracle on segment 8 and is more orange-yellow than orange-brown. Accordingly, the name Dichelobius relictus (Chamberlin, 1920) is applied to Anopsobiinae sp. 2 of Mesibov (1986). Attems’ (1928:74) key to anopsobiine genera followed Chamberlin’s (1920) in distinguishing Dichelobius and Tasmanobius based on the former having a 1-jointed tarsus 13 and the latter a 2-jointed tarsus 13. This distinction is inconsistent with the referral of D. bicuspis, which has a 2-jointed tarsus 13 (even fide Attems 1928:77). The supposed difference between these species seems to be nothing more than a terminological difference in what constitutes a ‘Soint’, since D. flavens, D. bicuspis and D. relictus are, upon direct comparison, identical with respect to the segmentation of leg 13. All have a distinct articulation on the tarsus of leg 13, though it is less flexed than is the articulation on leg 14. Other ambiguities concerning Attems’ description and illustrations of Dichelobius flavens have plagued previous interpretations of the genus, and exaggerated differences between D. flavens and other species. Interpretation of D. flavens is based on examination of syntypes from Lion Mill (WAM), Freemantle and Eradu (NMW), and large new collections from the southwest of Western Australia (AM, ANIC, WAM). Dichelobius bicuspis and D. schwabei were distinguished from D. flavens by the first two species having two coxal pores on legs 14 and 15 in the female, versus a single pore on each of the coxae in D. flavens. This cannot be upheld, since large females of D. flavens characteristically have two 191 A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS coxal pores on both legs 14 and 15. Attems’ (1911:157, Fig. 10) described and figured a single spur on the female gonopod in D. flavens, which Ribaut (1923) and Verhoeff (1939) cited as a distinction from the pair of spurs in D. bicuspis and D. schwabei, respectively. Large specimens of Dichelobius flavens resemble congeners (and indeed all other Anopsobiinae) in having a pair of spurs. The specimen drawn by Attems, with a single spur and single coxal pore, is typical of immature stadia of all Dichelobius species (see Archey 1937:pl. 23, fig. 6, for a comparable stage in Anopsobius neozelanicus). Ribaut (1923:27) distinguished D. bicuspis by its plumose setae along the length of the inner margin of the distal article of the telopodite of the first maxilla versus only three plumose setae confined to the distal end of this article in D. flavens (Attems 1911:Fig. 3). Either Attems’ drawing is erroneous or else the illustrated al/ Figure 3. Pretarsal claws in Anopsobiinae. a, Dichelobius relictus (Chamberlin, 1920). Leg 14, posterior side. b, c, Dichelobius flavens Attems, 1911. Leg 14, posterior and anterior sides. d, Anopsobius neozelanicus Silvestri, 1909. Leg 14, posterior side. e, f, Shikokuobius japonicus (Murakami, 1967). Leg 13, posterior and anterior sides. Scales 10 jum except b, 5 um. 192 Proc. Linn. Soc. N.S.W., 125, 2004 G.D. EDGECOMBE specimen is anomalous, because D. flavens has plumose setae all along the inner margin of this article, the same as D. bicuspis (Ribaut 1923:Figs. 30, 31) and other congeners. Certain characters of the pretarsus (claws) conflict with the monophyly of Dichelobius as grouped herein. Dichelobius flavens (Fig. 3b, c) and D. bicuspis differ from D. giribeti (Fig. 8b) and D. relictus (Fig. 3a) in having a long, needle-like spine (=’’sensory spur” of Eason 1964:Fig.486) originating ventrally on the posterior side of the main claw. In the latter two species, the posteroventral spine is short, and a short spine is shared by species of Anopsobius, such as A. neozelanicus Silvestri, 1909a (Fig. 3d) and A. wrighti (Edgecombe 2003:Figs.30, 31). The short spine appears to be apomorphic within the Gondwanan group of Anopsobiinae (i.e., a clade composed of Anopsobius + Dichelobius) because the Japanese anopsobiine Shikokuobius japonicus resembles Dichelobius flavens and D. bicuspis in possessing a greatly elongated posteroventral spine (Fig. 3e, f). The cladogram implied by this character, in which D. giribeti is more closely related to Anopsobius than to D. flavens (Fig. 2b), is retrieved under several parameter sets for combined morphological and molecular data (Fig. 2c). This cladogram would favour the assignment of D. giribeti to another genus. Should this topology find further support from additional data, Tasmanobius Chamberlin, 1920, could be rediagnosed to receive D. giribeti. A rediagnosed concept of that genus might emphasise the shared 14-15 antennal articles, short pretarsal posteroventral spine, absence of a distal spinose projection on the tibia of leg 12, and lack of spiracles on segments 5 and 8. Key to Dichelobius species la. Dental margin of maxillipede coxosternite lacking median notch ....... schwabei Verhoef, 1939 [Chile] 1b. Dental margin of maxillipede coxosternite with median notch...... 2 2a. 14-15 (usually 15) antennal articles; pretarsus with short posteroventral spine, not more than one- eighth length of main claw (Fig. 8b)...... 3 2b. 17 antennal articles; pretarsus with needle-like posteroventral spine nearly as long as main claw (Fig. 3c)..... 4 3a. Spiracle absent on segment 14...... giribeti n. sp. [southeastern Australia, Lord Howe Island] 3b. Spiracle present on segment 14...... relictus Chamberlin, 1920 [Tasmania] Proc. Linn. Soc. N.S.W., 125, 2004 4a. Tibia of leg 12 with short, blunt distal projection...... flavens Attems, 1911 [Western Australia] Ab. Tibia of leg 12 with spinose distal projection...... bicuspis Ribaut, 1923 [New Caledonia] Dichelobius giribeti n. sp. Dichelobius sp. Edgecombe, 2004:Fig. 38A. Dichelobius sp. ACT. Edgecombe and Giribet, 2003:Figs. 1-3. Etymology For Gonzalo Giribet, my collaborator in henicopid phylogeny, who sequenced DNA from this species. Diagnosis Dichelobius usually with 15 antennal articles; head pale orange, tergites orange-yellow; four to six (most commonly five) teeth on each dental margin of maxillipede; spiracle lacking on segment 14; two coxal pores on legs 14 and 15 in females, one or two pores on both legs in males; short posteroventral spine on pretarsus. Type material : Holotype: AM KS 82628, female (Fig. 4b), Badja SF, NSW, Peters Rd, 36°08’52"S 149°32’09"E, J. Tarnawski and S. Lassau, 13.11.1999; length of body 5.1 mm. Paratypes, all from type locality, same collection: AM KS 82629, male (Fig. 4c), KS 82630, male (Fig. 5b-e), KS 82631, female (Figs. 6a-g, 7a, b, d, h, j-l, 8k), KS 82632, female (Fig. 81, j,n), KS 82633, male (Fig. 81), KS 82634, 10 females, 1 male. Other material NSW: AM KS 82635, Kanangra-Boyd NP, Empress Fire Trail turnoff, 33°59’S 150°08’E, M. Gray, G. Hunt and J. McDougall, 27.11.1976, Eucalyptus pauciflora; AM KS 82636, female (Figs 4a, 5a), KS 82637, female (Fig. 8b, e), KS 82638, male (Fig. 61, j), Monga SF, NSW, Link Rd, 35°34’04"S 149°54’14"E, R. Harris and H. Smith, 16.11.1999; AM KS 82639, Buckenbowra SF, Macquarie Rd, 70 m S from junction with Milo Rd, 35°38’ 15"S 149°53’27"E, 1020 m, L. Wilkie and R. Harris, 16.11.1999; AM KS 82640, Tallaganda SF, South Forest Way, 35°42’50"S 149°32’20"E, J. Tarnawski and S. Lassau, 15.11.1999; AM KS 82641, Dampier SF, Coomerang Rd, 36°04’01"S 149°54°57"E, R. Harris and H. Smith, 11.11.1999; AM KS 82642, Badja SF, Wiola Ck Fire Trail, 36°05.56’S 149°35.09’E, J. Tarnawski and S. Lassau, 13.11.1999; AM KS 82643, Badja SF, Burkes 193 A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS Figure 4. a-c, Dichelobius giribeti n. sp. a, AM KS 82636, female, Monga SF, NSW. b, holotype AM KS $2628, female, Badja SF, NSW, terminal segments and gonopods; c, AM KS 82629, male, Badja SF, NSW, terminal segments and gonopods. All scales 100 um. 194 Proc. Linn. Soc. N.S.W., 125, 2004 G.D. EDGECOMBE Rd, 36°10°33"S 149°31°58"E, J. Tarnawski and S. Lassau, 13.11.1999; AM KS 82644, Badja SF, Burkes Rd, approx. 1.3 km E from junction with Peters Rd, 36°10.55’S 149°31.97’E, 992 m, J. Tarnawski and S. Lassau, 13.111.1999; AM KS 82645, Bodalla SF, 300 m along Reservoir Link Rd from junction with Big Rock Rd, 36°07.25’S 150°2.82’E, 121 m, L. Wilkie and R. Harris, 09.11.1999; AM KS 82646, Bodalla SF, Orange Ridge Rd, 36°16’55"S 149°53’31"E, R. Harris and H. Smith, 12.iii.1999; AM KS 82647, Wadbilliga NP, 9.6 km N on Bumberry Ck Fire Trail, 36°14.33’S 149°33.60’E, 1059 m, L. Wilkie and R. Harris, 13.11.1999. ANIC (ex. Berl. 855), Kanangra-Boyd NP, W Morong Creek, 33°58’S 150°04’E, 1200 m, L. Hill, 03.x.1982; ANIC (ex. Berl. 829), Kanangra-Boyd NP, Kanangra Brook and Rocky Spur, 34°00’S 150°06’E, L. Hill, 20.11.1982, closed forest; ANIC (ex. Berl. 852) Twin Falls, 14 km SE Moss Vale, 34°39’S 150°28’E, 600 m, L. Hill, 11.vii.1982; ANIC (ex. Berl. 663), Pigeon House Range via Nerriga, 35°02’S 150°08’E, J.C. Cardale, 22.xi.1979; ANIC (ex. Berls 2, 18, 34, 78A, 206A, 222, 246, 468, 657, 851), Clyde Mt, 35°33’S 149°57°E, 500-c. 800 m, various collections 1966-1982, dry sclerophyll, wet sclerophyll, rf; ANIC (ex. Berl. 877), 2 km N Monga, 35°34’S 149°56’E, M.S. Harvey, 18.1x.1983, wet sclerophyll; ANIC (ex. Berl. 594), Monga, 35°35’S 149°55’E, JFL and T. Weir, 10.11.1978, wet sclerophyll; ANIC (ex. Berl. 739), Tallaganda SF, 7 km ENE Captains Flat, 35°34’S 149°31’E, W. Allen, 29.viii.1981; ANIC (ex. Berl. 1069), Kioloa SF, 35°35’S 150°18’E, JFL and N. Lawrence, 4-5.111.1986; ANIC (ex. Berl. 927), Milo Forest Preserve, 1.6 km S Monga, 35°36’S 149°55’E, L. Hull, 25.xii.1983; ANIC (ex. Berl. 218), 8.8 km ESE Captains Flat, 35°38’S 149°31’E, 940 m, RWT, 10.1.1970, dry sclerophyll; ANIC (ex. Berl. 891), Rosedale, 35°49’S 150°14’E, R.J. Moran, 20.xi.1983, eucalypt litter; ANIC (ex. Berl. 933), Kosciusko NP, 1 km ENE Mt Sunrise, 36°22’S 148°29’E, L. Hill, 411.1984; ANIC (ex. Berl. 935), Kosciusko NP, 4 km NNE Mt Perisher, 36°22’S 148°29’E, L. Hill, 4.11.1984; ANIC (ex. Berl. 10), Brown Mt, 36°36’S 149°23’E, c. 3000 ft., RWT, 5.1.1967, wet sclerophyll; ANIC (ex. Berl. 20), Brown Mt, c. 2800 ft., RWT and R.J. Bartell, 30.11.1967, rf; ANIC (ex. Berl. 24), Brown Mt, 2500- 3000 ft., RWT and R.J. Bartell, 11.1v.1967; ANIC (ex. Berl. 41), Brown Mt, Rutherford Creek, 2700 ft., RWT and RJB, 9.x1i.1967, rf; ANIC (ex. Berl. 42), Brown Mt, c. 3000 ft., RWT and RJB, 9.x1i.1967, rf. ACT: ANIC (ex. Berl. 283), Black Mt, eastern slope, 35°16’S 149°06’E 750 m, J. Simmons, 26.v.1970, dry sclerophyll; ANIC (ex. Berl. 228), - Uriarra to Piccadilly Circus, 35°19’S 148°51’E, 700 m, RWT, 27.i.1970, dry sclerophyll; ANIC (ex. Berl. Proc. Linn. Soc. N.S.W., 125, 2004 225), Uriarra to Piccadilly Circus, 35°20’S 148°50’E, 500 m, RWT, 16.1.1970, wet sclerophyll; ANIC (ex. Berl. 231), Uriarra to Piccadilly Circus, 35°20’S 148°50’E, 1000 m, RWT, 16.1.1970, wet sclerophyll; ANIC (ex. Berl. 999), Wombat Creek, 6 km NE Piccadilly Circus, 35°19’S 148°51’E, 750 m, JFL, T. Weir and M.-L. Johnson, 30.vi.1984, open forest; ANIC (ex. Berl. 1001), Piccadilly Circus, 35°22’S 148°48’E, 1240 m, JFL, T. Weir and M.-L. Johnson, 30.vi.1984, subalpine eucalypt litter; ANIC (ex. Berl. 1000), Blundells Creek, 3 km E Piccadilly Circus, 35°22’S 148°50°E, 850 m, JFL, T. Weir and M.-L. Johnson, 30.vi.1984, open forest; ANIC (ex. Berl. 821), Brindabella Range, Franklin Rd, N end Moonlight Hollow, 2 km SW Bulls Head, 35°24’S 148°48’E, M.S. Harvey and R.J. Moran, 3.iv.1983; ANIC (ex. Berl. 926), Ginini Flat, 2 km NE Mt Ginini, 35°31’S 148°46’E, 1580 m, L. Hill, 20.viii.1983; ANIC (ex. Berl. 659), Mt Ginini, 35°32’S 148°46’E, 1660 m, JFL and T. Weir, 16.x.1979; ANIC (ex. Berl. 1068), 1 km S Mt Ginini, 35°33’S 148°46’E, JFL, 11.xi.1986; ANIC (ex. Berl. 704, 705), 1 km N Mt Gingera, 35°33’S 148°47°E, A.A. Calder, 18.11.1981; ANIC (ex. Berl. 26), Mt Gingera, 35°34’S 148°47’E, c. 5500 ft., E.B. Britton, 13.iv.1967, wet sclerophyll; ANIC (ex. Berl. 50), Mt Gingera, summit, E.B. Britton and Misco, 19.vii.1967; ANIC (ex. Berl. 661), Mt Gingera, E.C. Zimmerman, 20.xi1.1979; ANIC (ex. Berl. 830, 831), Mt Gingera, 1620-1700 m, L. Hill, 6.111.1982; ANIC (ex. Berl. 1084), Snowy Flat Creek, 0.5 km NE Mt Gingera, 35°35’S 148°47’E, A.A. Calder, 28.vi.1988. VIC: ANIC (ex. Berl. 1045), Cobb Hill, 14 km SE Bonang, Goonmirk Ra, 37°18’S 148°50’E, JFL and N. Lawrence, 24.x1.1985. LORD HOWE ISLAND: AM KS 35592, NE area of Mt Gower summit, moss forest near campsite, 31°35.2’S 159°04.7°E, 855 m, M.R. Gray, 12-15.11.1971; AM KS 35589, creek crossing above Boat Harbour, 31°33.5"’°S 159°05.5’E, 60 m, M.R. Gray, 8.11.1971; AM KS 82998, female (Figs. 6h, 8a, d, f, g), KS 82999, male (Figs. 6k, 1, 0, 7g, m, 8c, h, m), KS 83000, male (Figs. 6m, n, 7c, e, f, i), west end of Mt Gower summit on south edge, 31°35.32’S 159°04.2’E, I. Hutton, 15.v.2001; AM KS 84206- 84233, additional localities/samples on Mt Gower, AM KS 84234-84237, four localities on Mt Lidgbird, I. Hutton and CBCR, 2000-2002; AM KS 84238, North Hummock, trail to Intermediate Hill, 31°32754"S 159°04’58"E, CBCR, 3.xii.2000, mixed rf; AM KS 84239, western slope of Malabar Ridge, 31°30°57"S 159°03’31"E, CBCR, 24.xi.2000, broad megaphyllous closed sclerophyll forest; AM KS 84240, Transit Hill, 31°32’01"S 159°04’40"E, I. Hutton, 14.iv.2002; AM KS 84241, Little Island, below Far Flats, 31°34’08"S 159°04’32"E, I. Hutton, 10.viii.2001, under Ficus 195 A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS columnaris. Description Length (anterior margin of head shield to telson) up to 6.6 mm; length of head shield up to 0.7 mm; leg 15 33-40% length of body. Colour: head shield and maxillipede pale orange; antenna and most tergites orange-yellow, T14 and tergite of intermediate segment deeper orange; legs 1-13 pale yellow to pale orange, legs 14 and 15 may be deeper orange. Head shield (Fig. 5a) smooth, of equal length and width, slightly wider than T1, median notch contributing to biconvex anterior margin; longitudinal median furrow incised to transverse suture, about one- third length of head shield; posterior two-thirds of region distal to antennocellar suture desclerotised; setae on head shield arranged with bilateral symmetry, four larger pairs anterior to antennocellar suture, ten pairs behind suture, including four evenly spaced submarginal pairs; head shield lacking posterior and lateral borders. Antenna 27-32% length of body, 2.5-3.3 times length of head shield, composed of 14 or (usually) 15 articles; basal two articles enlarged, most articles in distal half moniliform, sclerotised part generally of subequal length and width; ultimate article about twice length of penultimate. Basal article bearing about a dozen sensilla microtrichoidea proximally on dorsal side (Fig. 6a). Trichoid sensilla arranged in three whorls per article; one or occasionally two curved, Figure 5. a-e, Dichelobius giribeti n. sp. a, AM KS 82636, female, Monga SF, NSW, head shield, maxillipede segment and T1; b-e, AM KS 82630, male, legs 12-15, Badja SF, NSW. All scales 100 um. 196 Proc. Linn. Soc. N.S.W., 125, 2004 G.D. EDGECOMBE Figure 6. Dichelobius giribeti n. sp. Scanning electron micrographs. a-g, Badja SF, NSW; h, k-o, Mt Gower, Lord Howe Island; i, j, Monga SF, NSW. a-g, AM KS 82631, female. a, cluster of sensilla microtrichoidea on proximal part of antenna, dorsal side, scale 10 tm; b, clypeus, scale 50 [im; c, posterior part of clypeus and labrum, scale 50 im; d, labral margin, scale 10 tm; e, antennal articles 10-13, dorsal side, scale 30 jum; f, basiconic sensillum at anterior edge of antennal article 12, dorsal side, scale 5 um; g, tip of terminal antennal article, scale 10 um. h, AM KS 82998, female, dental margin of maxillipede, scales 100 um, 30 um. i, j, AM KS 82638, male, dental margin and ventral view of maxillipede, scales 50 tum, 100 pm. k, I, o, AM KS 82999, male. k, porodont, scale 10 jum. 1, dental margin of maxillipede, scale 50 uum. 0, anterior angle of telopodite of first maxilla, scale 10 um. m, n, AM KS 83000, male, telopodite of -maxillipede and detail of tarsungulum, showing sensilla coeloconica, scales 50 jum, 5 um. Proc. Linn. Soc. N.S.W., 125, 2004 | 197 A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS digitiform sensilla near anterior edge on dorsomedial side of a few, variable antennal articles (Fig. 6e); four or five articles with a single, short, fusiform sensillum at anterior edge on dorsal side (Fig. 6f), most consistent on articles 11, 12 and 14; digitiform and fusiform sensilla sometimes cooccur on a single article (article 7 or 9); ultimate article with cluster of 8 or 9 trichoid sensilla at apex, one or two curved, digitiform sensilla behind apical cluster (Fig. 6g). Clypeus with apical cluster of three setae on ventral side near lateral margin, single seta medially (Fig. 6b); transverse band of four setae in front of labrum, outer pair slightly to distinctly smaller than inner (Fig. 6c); transverse seta projecting from sidepiece; labral margin moderately concave where cluster of 7-13 bristles projects; bristles with numerous short, spine-like projections along lateral margins and on ventral surface along their lengths (Fig. 6d). Tomosvary organ large, longitudinally ovate, outer edge at lateral margin of cephalic pleurite (Fig. 8k), Maxillipede (Figs 6h-n): coxosternal width across dental margin 39-44% maximum width; lateral margin flexed inward at base of dental projections and less convergent than against posterior part; each dental margin convex, usually with 5+5, 445 or 5+4 teeth, sometimes 4+4, 6+5, 5+6 or 6+6; inner tooth smaller than others, its apex well posterior to base of outer tooth; median notch varying from broadly V-shaped (Fig. 6h) to deeply parabolic (Fig. 61); porodont of similar length and thickness to largest coxosternal setae, its socket at posterolateral edge of outermost tooth (Fig. 6k); setae relatively sparsely, fairly evenly scattered on coxosternite; tarsal and pretarsal parts of tarsungulum of about equal length (Fig. 6m). Dorsal and ventral sides of tarsungulum with several sensilla coeloconica (Fig. 6n). Bands of pleural collar separated by longitudinal median suture (Fig. 6)). Mandible: Six curved aciculae (Fig. 7j), all with many (up to 18) short, blunt denticles along both margins (Fig. 7i) on distal half to two-thirds. Four paired teeth, dorsal three with accessory denticle field delimited by deep groove; dorsalmost tooth and basal part of second and third teeth composed of densely tuberculate rhomboid and polygonal scales (Fig. 71), becoming denticulate near furry pad (Fig. 7m). Fringe of branching bristles terminates against dorsalmost acicula (Fig. 7f); ventralmost bristles in fringe with flattened bases lacking spines, distal two-thirds with short spines along both margins and on outer face; bristles multifurcating at their distal tips, with three or. four spines that are longer and thicker than those more proximally (Figs 7f, k); more dorsal bristles gradually become more uniformly spinose to their broader bases, with more numerous distal spines (Fig. 7k), grading 198 into wide scales that form a nearly continuous double- fringe of hair-like spines, each scale composed of a narrow outer fringe and a wider inner fringe, each with 12-15 spines per scale (Fig. 71); fringe terminates at edge of dorsalmost tooth, against a large, smooth scale that separates dentate lamina from furry pad (Fig. 7m). Furry pad composed of a few scales with distal spines and cluster of six or seven mostly simple, elongate spines. First maxilla: sternite indistinctly delimited from coxa (Fig. 7a), short, wide. Coxal projections tapering, with rounded apex bearing four or five simple setae; one small seta along inner margin near base of coxal projection. Telopodite strongly delimited from coxal projection; basal article of telopodite with single marginal seta anterolaterally or lacking setae; distal article with one or two setae near outer margin, anterior angle terminating as a long, stout spine; entire inner margin fringed with row of six or seven plumose setae (Fig. 7b), paired in posterior part of row, with slender branchings along more than half of their length (Fig. 7c); five shorter simple setae inserting near bases of plumose setae on ventral side; anterior plumose setae fringed on dorsal side by a few elongate spines. Second maxilla: anterior margin of coxa gently concave; band of four or five small setae across anterior part of coxa. Inner edge of tarsus with a row ~ of five or six brush-like setae with abundant, slender branchings nearly to their bases (Fig. 7d, h). Claw composed of up to five digits with concave, scoop- like inner surfaces (Fig. 7g); large, curved medial digit with furrows or sutures running along its length (Fig. 7e); outer digits shorter, separated from medial digit by a slender, spine-like digit. Tergites smooth, all with rounded posterior angles, lacking projections; Tl about 85% width of widest tergite (TT10 or 12). Posterior margins of TT1, 3, 5 and 7 transverse (Fig. 4a); TT8, 10 and 12 gently concave; TT9, 11, 13 and 14 transverse to weakly concave; tergite of intermediate segment transverse or gently concave, posterior angle rounded. Two or three moderately long setae on lateral margins of long tergites, usually with short setae between these; posterior margins of tergites fringed with four to twelve setae, generally more abundant on more posterior segments (maximal number typically on T13); setae on inner part of long tergites include transverse band of up to six setae across anterior third, two or three pairs in two bands behind this. Legs 12-15 (Fig. 5b-e) with length ratios 1: 1.1 : 1.3-1.4: 1.7. Leg 15 basitarsus 85-115% length of distitarsus (Fig. 5e); basitarsus 70-75% length of tibia; tibia 2.9-3.4 times longer than maximal width, basitarsus 3.4-4 times, distitarsus 5.2-5.7 times. Proc. Linn. Soc. N.S.W., 125, 2004 G.D. EDGECOMBE Figure 7. Dichelobius giribeti n. sp. Scanning electron micrographs. Scales 10 um except where indicated. a, b, d, h, j-l, AM KS 82631, female, Badja SF, NSW; c, e-g, i, m, Mt Gower, Lord Howe Island. a, ventral view of first maxillae, scale 50 tm; b, distal article of telopodite of first maxilla; d, h, tarsus and claw of second maxilla, scales 10 um, 30 jum; j, aciculae; k, 1, ventral and dorsal parts of fringe of branching bristles on mandible. c, e, f, i, AM KS 83000, male. c, plumose setae on inner margin of telopodite of first maxilla; e, claw of second maxilla; f, aciculae and fringe of branching bristles on mandible; i, aciculae. g, m, AM KS 82999, male. g, claw of second maxilla, scale 10 im; m, dorsalmost tooth of mandible and furry pad. Basitarsus 90% length of distitarsus on leg 14 (Fig. surface with fine longitudinal grooves and ridges like 5d). Coxal projections on leg 15 tapering (in ventral _ those on pretarsal claws. Trochanter of leg 15 with view) at about 25-30 degrees; terminal spine with small ventrodistal spur (Figs 5e, 8h). Prefemur of legs distinct (Fig. 8e) or indistinct (Fig. 8i) basal joint, its 14 and 15 with large ventrodistal spur; leg 15 spur : Proc. Linn. Soc. N.S.W., 125, 2004 | 199 A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS with basal width about 25% maximum width of prefemur (Fig. 4b). Sharp distal spinose projections on tibiae of legs 1-11, absent on legs 12-15. Two tarsomeres of leg 13 defined by distinct constriction in width and weak articulation without flexure; articulation between tarsomeres stronger on leg 14. Setae fairly evenly distributed on all podomeres along leg, tarsal setae only slightly more slender than those on prefemur-tibia; proximo-distal gradient in setal thickness enhanced on legs 14 and, especially, 15, with distinctly thickened prefemoral setae, including on dorsal side of leg. Anterior and posterior accessory claws present on all legs, 25-40% length of main claw (Fig. 8a, b); accessory claws with closely-spaced linear ridges on their surface except for pitted proximoventral part separated by a shallow suture (Fig. 8c). Main claw curved, subdivided by sutures; deepest sutures define an elongate scute on both lateral sides of claw, proximal end of this scute at about distal end of shorter accessory claw; large pore or pair of pores at proximal end of scute on both sides of leg (Fig. 8c); strong suture extends from lateral pore across ventral surface of main claw (Fig. 8d), defining proximal end of an elongate, triangular ventral scute (Fig. 8g). Proximal part of main claw densely pitted; on ventral side of claw, ornament changes abruptly at suture delimiting lateral scute, becoming linear grooves and ridges as on accessory claws (Fig. 8d), with these lineations well developed on lateral scute and along length of claw on dorsal side; change from pitted to linear ornament gradual on dorsal side of claw, with pits irregular proximally, becoming aligned as rows of pits, then linear grooves. Pair of distally-directed spines proximoventrally, at distal end of a curved suture (Fig. 8d); larger spine not more than not more than one-eighth length of main claw, with tiny subsidiary spine at its base (Fig. 8b). Coxal pores: on legs 14 and 15; 2,2/2,2 in females (Fig. 4b), 1,1/1,1 in small males, either 1,1/ 1,1 or 2,2/2,2 (Fig. 4c) in large males, occasionally one and two pores on opposing sides of either leg or 1,2/1,2; pores round, separated by less than their diameter when paired; inner pore often smaller than outer pore in male, inner pore sometimes larger than outer pore in female. Female (Fig. 4b): Sternite of segment 15 gently convex posteromedially, fringed by a submarginal setal band that extends along entire posterolateral and posterior margin; several setae scattered on inner part of sternite. Posterior margin of first genital sternite moderately embayed between . gonopod articulations, sternite bearing 6-11 setae. Gonopod with pair of spurs at terminus of a short (Fig. 8n) to moderately long (Fig. 8e, f) projection; bases of spurs nearly touching each other; inner spur 200 substantially shorter and narrower than outer spur, both bullet-shaped, pointed (Fig. 8n); four or five setae on basal article of gonopod, three large setae on second article, one large seta on third (Fig. 8j); second and third articles variably with one and two smaller setae, respectively, on ventromedial face (Fig. 8n); claw simple. Male (Fig. 4c): Posterior margin of sternite 15 evenly convex; 10-13 setae fringing margin of sternite, 10-12 additional setae scattered over its ventral surface; first genital sternite entire medially, bearing 6-12 setae aligned in two imprecisely-defined transverse rows; gonopod bearing two or three setae on first article, two on second article, none or one on third article, which grades into long, flagelliform terminal process, up to 80% length of rest of gonopod (Fig. 81); terminal process bearing numerous slender spines proximally (Fig. 8m). Larvae: five larval stadia (ANIC Berl. 18 and 231) identified as LO-LIV by comparison to limb development in other Lithobiomorpha (Table 1). LI with 11 antennal articles; LII-LIV all with 14 articles. LII and LIII with 2+2 teeth on dental margin of maxillipede; LIV with 3+3 teeth. Discussion Specimens from Lord Howe Island resemble those from the Australian mainland in all meristic characters and in fine detail. Intrapopulation variation is observed with respect to the number of teeth on the maxillipede coxosternal margin, the depth of the median notch in the maxillipede coxosternite (relatively shallow in Fig. 6h, relatively deep in Fig. 61), the concavity of the posterior margins of the short tergites, and the length of the spur-bearing process on the female gonopod. Samples vary in the frequency with which large males have either one or two coxal pores on legs 14 and 15 (usually two in Lord Howe specimens versus one in the large sample from Clyde Mountain, NSW, but also two in large specimens from the type locality and in the Brindabella Range, e.g., Piccadilly Circus, Mt Gingera and Mt Ginini). Distinction from other congeners is indicated in key above. Dichelobius relictus and D. giribeti are consistently distinguished by the presence of a spiracle on segment 14 in the former, and D. relictus is generally a deeper brown colour. The two species share minute details of mandibular and maxillary structure, indeed to the extent that description of the mouthparts for D. giribeti serves for D. relictus as well. The early larval stadia of Dichelobius giribeti differ in detail from those of Lithobiidae and Henicopinae (see Table 1) with respect to limb development. Segmentation of LO is matched by Proc. Linn. Soc. N.S.W., 125, 2004 G.D. EDGECOMBE Pk A Figure 8. Dichelobius giribeti n. sp. Scanning electron micrographs. a, c, d, f-h, m, Mt Gower, Lord Howe Island; b, e, Monga SF, NSW; i-l, n, Badja SF, NSW. a, d, f, g, AM KS 82998, female. a, pretarsus of leg 14, scale 10 um. d, g, ventral views of pretarsus of leg 14, scales 10 um; f, gonopods, scale 30 Lum. b, e, AM KS 82637, female. b, pretarsus of leg 14, posterior view, scale 10 jum; e, ventrolateral view of first genital sternite and gonopods, scale 100 tum. c, h, m, AM KS 82999, male. c, pretarsus of leg 15, detail of anterior accessory claw, scale 5 um; h, prefemur of leg 15, anterior side, scale 100 um; m, terminal process on gonopod, scale 10 pum. i, j, n, AM KS 82632, female. i, leg 15 coxal process, scale 30 um; j, n, lateral and ventral views of gonopod, scales 50 um, 10 um. k, AM KS 82631, female, cephalic pleurite with Témoésvary organ, scale 50 um. 1, AM KS 82633, male, gonopod, scale 30 Lm. Proc. Linn. Soc. N.S.W., 125, 2004 , 201 A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS Table 1. Comparison of limb development in larval stadia of Lithobiomorpha. Modified from Andersson (1979:Table II), adding data for Dichelobius giribeti. Lamyctes emarginatus Lamyctes coeculus Dichelobius Lithobius 8 spp. Lamyctes coeculus, but larval stadium LI has a unique combination of half-developed legs and limb-buds in D. giribeti. Segmentation of stadia LII-IV is as in other lithobiomorphs. Four larval stadia identified by Eason (1993) for Anopsobius macfaydeni have seven, eight, ten and twelve pairs of legs, the last three obviously being LII-LIV. The taxonomic significance of the distinction between six- and seven-legged first larval stages in Dichelobius giribeti and Anopsobius macfaydeni is unclear without additional data for Anopsobiinae. Dichelobius bicuspis Ribaut, 1923 Dichelobius bicuspis Ribaut, 1923:24, Figs. 27-34. Dichelobius bicuspis: Wiirmli, 1974:526. Material NEW CALEDONIA: PROV. NORD: AM KS 83001, 1 female, 1 male, Mt Panié, nr summit, 20°34’S 164°46’E, 1500 m, C. Burwell, 9.xi.2001, rf; MNHN, | female, 1 larval stadium LIV, Mt Panié, 20°34’53"S 164°45’°38"E, 1350 m, J. Chazeau, A. & S. Tillier, 18.xi.1986, wet Agathis forest; QM S60653, 1 female, Pic d’Amoa, N slopes, 20°58’S 165°17’E, 500 m, GBM, 10.xi.2001, rf; QM S60654, 1 male, Me Maoya, summit plateau, 21°12’S 165°20’E, 1400 m, GBM, 12.xi.2002, rf. PROV. SUD: MNHN, 3 females, Mt Do, 21°45°37"S 165°59’33"E, 840 m, A. & S. Tillier & Monniot, 2.iv.1987, wet Araucaria forest; QM S60655, 1 male, Mt Humboldt refuge, 21°53’S 166°24’°E, 1300 m, GBM, 7-8.xi.2002, rf; AM KS - 83002, | male, R Bleue, Pourina Track, 22°04’S 166°38’E, 900 m, GBM, 18.xi.2001, rf; AM KS 83003, | male, Mt Ouin, 22°01’S 166°28’E, 1100 m, GBM, 9.xi.2002, rf; AM KS 83004, 1 female, 1 male, QM 202 giribeti $60656, 1 male, Mt Mou base, 22°05’S 166°22’E, 200 m, GBM, 30.x.2001, 15.xi.2001, rf; MNHN, 3 females, 1 juvenile, Riviére Bleue, 22°06’13"S 166°39’16"E, 160 m, A. & S. Tillier, 1.viii.1986-30.1v.1987; QM S$60657, 1 male, Mt Koghis, 22°11’S 166°01’E, 750 m, GBM, 29.xi.2000, rf; AM KS 83005, 1 female, Yahoué, 22°12’S 166°30’E, 100 m, GBM, 4.xi.2001, rf. Remarks Dichelobius bicuspis was based on a few specimens from Mt Humboldt (the type locality) and Mt Canala, New Caledonia, with Wtirmli (1974) adding a record at Nékliai. New collections are listed above to indicate that the species has a more widespread distribution. ACKNOWLEDGEMENTS I thank Suzanne Bullock (Scientific Interface) and Sue Lindsay and Yongyi Zhen (Australian Museum) for assistance with illustrations and electron microscopy, and the referees for useful suggestions. Geoff Monteith (Queensland Museum) and Jean-Paul Mauriés (Museum National d’ Histoire Naturelle, Paris) kindly provided material from New Caledonia. Collection study was hosted and loans were arranged by Matthew Colloff (Australian National Insect Collection), Gonzalo Giribet and Laura Leibensperger (Harvard University), and Mark Harvey and Julianne Waldock (Western Australian Museum). Verena Stag] (Naturhistorisches Museum Wien) is thanked for arranging loan of C. Attems’ types. Proc. Linn. Soc. N.S.W., 125, 2004 G.D. EDGECOMBE REFERENCES Andersson, G. (1979). On the use of larval characters in the classification of lithobiomorph centipedes (Chilopoda, Lithobiomorpha). In ‘Myriapod Biology’ (Ed M. Camatini) pp. 73-81. (Academic Press: London). Archey, G. (1917). The Lithobiomorpha of New Zealand. Transactions and Proceedings of the New Zealand Institute 49, 303-318. Archey, G. (1937). Revision of the Chilopoda of New Zealand. Part 2. Records of the Auckland Institute and Museum 2, 71-100. Attems, C. (1911). Myriopoda exkl. Scolopendridae. Die Fauna Siidwest-Australiens. 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(1986). “A Guide to Tasmanian Centipedes.’ (Privately published: Zeehan, Tasmania). Ribaut, H. (1923). Chilopodes de la Nouvelle-Calédonie et des Iles Loyalty. In “Nova Caledonia. Recherches scientifique en Nouvelle-Calédonie et aux Iles Loyalty. A. Zoology 3(1)’ (Eds F. Sarasin and J. Roux) pp. 1-79. (C.W. Kreidel’s Verlag: Berlin, Weisbaden). Shinohara, K. (1982). A new genus of centipede of the subfamily Anopsobiinae (Henicopidae, Chilopoda). Proceedings of the Japanese Society of Systematic Zoology 24, 41-46. Silvestri, F. (1899). Contribucion al estudio de los quilopodos chilenos. Revista Chilena de Historia Natural 3, 141-152. Silvestri, F. (1909a). Contribuzioni alla conoscenza dei Chilopodi. III. Descrizione di alcuni generi e specie di Henicopidae. Bolletino del Laboratorio di Zoologia generale e agraria, Portici 4, 38-50. Silvestri, F. (1909b). Descrizioni preliminari di vari Arthropodi, specialmente d’America. V. Nuovi genere di Henicopidae (Chilopoda). Rendiconti della R. Accademia dei Lincei 18, 270-271. Silvestri, F. (1933). Nuovi contributi alla conoscenza della fauna delle isole italiane dell’Ageo. I. Descrizione di un nuovo genere di Chilopodo Henicopino. Bolletino del Laboratorio di Zoologia generale e agraria, Portici 27, 58-60. Verhoeff, K.W. (1925). Systematik. C. Supplement zu den Lithobiomorpha. In “Klassen und Ordnungen des Tier-Reichs’ (Ed H.G. Bronn) pp. 595-603. (Akademische Verlagsgesellschaft: Leipzig). Verhoeff, K.W. (1939). Von Dr. G. H. Schwabe in Chile gesammelte Isopoda terrestria, Diplopoda und Chilopoda. Archiv fiir Naturgeschichte, Zeitschrift fur Wissenschaftliche Zoologie Abteilung B 8, 301-324. Wiirmli, M. (1974). Ergebnisse der Osterreichischen Neukaledonien-Expedition 1965. Chilopoden. Annalen des Naturhistorisches Museums in Wien 78, 523-533. Zalesskaja, N.T. (1975). New genera and species of Chilopoda (Lithobiomorpha) from central Asia and Far East. 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Wits TH) TAN LAS SMa : © yee she, SRC hud, 29 PIS b, Ol ae TM: renal et : j 7 ba ; ett ¥ > . \ *" " : bs c é : { mae, {} vi ay sw iy &’ a 4 ’ “ Mi rf , + i . “4 # » be - y 204 } : és or Pa i 7 : he ‘ Fi) x ¥ = A Survey of Ectoparasite Species on Small Mammals During Autumn and Winter at Anglesea, Victoria Haylee J Weaver'’ and John G Aberton’. ‘School of Ecology and Environment, Deakin University, Geelong VIC 3217; *Present address: School of Biological and Environmental Sciences, Central Queensland University, Rockhampton QLD 4702 (h.weaver @cqu.edu.au) Weaver, H.J. and Aberton, J.G. (2004). A survey of ectoparasite species on small mammals during autumn and winter at Anglesea, Victoria. Proceedings of the Linnean Society of New South Wales 125, 205-210. A survey of the ectoparasites of small native mammals was carried out between April and August 2002, in heathlands surrounding Anglesea, Victoria. Antechinus minimus, A. agilis, Rattus lutreolus, R. fuscipes, Sminthopsis leucopus and Isoodon obesulus were the dominant host mammal species examined. A total of 921 ectoparasites were collected and identified as five flea species, seven mite species and two species of tick. Isoodon obesulus was found to have the highest ectoparasite species richness, with eleven of the fourteen species present; while S. leucopus displayed the lowest ectoparasite species richness with only three species found on the hosts examined. The flea Pygiopsylla hoplia was the only ectoparasite species in this study to have a distribution across all host mammal species. A new distribution record was made for a Haemaphysalis tick species. Manuscript received 16 October 2003, accepted for publication 8 January 2004. KEYWORDS: Anglesea, ectoparasites, host specificity, marsupials, rodents, species richness. INTRODUCTION The main groups of ectoparasitic arthropods encountered on Australian mammals include fleas (order Siphonaptera), mites (order Acariformes), ticks (order Parasitiformes) and lice (order Phthiraptera). These ectoparasites, as a group, have evolved specialised piercing and sucking mouthparts, designed for the extraction of blood from a host, with the degree of host specificity displayed by ectoparasites varying amongst species (Kemp et al. 1982; Dunnet and Mardon 1991). Many species of ectoparasites are of considerable medical and veterinary importance. Fleas are capable of transmitting various rickettsial, filarial and protozoan diseases (Dunnet and Mardon 1991), and ticks can transmit pathogenic filariae, bacteria, protozoa, rickettsiae and viruses to wild and domestic animals and humans (Obenchain and Galun 1982; Aeschlimann 1991). Previous research on ectoparasites in the Anglesea region has been limited to flea surveys as a precursor to the introduction of myxomatosis (Dunnet and Mardon 1991) and calicivirus (F. Bartholomaeus pers. comm.), and basic natural history of ticks (Roberts 1970). Ectoparasites also negatively impact on the health of both domestic and wild animals through large infestations, which are of importance in management considerations of rare or endangered small mammal species present at Anglesea as increases in host densities may increase ectoparasite loads. The objective of this study was to survey ectoparasite species on small native mammals near Anglesea, Victoria because an awareness of the ectoparasites is important for the potential transmission of disease to humans, domestic animals and livestock. It is also of interest to the general ecology of small mammals in the region. METHODS Ectoparasites were removed from small mammals trapped at two sites at Anglesea, Victoria (Fig. 1). The sites chosen for study were the Eumeralla Scout camp (38°24’0"S, 144°12’36”E) and Bald Hills Road (38°23’24’S, 144°8’24”E) at the Alcoa Lease. Both sites were selected using knowledge that they contained many host species, and these species were all relatively abundant. The Eumeralla Scout camp consisted of a coastal tea tree, Leptospermum continentale shrub layer, with plants varying from 20 centimetres to over two metres in height and Eucalyptus obliqua at a height of over two metres ECTOPARASITES ON SMALL MAMMALS Bald Hills Rd Eumeralla Alcoa Lease boundary —» Anglesea Great Ocean Rd Airey's Inlet Figure 1. Location map of study area. forming the canopy. The site was a flat open heathland with woodland dispersed through it, and a swamp consisting mainly of Gahnia radula and also L. continentale. The Bald Hills Rd site on the Alcoa Lease was situated on a slope of approximately 30° in a southwesterly direction. The heathland was dominated by L. continentale, Epacris impressa, Conospermum mitchelli, L. myrsinoides, Platylobium obtusangulum and G. radula were the main species present in the understorey. Stands of Eu. willisi and Banksia marginata were present at the study site. Trapping of small mammals was carried out during Autumn and Winter 2002, due to the study being an honours project requiring completion during an academic year. Trapping sessions of three nights each were carried out at Eumeralla in June (3-6.6.02), July (8-11.7.02) and August (6-9.8.02), with a total of 131 mammals captured over the three sessions and at the Bald Hills Rd site in April/May (29.4-2.5.02), July (22-25.7.02) and August (19-22.8.02), with 161 206 captures recorded. Any previously trapped mammals captured again in following sessions were re-examined for ectoparasites and were counted accordingly. Fifty aluminium Elliott traps (32 x 9 x 10 cm) were placed in transects across the Eumeralla site. The site at Bald Hills Rd consisted of 100 traps set in a grid pattern (100 m x 100 m) at ten metre intervals. Traps were baited using a rolled oats, peanut butter and honey mix and were cleared within three hours of sunrise. Upon capture, mammals were transferred from the trap into a lightweight mesh bag, identified, ear notched for identification purposes, weighed, sexed and inspected for ectoparasites. As ticks were physically attached to the host, they were removed using fine forceps to grip the tick as close to the host’s skin as possible and flipping it over to remove the tick while leaving the mouthparts intact. Fleas and mites were removed by ruffling the host’s pelage with fingers in order to dislodge the ectoparasites, or the host was combed using Licemeister combs or animal flea combs. Numbers of each ectoparasite taxa were recorded from each mammal and all ectoparasites collected were placed in labelled containers of 70% ethanol. Identification of fleas; mites and ticks were carried out using descriptions provided by Dunnet and Mardon (1974), Domrow (1987, 1991) and. Roberts (1970) respectively. A linear regression on host mammal body weight and ectoparasite species richness was carried out using log transformed data. RESULTS AND DISCUSSION A total of 292 individual mammals were trapped over 1350 trap nights from the two sites. The host mammals trapped included Antechinus minimus (74), A. agilis (69), Sminthopsis leucopus (4) (Dasyuriomorphia: Dasyuridae), Isoodon obesulus (10) (Peramelemorphia: Peramelidae), Rattus fuscipes (50) and R. lutreolus (85) (Rodentia: Muridae). Examination of 296 host mammals yielded 364 fleas and 557 acari (mites and ticks) in total. From this, five flea species were identified, along with seven Proc. Linn. Soc. N.S.W., 125, 2004 H.J. WEAVER AND J.G. ABERTON mite species and two tick species. Of these, two species of mites were unable to be identified to species level; these were referred to by their family names as unidentified Laelapidae and _ unidentified Trombiculidae. Table 1 shows the number of examinations of each host mammal species and the species of ectoparasites removed from the host species. The most common host examined for ectoparasites was Rattus lutreolus, with 85 examinations and the host examined least was Sminthopsis leucopus with only four examinations. Sminthopsis leucopus is an uncommon mammal in the Anglesea area. Lunney (1995) states although it has a wide distribution throughout southern Australia, it prefers sparse ground to forage, whereas the sites in this study had very dense ground cover. Figure 2 shows I. obesulus as having the greatest ectoparasite species richness and S. leucopus the smallest. A significant linear association was found between host weight and ectoparasite species richness (MS=0.098, F=7.966, df=1, P=0.048) with 64.32% of the variation in ectoparasite species richness accounted for by mean body weight of the hosts. This is consistent with previous studies showing that host body size determines ectoparasite species richness (Kuris et al. 1980, cited in Stanko et al. 2002). Another factor that can influence ectoparasite species richness is the social behaviour of the host. Stanko et al. (2002) found that higher host densities generally equated to lower species richness on individuals, possibly because of anti- parasitic behaviours such as grooming. As bandicoots have a reputation of ‘pugnacious behaviour between conspecifics’ (Lobert 1990) and indicate a low social tolerance (Thomas 1990), it could be that the bandicoots examined in this study had:a higher species richness of ectoparasites and a higher abundance of each species in part due to a combination of larger body size and lack of social grooming. The most common ectoparasite collected was the flea Pygiopsylla hoplia, which was recorded on every host mammal species. According to Dunnet and Mardon (1974), P. hoplia is the most commonly collected Australian species of flea. It has a distribution across Australia, excluding the Northern Territory, and has been recorded on many species of peramelids, dasyurids and rodents (Dunnet and Mardon 1974). In contrast, Stephanocircus dasyuri was mostly recorded on I. obesulus, and occasionally on A. minimus. The similar foraging nature of both these mammal species may be the reason why this species of flea was not recorded on any other hosts. Macropsylla hercules was only recorded on Rattus spp. and I. obesulus, perhaps due to the size of the host animals, as this flea is very large. Macropsylla hercules is commonly collected Proc. Linn. Soc. N.S.W., 125, 2004 from various native Rattus species from southern Australia (Dunnet and Mardon 1974). The other species of flea collected, Acanthopsylla rothschildi rothschildi and Bibikovana rainbowi appeared to display little host specificity, as they were recorded from the majority of the host species. Host specificity for acarine ectoparasites collected varied. The highly host specific Androlaelaps marsupialis was only found between the groove of the tibia and fibula on the hind legs of I. obesulus where grooming is difficult (pers. obs.). Similarly, Mesolaelaps anomalus was recorded only on I. obesulus. In contrast, the trombiculid mites and the tick Ixodes tasmani showed a broad host range, being found on all host species except for S. leucopus and A. minimus respectively. The trombiculids were found most frequently inside the ears of hosts during this study, but can be found on any exposed skin including legs, feet and tails (pers. obs.). Trombiculid mites are parasitic during their larval stage and later live in the soil as free living adults (Domrow 1962). One small infestation was recorded in the pouch of a female /. obesulus, and it has been suggested that larval trombiculids occurring in the pouches of A. minimus can directly infest any pouch young present (B. Wilson, Deakin University, pers. comm.). Ixodes tasmani is a common species of tick with a distribution widespread across southern Australia with a wide range of hosts (Roberts 1970). The species of Haemaphysalis collected from I. obesulus was identified as H. humerosa, but differences in the spiracular plate between the Anglesea specimens and specimens from known populations in Queensland have been observed. An alternative identification is H. ratti. Further research is being carried to provide a definite identification of the specimens (I. Beveridge, University of Melbourne, pers. comm., D. Kemp, CSIRO, pers. comm). Other ectoparasitic arthropods were collected from host mammals studied. Lice (Phthiraptera, species unknown) were collected from R. lutreolus on three occasions; but were not observed on any other host mammals examined. The rove beetle species Myotyphlus jansoni (Coleoptera: Staphylinidae) was collected from Rattus lutreolus on five occasions. However, M. jansoni is not an obligate ectoparasite. Myotyphlus jansoni has only been recorded on a very small number of individual native Rattus species previously (Hamilton-Smith and Adams 1966). The beetles are usually collected near the anus or tail (as they were in this study) and have also been recorded in bat guano in a cave near Warrnambool, Victoria; thus it may be assumed that the beetles feed on the excreta of the rats, which is not strictly an ectoparasitic 207 ECTOPARASITES ON SMALL MAMMALS Host species No. of mammals Body weight (g) Siphonaptera Number Acari Number examined (Mean + SD) Antechinus minimus 74 50+ 13 Pygiopsylla hoplia 36 Andreacarus tauffliebi 5 (swamp antechinus) Acanthopsylla rothschildi 5 Mesolaelaps sminthopsis 1 : Bibikovana rainbowi 1 Androlaelaps telemachus 5 Stephanocircus dasyuri 3 Trombiculidae 17 Antechinus agilis 67 31+9 Pygiopsylla hoplia 11 Mesolaelaps sminthopsis 4 (agile antechinus) Acanthopsylla rothschildi 39 Androlaelaps telemachus 3 Trombiculidae 8 Ixodes tasmani 6 Rattus fuscipes 50 106 + 23 Pygiopsylla hoplia 4 Androlaelaps telemachus 1 (bush rat) Bibikovana rainbowi 5 Trombiculidae 23 Macropsylla hercules 7 Ixodes tasmani I Rattus lutreolus 85 95 +21 Pygiopsylla hoplia 3 Andreacarus tauffliebi 50 (swamp rat) Bibikovana rainbowi 7 Mesolaelaps sminthopsis 2 Macropsylla hercules 6 Trombiculidae 12 Ixodes tasmani 2 unidentified Laelapidae 10 Sminthopsis leucopus 4 prea 3) Pygiopsylla hoplia 1 Ixodes tasmani 1 (white footed dunnart) Acanthopsylla rothschildi 1 Isoodon obesulus 10 400 + 173 Pygiopsylla hoplia 90 Androlaelaps marsupialis 10 (southern brown bandicoot) Acanthopsylla rothschildi 2 Mesolaelaps anomalus 55 Bibikovana rainbowi 2 Mesolaelaps sminthopsis 317 Macropsylla hercules 1 Trombiculidae 5 Stephanocircus dasyuri 140 Ixodes tasmani 2 Table 1. Species of ectoparasites collected from host mammals. Haemaphysalis sp. Proc. Linn. Soc. N.S.W., 125, 2004 208 H.J. WEAVER AND J.G. ABERTON Total parasite spp. y = 0.017x + 5.722 R’ = 0.643 0 50 100 150 200 250 300 350 400 Mean body weight (g) Figure 2. Relationship between ectoparasite species richness and body weight of host mammals. SI = Sminthopsis leucopus, Aa = Antechinus agilis, Am = Antechinus minimus, R\ = Rattus lutreolus, Rf = Rattus fuscipes, lo = Isoodon obesulus. relationship (Hamilton-Smith and Adams 1966; Lawrence and Britton 1991). The ectoparasite species collected during this study were all considered to be common throughout the region (Roberts 1970; Dunnet and Mardon 1974) and all are theoretically able to transmit pathogens to animals or humans. Generally, fleas are known to be intermediate hosts for the cosmopolitan rodent tapeworm, Hymenolepis diminuta and the canine tapeworm Dipylidium canium, along with being able to transmit various filarial, rickettsial and protozoan pathogens, however native Australian fleas have not been found to contribute epizootics in the field (Dunnet and Mardon 1991). Ixodes tasmani has been recorded as an intermediate host of various rickettsiae, including Rickettsia australis, the organism which causes Queensland tick typhus (Campbell and Domrow 1974, cited in Cavanagh 1999). Haemaphysalis ticks are vectors of Coxiella burnetii (Q fever), in bandicoots and macropods and domestic livestock (Kettle 1995). Therefore it is recommended that care be taken when in areas where ticks are present, especially at the Proc. Linn. Soc. N.S.W., 125, 2004 Eumeralla Scout Camp where groups of scouts may come into contact with ticks while carrying out activities in the area. In conclusion, it was found that there was a significant relationship between ectoparasite species richness and body weight of host mammal species. There was no difference in the species of ectoparasites collected from both study sites, except for M. jansoni, which was only found on R. lutreolus at the Bald Hills Rd site. As there have been no other studies carried out of this type in the region, it is recommended that a study over a longer time frame be carried out in order to accurately assess seasonal variations of ectoparasite numbers. ACKNOWLEDGEMENTS The authors are grateful to F. Bartholomaeus (South Australian Research and Development Institute) and M. Shaw (University of Queensland) for additional flea and mite identification respectively, and Dr Ian Beveridge (University of Melbourne) and Dr David Kemp (CSIRO) 209 ECTOPARASITES ON SMALL MAMMALS for assistance with the identification of Haempahysalis sp. We appreciate Philip Barton’s comments on the manuscript. The study formed part of the principal author’s BEnvSc Honours thesis and was carried out in accordance with Deakin University Animal Ethics committee guidelines and conducted under Victorian Department of Natural Resources and Environment wildlife permit no. 10001759. REFERENCES Aeschlimann, A. (1991). Ticks and Disease: Susceptible - hosts, reservoir hosts and vectors. In ‘Parasite-host associations, coexistence or conflict?’ (Eds C.A. Toft, A. Aeschlimann and L. Bolis) pp. 148-156. (Oxford University Press: Oxford). Campbell, R.W. and Domrow, R. (1974) Rickettsioses in Australia: Isolation of Rickettsia tsutsugamushi and R. australis from naturally infected hosts. Transactions of the Royal Society of Tropical Medicine and Hygiene 68, 397-402. Cavanagh, F.A. (1999) The common marsupial tick, Ixodes tasmani, and factors that influence the degree of infestation of the common brushtail possum, Trichosurus vulpecula. BSc Honours thesis, University of Sydney, Sydney. Domrow, R. (1962). The mammals of Innisfail. 2: Their mite parasites. Australian Journal of Zoology 2, 268-307. Domrow, R. (1987). Acari Mesostigmata parasitic on Australian vertebrates: An annotated checklist, keys and bibliography. Invertebrate Taxonomy 1, 817-948. Domrow, R. (1991). Acari Prostigmata parasitic on Australian vertebrates: An annotated checklist, keys and bibliography. Invertebrate Taxonomy 4, 1283-1376. Dunnet, G.M. and Mardon, D.K. (1974). A monograph of Australian fleas (Siphonaptera). Australian Journal of Zoology Supplementary Series 30, 1- DPS - Dunnet, G.M. and Mardon, D.K. (1991). Siphonaptera. In ‘The insects of Australia: a textbook for students and research workers Vol II.’ (CSIRO) pp. 705- 716. (Melbourne University Press: Melbourne). Hamilton-Smith, E. and Adams, D.J.H. (1966). The alleged obligate ectoparasitism of Myotyphus jansoni (Mathews) (Coleoptera: Staphylinidae) Journal of the Entomological Society of Queensland 5, 44-45. Kemp, D.H., Stone, B.F. and Binnington, K.C. (1982). Tick Attachment and Feeding: Role of the Mouthparts, Feeding Apparatus, Salivary Gland Secretions and the Host Response. In ‘Physiology of Ticks’. (Eds F.D. Obenchain and R. Galun) pp.119-168. (Pergamon Press: Oxford). Kettle, D.S. (1995) ‘Medical and Veterinary Entomology (2 edn.)’. (CAB International: United Kingdom). Kuris, A.M., Blaustein, A.R. and Ali6, J.J. (1980). Hosts as islands. American Naturalist 116, 570-586. Lawrence, J.F. and Britton, E.B. (1991). Coleoptera. In “The insects of Australia: a textbook for students and research workers Vol II.’ (CSIRO) pp. 543- 683. (Melbourne University Press: Melbourne). Lobert, B. (1990). Home range and activity period of the Southern brown Bandicoot (/soodon obesulus) in a Victorian heathland. In “Bandicoots and Bilbies.’ (Eds J.H. Seebeck, P.R. Brown, R.L. Wallis and C.M. Kemper) pp. 319-325. (Surrey Beatty and Sons: Sydney). Lunney, D. (1995). White-footed Dunnart. In “The mammals of Australia.’ (Ed. R. Strahan) pp.143- 145. (Australian Museum/Reed New Holland: Sydney). Obenchain, F.D. and Galun, R. (1982). Preface. In ‘Physiology of ticks.’ (Eds F.D. Obenchain and R. Galun) pp. vii-ix. (Pergamon Press: Oxford). Roberts, F.H.S. (1970). ‘Australian ticks.” (CSIRO: Melbourne). Stanko, M., Miklisova, D., Gotiy de Bellocgq, J. and Morand, S. (2002). Mammal density and patterns of ectoparasite species richness and abundance. Oecologica 131, 289-295. Thomas, L.N. (1990). Stress and population regulation in Isoodon obesulus (Shaw and Nodder). In ‘Bandicoots and bilbies.’ (Eds J.H. Seebeck, P.R. Brown, R.L. Wallis and C.M. Kemper) pp. 335- 343. (Surrey Beatty and Sons: Sydney). Proc. Linn. Soc. N.S.W., 125, 2004 Occurrence and Conservation of the Dugong (Sirenia: Dugongidae) in New South Wales Smmon ALLEN!, HELENE MArsH23 AND AMANDA HopcGson23 1Graduate School of the Environment, Macquarie University, NSW 2109; *School of Tropical Environment Studies and Geography, James Cook University, Townsville, Qld 4811; 3CRC Reef Research Centre, PO Box 772, Townsville, Qld 4810 Allen, S., Marsh, H. and Hodgson, A. (2004). Occurrence and conservation of the dugong (Sirenia: Dugongidae) in New South Wales. Proceedings of the Linnean Society of New South Wales 125, 211- 216. Recent sightings of dugongs well beyond the southern limit of their accepted range (~27°S) on the Australian east coast prompted a review of past records of dugongs and their current conservation status in New South Wales. While archaeological analyses have identified bones of Dugong dugon in Aboriginal middens at Botany Bay (~34°S) and colonial records indicate stranded animals as far south as Tathra (~36.5°S), there were no verified sightings of live individuals in NSW waters for some years; however, five separate sightings of individuals and pairs were documented in the austral summer of 2002/03 in estuaries on the NSW central coast (~32-33.5°S). Itis suggested that conditions such as warm sea temperatures and low rainfall (promoting seagrass growth) may be facilitating explorative ranging south by dugongs. The IUCN lists dugongs as ‘vulnerable’ at a global scale and they are also classified ‘vulnerable’ under the Threatened Species Conservation Act NSW 1995, yet they are not routinely considered in risk assessments for inshore development in this State. Threatening processes such as shark meshing persist. The importance of considering dugongs in future impact assessments for inshore marine and estuarine developments is emphasized. Manuscript received 17 October 2003, accepted for publication 8 January 2004. KEYWORDS: conservation, distribution, dugong, Dugong dugon, risk assessment, sightings, status, vulnerable. INTRODUCTION The dugong (Dugong dugon), along with all other extant Sirenians, is regarded as a shallow water, tropical and sub-tropical species (Martin and Reeves 2002; Rice 1998). Dugongs are thought to be strictly marine, inhabiting the coasts of some 37 countries and territories (Marsh et al. 2002). Despite their widespread distribution, dugong numbers have declined in most of their known range and they are believed to be represented by fragmented, relic populations in most countries. Likely causes for this decline and continuing threats include: large-scale destruction of seagrass as a result of sedimentation, dredging, mining, trawling, and pollution; incidental take as by-catch in commercial and recreational gill and mesh nets as well as shark nets set for bather protection; direct takes from indigenous hunting, and vessel strikes and disturbance (Marsh et al. 1999, 2002; Hodgson 2003). Australian waters are the dugong’s stronghold, where their distribution is described as extending from Shark Bay in Western Australia (25°S) around northern Australia to Moreton Bay in southern Queensland (27°S) (Marsh et al. 2002). Dugongs are a ‘listed marine species’ under the Australian Environment Protection and Biodiversity Conservation Act 1999 (EPBC Act). The EPBC Act reflects Australia’s commitments under various international conventions including the Bonn Convention on the Conservation of Migratory Species of Wild Animals, which lists the dugong on Appendix 2. Dugongs are also considered ‘vulnerable’ under the Threatened Species Conservation Act NSW 1995 and under the Nature Conservation Act Qld 1992. Evidence of a decline in dugong numbers along the urban coast of Queensland (Marsh et al. 2001) led to the establishment of a series of dugong protection areas in some key dugong habitats in Queensland (Marsh et al. 1999; Marsh 2000). No similar protection has been afforded dugongs in NSW, presumably on the assumption that only vagrants of the species range into NSW waters. Dugongs have been considered in some impact assessments for aquaculture developments in NSW (e.g. Anon. 2001a), but not others (e.g. Anon. 2001b). These assessments occurred in the same location, suggesting consideration of dugongs and potential impacts thereon is inconsistent in NSW. DUGONGS IN NEW SOUTH WALES In this paper, we highlight past and present evidence that the dugong’s range on the east coast of Australia extends into NSW waters, including estuaries, when environmental conditions are suitable. Given their conservation status under both international conventions and national acts, we suggest that occasional visitation warrants adherence to the legal obligation of considering dugongs and their preferred habitats in future impact assessments. EARLY RECORDS TO RECENT SIGHTINGS Dugong bones have been found associated with edge-ground hatchet heads in Aboriginal middens near Sydney, indicating that at least small numbers of dugongs have utilized NSW waters for many centuries (Etheridge et al. 1896). In 1799 Flinders described the catching of dugongs by Aborigines in Moreton Bay, southeast Queensland (Mackaness 1979). Aborigines in NSW also caught dugongs in more recent times, Pell eee with bones having been found in middens as far south as Botany Bay in the late 18" Century (Troughton 1928). There are currently two sources of dugong sightings in NSW: the Atlas of NSW Wildlife and records of by-catch from shark meshing supervised by NSW Fisheries. The Atlas of NSW Wildlife yields 83 reports of live, stranded and dead animals for the period 1788 to 2003 (Anon. 2003b; Fig. 1). A significant portion of these reports (63) occurred in late 1992 and throughout 1993. This influx of animals occurred after the loss of 1,000 km? of seagrass from Hervey Bay in southeast Queensland following floods (Preen and Marsh 1995). Two dugongs were caught in NSW shark meshing during this time (Swansea in November 1992 and January 1993). Three earlier captures were also made in shark nets (Bronte in July 1951, Bondi in July 1951, Queenscliff in April 1971) (Krogh and Reid 1996). Only two records of dead and stranded individuals have been reported to the NSW National be Pe set les Figure 1. Past records of dugongs on the NSW coast from 1788 to 2003 (open circles; Anon. 2003b) and dugong sightings in central NSW estuaries during summer 2002/03 (filled circles). 212 Proc. Linn. Soc. N.S.W., 125, 2004 S. ALLEN, H. MARSH AND A. HODGSON 32°11.0° 152 30.27 Wallis Late Oct. Lak 2002 € ‘ort e rt 32°42.8° ~ 152°06.7” 33°20.5” P 10° Jan. ‘| Lak : Po 1* Feb. Brisbane 150°29.8° 32°41.8° 152°03.27 33°30.1° Water 152°20.3’ | beach Kayak tour operator reports dugong/s over seagrass beds within Wallis Lake Dolphin watch operators report two adult dugongs near Manton Bank Recreational travelling seaward out Swansea Channel Dolphin watch operator report dugong/s in upper estuary west of Soldiers Point Resident reports dugong/s off Orange Grove S. Smith, pers. comm. D. Aldritch, pets. comm. fishers report cow-calf pair | B. Roche, pers. comm. D. Aldritch, pers. comm. Anon. 2003b Table 1. Dugong sightings in central NSW estuaries in the austral summer of 2002/2003. Parks and Wildlife Service (NPWS) in the last decade, with no live sightings occurring until late 2002/03. Between late October 2002 and early February 2003, five separate sightings of individuals and pairs within (or swimming out of) central coastal estuaries were reported to NPWS and/or the authors (Table 1; Fig. 1). These occurred along ac. 200km stretch of coastline and we do not know if these sightings include repeat sightings of the same individual(s). SEAGRASS DISTRIBUTION AND WATER TEMPERATURES All the estuaries in which dugongs were sighted are known to support seagrass meadows (Table 2). Dugongs have been recorded eating the seagrasses listed in Table 2, with the exception of Ruppia spp. (Anderson 1986, Marsh et al. 1982, Lanyon et al. 1989). Species of the genus Halophila are preferred. The distribution of dugongs has been reported as being constrained to water temperatures >~18°C (Anderson 1986, 1994: Marsh et al. 1994; Preen et al. 1997). However, the water temperatures at the sites in Table 2 were above this thermal threshold in summer 2002/ 03. DISCUSSION The low abundance of dugongs in NSW waters may be the result of a number of factors Proc. Linn. Soc. N.S.W., 125, 2004 including limited availability of seagrass in the region, relatively low water temperatures during winter months and in open coastal waters between estuary and bay habitats, and/or human pressures. The entire NSW coast supports only 155 km? of seagrass (West et al. 1989), the major portion of which would be Posidonia australis and species of the Zosteraceae family, which are not favoured by dugongs. In relative terms, the amount of seagrass in NSW is much less than the total area of seagrass in Moreton Bay alone (250 km?: Abal et al. 1998) and would contain correspondingly small cover of Halophila spp. Troughton (1928) interpreted historical records as suggesting that dugongs may have occurred in greater numbers in NSW prior to European settlement. It has also been suggested (MacMillan 1955) that dugong populations on the tropical east coast were again beginning to expand into the northern rivers region of NSW. Any expansion of the dugong’s range into NSW waters further south than this region may have been inhibited by the loss of seagrass beds in areas such as Port Macquarie and Botany Bay to anthropogenic influences (Pointer and Peterkin 1996). The dugong observations in 2002/03 (Table 1) were in areas of NSW which have some of the largest seagrass beds, at least two of which include Halophila species — part of the preferred diet of dugongs (Marsh et al. 1982; Table 2). The increasing evidence that individual dugongs embark on movements over many hundreds of kilometres within tropical waters (N. Gales pers. comm; Marsh and Lawler 2001, 2002; Marsh DUGONGS IN NEW SOUTH WALES Estuary (latitude) Seagrass species and approximate area coverage Water temp. (°C) ne Zosteraceae, Posidonia australis, Ruppia and Halophila Wallis Lake (~32.29S) spp. ~30.785km? Port Stephens (~32.78) Halophila spp. ~7,453km? Lake Macquarie (~33.19S) spp. ~13.391km? Brisbane Water (~33.4S) Halophila spp. ~5.490km? Zosteraceae, Posidonia australis and Zosteraceae, Posidonia australis, Ruppia and Halophila Zosteraceae, Posidonia australis and October mean: 18.9 October 2002: 21.0 January mean: 24.1 - February mean: 24.6 January mean: 21.6 \ February mean: 22.1 Table 2. Extent of seagrass meadows and water temperatures at sighting ocations. Sources for seagrass coverage and water temperature data: West et al. (1985) and Anon. (2003a) respectively. Water temperatures are means from 1987-2002, unless otherwise stated. and Rathbun 1990; Marsh et al. 2002) suggests it is possible that dugongs explore and utilize these southern seagrass beds. Warm water temperatures during the summer months of 2002/03 may have encouraged this behaviour. Although only five dugongs have been reported drowned in shark nets in NSW over the last c. 50 years (Krogh and Reid 1996), such deaths are not inconsequential since few dugongs are commonly found south of Moreton Bay. Two of these mortalities coincided with a seagrass dieback event (Preen and Marsh 1995) and further impact on Queensland seagrass beds or increase in water temperature in NSW may see an increase in shark net capture of dugongs off NSW beaches. Such events will highlight negative effects on populations of non-target species, and the efficacy of shark control programs for bather protection in NSW and Queensland will again be called into question (Anon. 2002). The dugong is classified as ‘vulnerable’ at a global scale on the IUCN Red List of Threatened Species. As the only extant species in the family Dugongidae, the extinction of the dugong will result in biodiversity loss at the family and generic levels as well as at the species level. In the light of inconsistencies evident in risk assessments for inshore development in NSW, we re-iterate that dugongs should be considered occasional visitors to NSW coastal waters. Their limited numbers warrant the dugongs’ consideration in future impact assessments for estuarine and inshore marine developments. The estuarine nature of recent sightings suggests that 214 explorative ranging by dugongs is not necessarily limited to strictly marine environments, rather to areas where seagrass beds occur. This also adds weight to the importance of assessing potential impacts on seagrass habitats. ACKNOWLEDGMENTS We gratefully acknowledge all those that provided prompt and unambiguous reports of recent sightings. We would also like to thank Mick Murphy of Hunter Coast Area NSW National Parks and Wildlife Service for providing access to the relevant wildlife database and Jeanine Almany for information on seagrass distribution and water temperatures in NSW. SA was supported by an ARC SPIRT grant, HM and AH by funding from the CRC Reef Research Centre. This manuscript was greatly improved by comments from Robert Williams, John Merrick and two anonymous reviewers. REFERENCES Abal, E.G., Dennison, W.C. and O’ Donohue, M.H. (1998). Seagrasses and mangroves in Moreton Bay. In ‘Moreton Bay and Catchment’ (Eds LR. Tibbetts, N.J. Hall and W.C. Dennison) pp. 269- 278. (University of Queensland: Brisbane). Anderson, P. K. (1986). Dugongs of Shark Bay, Australia - seasonal migration, water temperature, and forage. National Geographic Research 2, 473- 490. Proc. Linn. Soc. 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West, R.J., Thorogood, C.A., Walford, T.R. and Williams, R.J. (1985). An estuarine inventory of New South Wales, Australia. Fisheries Bulletin 2. Department of Agriculture, NSW. 216 Proc. Linn. Soc. N.S.W., 125, 2004 Captures, Capture Mortality, Age and Sex Ratios of Platypuses, Ornithorhynchus anatinus, During Studies Over 30 Years in the Upper Shoalhaven River in New South Wales T.R. GRANT School of Biological, Earth and Environmental Sciences, University of NSW, NSW 2052 Email t.grant@unsw.edu.au Grant, T.R. (2004). Captures, capture mortality, age and sex ratios of platypuses, Ornithorhynchus anatinus, during studies over 30 years in the upper Shoalhaven River in New South Wales. Proceedings of the Linnean Society of New South Wales 125, 217-226. Data collected during a number of studies over a period of 30 years in the upper Shoalhaven River, New South Wales, are presented. A total of 700 individual platypuses (Ornithorhynchus anatinus) were captured during the studies, consisting of 137 juvenile females, 94 juvenile males, 292 adult females and 177 adult males. The overall sex ratios of both adults (1.65:1) and juveniles (1.46:1) were significantly biased towards females. Females were found to live up to 21 years. Very few recaptures of juvenile males made estimates of longevity equivocal, but three individuals were at least 7 years old when last captured. Capture and handling mortality during the various studies was low (0.86%). Sixty-two percent of platypuses marked in the study area were never recaptured, fewer adult males were recaptured than females (36% and 51% respectively) and recaptures of juveniles were much lower than for adults (32% females and 14% males). Recapture data suggest considerable mobility by adults and dispersal by juvenile platypuses along the upper Shoalhaven River and its tributaries. Manuscript received 2 September 2003, accepted for publication 8 January 2004. Keywords: Age, Capture, Marking, Mortality, Ornithorhynchus anatinus, Platypus , Sex Ratio INTRODUCTION Over the past 30 years a number of research projects has been carried out in a study area in the upper Shoalhaven River in New South Wales. Individual projects have included investigation of temperature physiology (Grant and Dawson 1978a,b; Grant 1983; Hulbert and Grant 1983a,b), diet (Faragher et al. 1979), movements and home ranges (Grant 1983, 1992), haematology and pathology (Munday et al. 1998; Whittington and Grant 1983, 1984, 1995; Whittington et al. 2002), lactation and milk composition (Grant et al. 1983, Gibson et al. 1988; Grant and Griffiths, 1992) and population genetics (Gemmell 1994; Akiyama 1999). During these studies, long-term data have been collected on recaptures, capture mortality, longevity and sex ratios of platypuses (Ornithorhynchus anatinus). During the later studies from 1987, the investigation and use of Passive Integrated Transponder (PIT) tags or “microchips” was begun, probably the first time that this marking method was used on a wild mammalian species in Australia (Grant and Whittington 1991). The long-term success of this method in the mark and recapture studies of the platypus is reported below. Collins (1973) tabulated the ages of eight platypuses kept in captivity in a variety of locations, including the Bronx and Budapest Zoos. These ranged from four to 17 years, although anecdotal information from zoos and sanctuaries indicates that the species may survive in captivity for up to 21 years (Whittington 1991). Concerning the longevity of platypuses in the wild, the naturalist Harry Burrell (1927) wrote that “the length of life of the platypus is not known. It is my intention to ring-mark some fully furred young as opportunity offers, and it may be that we shall gain some information on this point at a later date, if these marked individuals are captured”. Burrell did not later report the ages of platypuses he may have “ring- marked”. However, Grant and Griffiths (1992) reported the ages of platypuses marked in the upper Shoalhaven River as being between as much as 4 years for males and 8 for females. Since that report, a further 12 years of research has resulted in the data presented in this paper on the ages and sex ratios in this population of platypuses. Mortality of capture and handling of platypuses in previous studies has not CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN RIVER previously been discussed in the literature and this aspect of the studies is presented in the current paper. METHODS Between June 1973 and January 2004, platypuses were captured in 16 pools in the upper Shoalhaven River in New South Wales using the unweighted “gill” net methods outlined in Grant and Carrick (1974). Until 1987, individuals were marked using stainless steel leg bands (Grant and Carrick (1974) but these were phased out after trials on the use of Passive Integrated Transponder (PIT) tags proved to be successful (Life Chip tags; Destron Fearing Corporation Scanner; Grant and Whittington 1991). Sex was determined using the presence or absence of the adult spur or the morphology of juvenile spurs. Absolute ages were determined from individuals initially captured as juveniles and minimum ages for adults were estimated using the time of loss of the female spur, the morphological changes in the males spur (Temple-Smith 1973; Grant 1995), and subsequent recaptures. Females possessing a spur were categorised as being in their first year of life (O years of age) and males could be assigned to their first or second year of life (0 or 1 year of age). As the females in this area lose the spur between October and December in their first year after emergence from the nesting burrows, any female lacking a spur at first capture was considered to be = 1 year of age (i.e. in their second year of life). It should be noted that two female juvenile platypuses bred in captivity at Taronga Zoo in the 2002/2003 breeding season apparently lost their spurs within only 4 months after emergence from the nesting burrow (Adam Battaglia, Taronga Zoo, pers. comm.). Males with adult spur morphology were considered to be at least in their third year of life, or = 2 years old. Subsequent recaptures permitted minimum ages to be assigned to individuals, beginning with a minimum age at first capture of one year for adult Juvenile Males 94 Females 137 Total 231 Sex Ratio (F:M) 1.46:1 Chi? 8.00 p ** < 0.005 females and two years for adult males (Temple-Smith 1973; Grant and Griffiths 1992; Grant 1995). Recaptures were recorded for animals in all of the 16 pools of the 12.5 km section of the upper Shoalhaven River and 3.9 km of an adjacent creek. However, by 1987 a number of these pools had filled with sand and were no longer netted. By 1993, the previously largest and deepest pool (1 km long x 2-5 m deep) was completely filled with sand and was no longer sampled, although many of the platypuses originally captured in this pool were captured in the pools downstream. From 1988, when PIT tagging had become the predominant method of marking, sampling was mainly restricted to three pools in the Shoalhaven River itself and one in the adjacent creek. In most years after that time these pools were sampled late in the year (mainly December) when lactating animals were most likely to be caught and at the end of summer (mainly February or March) when juveniles had newly emerged (Grant and Griffiths 1992). RESULTS Sex ratios During the studies from June 1973 to January 2004, 700 individual platypuses were captured. Table 1 shows the numbers in each of four age/sex classes. All sex ratios were significantly biased towards females. The ratio of females to males was 1.58 females:1 male (Chi? = 34.87; p < 0.001) for all animals, 1.46:1 for juveniles (Chi? = 8.00; p < 0.01) and 1.65:1 for adults (Chi? = 27.38; p < 0.001). Age Only 45 individuals (41 females and 5 males), first marked as juveniles, were subsequently recaptured. Figure 1 shows the distribution of ages of these individuals at their latest recapture. Two juvenile females were recaptured regularly over periods of 13 and 16 years but were not captured again in 5-6 subsequent years. These animals were assumed to have Adult Total La 271 292 429 469 700 1.65:1 1.58:1 27.38 34.87 ** < 2.001 ** < 0.001 Table 1. Numbers and sex ratios of platypuses captured in the upper Shoalhaven River study area. ** significant at < 0.01 level. 218 Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT ~ Year Juvenile Juvenile Year Adult Adult (Actual) Female _ Male (Minimum) Female Male 0 96 89 0 - - 1 Di 4 1 181 28 2 3 if 2 33 117 3 §) - 3 18 | iu 4 3 - 4 18 9 5 D - 5 10 36 >5 ovens - >5 32 6 Total 137 94 Total 292 177 Table 2. Numbers of platypuses allocated to actual and estimated minimum age categories in the upper Shoalhaven River study area. Actual = ages of animals initially captured as newly-emerged juveniles; Minimum = minimum ages; calculated from years between initial and last capture of individuals first captured as adults. died. However, one was again recaptured and lactating at the end of the study, when her age was 21 years. As indicated in the Methods section, females without spurs are at least in their second year of life (>/=1 year old) and it is possible to attribute males to either their first second or third year of life (O, 1 or 2 years old) based on spur morphology changes. Ages of >/= 2 years could be attributed to 111 female adults caught and subsequently recaptured. Similarly 32 adult males were attributed to the >/= 3 year age category. The distribution of these minimum ages are shown in Table 2 and Fig. 1. While most platypuses caught in the study could only be attributed to the >/= 1-2 year age category, 9 females first captured as juveniles survived between 5-21 years and 32 of those initially captured as adults survived 5-15 years. One juvenile male was subsequently recaptured at 2 years of age but 32 males, first captured as adults, survived to minimum ages of 3-7 years. Recaptures The numbers of juvenile, adult male and adult female platypuses recaptured at least once in the latter 12 years (when most animals were marked with PIT tags) were not significantly different from those of the first 18 years of the study (when the majority were marked with leg-bands)(Table 3). Table 4 presents combined recapture data for both leg-banded and PIT tagged animals for the whole study period. Juvenile Juvenile Adult Adult oe Female Male Female . Male Leg-banded | Total recaptures 33 9 105 ? 53 & 1 recapture) Total captures 97 69 214 134 % recapture 34.0% 13.0% 49.1% 39.5% PIT tagged Total recaptures 24 © 2 47 iS (= 1 recapture) Total captures 54 32 86 41 % recapture 42.5% 6.3% 54.7% 36.6 Chie: 1.60 — 1.04 0.77 0.12 pra NS <0.20 NS <0.31 NS <0.38 NS <0.73 Table 3. Comparison between total number of recaptures (>/= 1) for leg-banded and PIT tagged platypuses in the upper Shoalhaven River study area. Animals marked with both bands and PIT tags between 1987 and 1991 are included in both sets of data. NS = not significant. Proc. Linn. Soc. N.S.W., 125, 2004 219 CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN RIVER Juvenile Females 200 180 160 140 120 100 Frequency 0 — she Dk 34 S62 7 Bk 9. 10" 24 12-1314 15) 16 17-18 19 20M Age (years) Adult Females Frequency Li Dicer AeeD) weOb dg: Bo Oe Od 2 Ss a ea Age (years) Figure 1. Actual and minimum age frequencies of platypuses in the study. Actual ages are shown for those animals initially caught as juveniles (Juvenile Females and Juvenile Males). Estimated minimum ages shown are for animals caught first as adults (Adult Females and Adult Males). CONTINUED ON FACING PAGE. Considerable numbers of both male and female adults and juveniles were captured only once. Table 4 shows that total recaptures (>/= 1 times) and recaptures in the categories of 1, 2-5 and >5 times were lower for adult males than for females and that total recaptures of juvenile males was less 220 than half (14%) that of juvenile females (32%). The majority of recaptured males were caught within the first months after initially emerging from the nesting burrows (0 years of age), while recaptures of juvenile females were spread across 0-21 years after emergence (Figure 1 and Table 4). Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT Juvenile males Frequency ee 3045 Sy Oo on. oO 10 1 2.13" 14 a5 16 Age (years) Adult Males Frequency a, Capture mortality Of the 700 platypuses captured during the various research projects six died as a result of capture and handling (0.86%). Two drowned as a result of netting, two died suddenly within a few hours of capture (sudden death; animals appeared healthy and no obvious cause of death was identified from post- mortem examination by veterinarians), one succumbed to anaesthesia and one became caught in submerged vegetation by its transmitter attachment and drowned during telemetry work. Details are in Table 5. Proc. Linn. Soc. N.S.W., 125, 2004 BAe 6 3 8 9 O02 t 12713 14. 15-16 Age (years) DISCUSSION Sex Ratios As reported by Grant and Griffiths (1992) for the first 18 years of the study, the sex ratios for both adults and juveniles were significantly biased towards females. Table 6 compares the sex ratios for captured adult platypuses in three other areas (Grant unpublished). Although based on much smaller sample 221 CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN RIVER Juvenile Juvenile Adult Adult Female Male Female Male 1 recapture 17 10 66 34 2-5 recaptures 19 3 58 20 >5 recaptures i 0 18 6 Total recaptures 43 13 142 60 (> 1 recaptures) » % recapture 32% 14% 51% 36% Total captures 135 94 278 165 Table 4. Total recaptures of male and female juvenile and adults platypuses in the upper Shoalhaven River study area. ‘Mortalities and some animals which would have been unlikely to have been recaptured after the netting of some pools was discontinued are not included in this total captures figure. sizes, none of these were significantly different from parity. Grant and Griffiths (1992) also reported no significant difference between males and females in total numbers of platypuses (juvenile and adults not specified) captured in various rivers of New South Wales and the Australian Capital Territory (Table 6). Like the situation in the upper Shoalhaven River, during the earlier years of a study (1986-90) in the Duckmaloi River on the central tablelands of New South Wales, a bias towards females in both adult and juvenile platypuses was found. However, in the later years (1991-2000) more adult males than females were recorded (David Goldney, University of Sydney, Orange, pers. comm.). The recapture data discussed below seems to indicate that female platypuses in the upper Shoalhaven River survived for significantly longer periods than males. Over time, this longer survival of females would presumably have led to a sex ratio weighted towards females in the population. However, this explanation does not account for the disparity between numbers of male and female juveniles in this population. Most juvenile male platypuses disappeared from the upper Shoalhaven River population in their first year (86%; Table 4). However, 13% of juvenile females were recaptured in the area up to age one year, 19% were recaptured later than two years after emerging from the nesting burrows and two even remained in the area up to age 13 and 21 years respectively. Twelve juvenile females (9%) bred in the area, eight of these over a number of breeding seasons. Differential dispersal may contribute to the difference in the adult sex ratio, but again this does not explain the significant bias to females in the numbers of juveniles captured at the time they were becoming independent (late January-late March), unless most male juveniles dispersed immediately after independence, with females dispersing later. Unfortunately the data from this study do not permit this hypothesis to be rigorously tested, as most sampling only occurred early and late each year. The possibility also exists that the uneven sex ratios are determined by differential fertilisation of eggs, development of embryos or pre-emergence survival of young but no explanation arises from the data collected in this study concerning the significant Cause of Death Juvenile Juvenile Male Adult Female Adult. Total Female Male Drowned in net 1 0 1 0 2 Sudden death 0 0 0 2 2 Anaesthesia (ether) 1 0 0 0 1 Snagged transmitter 0 0 1 0 1 Total 2 0 2 2) 6 Table 5. Capture and handling mortalities in platypuses during work on various projects in the upper Shoalhaven River study area 222 Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT Location Adult Female Adult Sex Ratio Chi Male (F:M) Probability Various streams NSW/ACT" 101 117 1.15:1 1.17 ps 0.28 Various streams NSW/ACT* 47 47 1:1 i Bamard River, NSW 24 22 1.10:1 0.09 p< 0.77 Thredbo River, NSW 14 10 1.4:1 0.67 ; p< 0.41 Wingecarribee River, NSW 29 30 1:1.03 0.02 ps 0.90 Shoalhaven River, NSW 285 173. 1.85:1 27.38** p< 0.001 Table 6. Comparison of sex ratios of adult platypuses in various studies in New South Wales and the Australian Capital Territory (ACT). Collected by ‘Temple-Smith and * Griffiths (from Grant and Griffiths 1992); ** significant at < 0.01 level. bias towards females in the sex ratio of juvenile animals. In the Barnard River study referred to in Table 6, where the adult sex ratio was not different from parity, the sex ratio for juveniles was heavily male- biased (14 males; 3 females) during the single breeding season studied. A similar result was also obtained once during a single breeding season (12 males to 3 females; 1978/79) in the current study, indicating differences between individual years. However, in the 24 years during the study in which juveniles were captured, the numbers of females exceeded males in 87.5% of those years. Considerable annual differences in recorded annual sex ratios were also found in the Duckmaloi River study (David Goldney, University of Sydney, Orange pers. comm.). Age The minimum estimated age for male platypuses in this study (7 years) was considerably less than for females (up to 21 years), with nine adult females surviving for a minimum of 10 years. These data suggest that females live longer than males in the wild. However, determination of actual age, or the estimated minimum age, depended on recapture data and, as discussed, recapture of males was much lower (36% recaptured >/= once) than for females (51% recaptured >/= once). After being regularly captured previously, 13 and 21 year old females (first marked as juveniles) had not been recaptured for the last 6 and 5 years respectively of the study. Except for one adult female, which had been captured in December 2002 Proc. Linn. Soc. N.S.W., 125, 2004 and was not caught in March 2003, all the other adult females >/= 10 years of age had also not been recaptured in the latter years of the study. These data appeared to indicate a life span of 10-16 years may represent an expected upper range of longevity for female platypuses in the wild. However, the final recapture of one female at the age of 21 years showed a maximum female longevity in the wild comparable to that in captivity. While the data for males in the wild appeared to show shorter life spans (up to 7 years), this could equally represent non-recapture of older males. Reports do not suggest differing longevity between the sexes in captivity (Collins 1973; Whittington 1991). Recaptures PIT tags were initially used because of the occurrence of notched and broken male spurs as a result of bands abrading the spur base. However, it was also suspected that the lower capture rates in males, particularly juveniles may have been attributable to band losses. Bands were normally fitted more loosely to males to permit the much greater radial growth of the hind legs in this sex. No significant differences between captures for banded and PIT tagged males (Table 3) indicated that band loss could not fully explain the lack of recaptures, although four female animals, initially marked with bands and PIT tags, were found to have lost their bands during the latter part of the study, indicating some band loss. Only one PIT 223 CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN RIVER Home Maximum distance Source Location range(km) (km) : 0.2-2.0 5.6 (24 hr max.=4.0) Grant, 1983 Shoalhaven River 0.4-2.3 23 Grant, 1983, Grant et al. 1992 Thredbo River 0.3-2.3 23 Serena, 1994 Badger Creek 2.9-7.0 15.0 Gardner and Serena, 1995 Watts River and Badger Creek 0.4-2.6 2.6 Gust and Handasyde, 1995 Goulburn River Adult: 24 hr max. = 10.4 Serena et al. 1998 Yarra River, Mullum Mullum Creek, 2.9-7.3 (male) Diamond Creek Juvenile: 24 hr max. = 4.0 1.4-1.7 (female) 40 in 18 months Australian Platypus Yarra catchment (juvenile) Conservancy 1999 Andersons to Steels Ck 48 in 7 months Australian Platypus Wimmera River (young male) Conservancy 2001 Table 7. Home ranges and maximum distances moved by platypuses in various studies, including the Shoalhaven River (bold). tag failure or loss was confirmed in 220 tags applied to animals during the studies. The lack of spur damage, some evidence of band loss and no significant differences being found between recaptures of leg- banded and PIT tagged animals confirmed PIT tagging as the preferred method of marking platypuses (Grant and Whittington 1991). Large numbers of both adult male (64%) and female (49%) platypuses were not recaptured in the study area after being marked either with leg bands, PIT tags or both. This observations suggests one, or a combination of the following: Loss of marks. Double marking indicated that some band loss did occur during the study but there was little indication that PIT tags were lost or failed. Mortality. Little is known about the causes and incidence of mortality in platypuses. Foxes (or dogs) will take platypuses on land, from shallow riffle areas and by digging into burrows (Serena 1994; Grant 1993; Anon. 2002). Large eagles may also be possible natural predators of platypuses (Rakick et al. 2001). The remains of a platypus near a burrow excavated by a fox or dog, an isolated skull in a pool and part of a skull in a pile of other mammalian bones (mainly cattle and sheep) were the only observed evidence of mortality found during the studies in the upper Shoalhaven River. While 50% of 131 individuals tested positive to leptospirosis antibodies (Leptospira interogans- serovar hardjo)(Munday et al. 1998), no clinical symptoms of the disease were observed and nothing is known of any disease organism, resulting in significant mortality in this population. The Mucor fungus, which has caused mortality in Tasmanian populations, has not so far been detected in mainland populations of platypuses, including those in the upper Shoalhaven River (Whittington et al. 2002). Mobility. Diurnal and longer-term mobility over distances of up to 5.6 kilometres have been previously reported in individuals in the upper Shoalhaven River (Grant 1983, 1992, 1995; unpublished) and in other — studies. These data are summarised in Table 7. After the marking of 700 individual platypuses (including significant numbers of new juveniles) during the 30 years, it was expected that the majority of the population would eventually be marked and that unmarked dispersing juveniles or adults might still enter the area but would be in fairly small numbers. In fact, considerable numbers of new adult animals were captured throughout the study. Some individuals were captured as many as 20 times over periods of up to 21 years and yet the times between recaptures of these individuals was often quite variable. For example, despite the pools being regularly netted during the study, two adult females were only subsequently recaptured nine and 10 years respectively after their initial capture in those pools. Even for females identified as breeding in particular pools during different breeding seasons, periods of time between some recaptures of these animals ranged from | to 10 years. These latter observations suggest that a great deal of mobility probably characterises the platypus populations in the upper Shoalhaven River, although the effects of mortality and/or dispersal cannot be ruled out as reasons for the influx of new animals and the lack of recapture of a significant proportion of the platypuses in the upper Shoalhaven River study area. All of these possibilities need further study. Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT While there was capture and handling mortality during the studies in the upper Shoalhaven River population, this was quite low (< 1%) due to the utilisation of methods developed through considerable experience by the author and other researchers over the past three decades. ACKNOWLEDGMENTS The many friends and colleagues who assisted with the field work over the years are too numerous to thank individually but all are gratefully acknowledged. The last 10 years of the study would not have been possible without the help and support of Colin, Kate, Sue and Tom Heath and Paul Anink, Marie-Louise Lissone and Gina Grant. The late Athol MacDonald and the Izzard and Laurie families are acknowledged for their permission for access to the river and creek, and for their friendship and assistance in various aspects of the field work. Unfortunately Bill and Ron Izzard both died in 2003 and this paper is dedicated to their memory. Adam Battaglia and David Goldney are acknowledged for their personal communications. Richard Whittington and Joanne Connolly carried out the post-mortem examinations of the two animals which died suddenly after capture. Peter Temple-Smith, Michael Augee and an anonymous referee provided valuable comments on the manuscript. Some of the work reported was done while in receipt of funding from the Environment Australia (then Australian National Parks and Wildlife Service) and the Australian Research Council (then Australian Research Grants Committee). This work was carried out under NSW National Parks and Wildlife Service Scientific Investigations Licence A184, New South Wales Fisheries Scientific Research Permit F84/1245 and University of New South Wales Animal Care and Ethics Approvals 94/91, 97/46 and 00/45. REFERENCES Akiyama, S. (1999). Molecular ecology of the platypus (Ornithorhynchus anatinus). PhD thesis, La Trobe University, Melbourne. Anonymous. (1999). Keeping tabs on a marathon swimmer. Ripples 113, 1 Anonymous. (2001). Platypus on the move. Ripples 18, 1 Anonymous. (2002). Foxes kill four platypus. Hastings Gazette. 19 December. Burrell, H. (1927). The platypus. Sydney: Angus and Robertson. Collins, L.R. (1973). ‘Monotremes and marsupials. A reference for zoological institutions’ . Smithsonian Institution Press, Washington DC. Faragher, R.A., Grant, T.R. and Carrick, F.N. (1979). Food of the platypus (Ornithorhynchus anatinus) with notes on the food of the brown trout (Salmo trutta) in the Shoalhaven River, New South Wales. Australian Journal of Ecology 4, 171-179. Proc. Linn. Soc. N.S.W., 125, 2004 Gardner, J.L. and Serena, M. (1995). Spatial organisation and movement patterns of adult male platypus, Ornithorhynchus anatinus (Monotremata: Omnithorhynchidae). Australian Journal of Zoology 43, 91-103. Gibson, R.A., Neumann, M., Grant, T.R. and Griffiths, M. (1988). Fatty acids of the milk and food of the platypus (Ornithorhynchus anatinus). Lipids 23: 377-379. Gemmell, N.J. (1994). Population and evolutionary investigations in the platypus (Ornithorhynchus anatinus): a molecular approach. PhD thesis, La Trobe University, Melbourne. Grant, T.R. (1983). Body temperature of free-ranging platypuses, Ornithorhynchus anatinus, with observations on their use of burrows. Australian Journal of Zoology 31, 117-122. Grant, T.R. (1992). Captures, movements and dispersal of platypuses, Ornithorhynchus anatinus, in the Shoalhaven River, New South Wales, with evaluation of capture and marking techniques. In. Platypus and Echidnas. (Ed M.L. Augee). pp. 255-262. (Royal Zoological Society of NSW: Sydney). Grant, T.R. (1993). “The Bellinger River Water Supply Project Aquatic Studies - The Platypus’. Report to Mitchell McCotter on behalf of the Coffs Harbour City Council and Department of Public Works by Mount King Ecological Surveys, May, 1993. Grant, T.R. (1995). “The platypus. A unique mammal’. 2nd edition. Sydney: University of NSW Press: Sydney Grant, T.R. and Carrick, F.N. (1974). Capture and marking of the platypus, Ornithorhynchus anatinus, in the wild. Australian Zoologist 18: 133-135 Grant, T.R. and Dawson, T.J. (1978a). Temperature regulation in the platypus, Ornithorhynchus _anatinus, maintenance of body temperature in air and water. Physiological Zoology 51:1-6. Grant, T.R. and Dawson, T.J. (1978b). Temperature regulation in the platypus, Ornithorhynchus anatinus, production and loss of metabolic heat in air and water. Physiological Zoology 51: 315- 332. Grant, T.R. and Griffiths, M. (1992). Aspects of lactation and determination of sex ratios and longevity in a free-ranging population of platypuses, Ornithorhynchus anatinus, in the Shoalhaven River, New South Wales. In. Platypus and Echidnas. (Ed M.L.Augee). pp. 80-89. (Royal Zoological Society of NSW: Sydney). Grant, T.R., Griffiths, M. and Leckie, R.M.C. (1983). Aspects of lactation in the platypus, Ornithorhynchus anatinus (Monotremata), in waters of eastern New South Wales. Australian Journal of Zoology 31: 881-889. 225 CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN RIVER Grant, T.R. and Whittington, R.J. (1991). The use of freeze-branding and implanted transponder tags as a permanent marking method for platypuses, Ornithorhynchus anatinus (Monotremata: Ornithorhynchidae). Australian Mammalogy 14: 147-150. Gust, N., Handasyde, K. (1995). Seasonal variation in the ranging behaviour of the platypus (Ornithorhynchus anatinus) on the Goulburn River; Victoria. Australian Journal of Zoology 43, 193-208. Hulbert, A.J. and Grant, T.R. (1983a). A seasonal study of body condition and water turnover in a free- ranging population of platypuses, Ornithorhynchus anatinus. Australian Journal of Zoology 31: 109-116. Hulbert, A.J. and Grant, T-R. (1983b). Thyroid hormone levels in the egg-laying monotreme, the platypus, Ornithorhynchus anatinus. General and Comparative Endocrinology 51: 401-405. Munday, B.L., Whittington, R.J. and Stewart, N.J. (1998). Disease conditions and subclinical infections of the platypus (Ornithorhynchus anatinus). Philosophical Transactions of the Royal Society London. Biological Sciences 353, 1093-1099. Rakick, R., Rakick, B., Cook, L. and Munks, S. (2001). Observations of a platypus foraging in the sea and hunting by a wedge-tailed eagle. Tasmanian Naturalist 123, 3-4. Serena, M. (1994). Use of time and space by platypus (Ornithorhynchus anatinus; Monotremata) along a Victorian stream. Journal of Zoology (London) 232, 117-131. Serena, M., Thomas, J.L., Williams, G.A., Officer, R.C.E. (1998). Use of stream and river habitats by the platypus, Ornithorhynchus anatinus, in an urban fringe habitat. Australian Journal of Zoology. 46, 267-282 Temple-Smith, P.D. (1973). Seasonal breeding biology of the platypus, Ornithorhynchus anatinus Shaw 1799, with special reference to the male. PhD Thesis. Australian National University: Canberra. Whittington, R.J. (1991). the survival of platypuses in captivity. Australian Veterinary Journal 68, 32- 35. Whittington, R.J. and Grant, T.R. (1983). Haematology and blood chemistry of free-living platypuses, Ornithorhynchus anatinus (Shaw)(Monotremata: Ornithorhynchidae). Australian Journal of Zoology 31: 475-482. Whittington, R.J. and Grant, T.R. (1984). Haematology and blood chemistry of the conscious platypus, Ornithorhynchus anatinus (Shaw)(Monotremata: Ornithorhynchidae). Australian Journal of Zoology 32: 631-635. Whittington, R.J. and Grant, T.R. (1995). Haematological changes in the platypus (Ornithorhynchus anatinus) following capture. J. Wildlife Diseases 31: 386-390. Whittington, R.J., Connolly, J.H., Obendorf, D.L., Emmins, J., Grant, T.R. and Handasyde, K.A. (2002). Serological responses against the pathogenic fungus Mucor amphibiorum in populations of platypus (Ornithorhynchus anatinus) with and without ulcerative mycotic dermatitis. Veterinary Microbiology 87, 59-71. Proc. Linn. Soc. N.S.W., 125, 2004 Breeding in a Free-ranging Population of Platypuses, Ornithorhynchus anatinus, in the Upper Shoalhaven River, New South Wales - a 27 Year Study T.R. Grant!, M. GriFFitHs? AND P.D. TEmMPLE-SMITH? ' School of Biological, Earth and Environmental Sciences, University of NSW, Sydney 2052, t.grant @unsw.edu.au; *80 Dominion Circuit, Deakin, ACT, 2600 *Department of Conservation and Research, Zoological Parks Board of Victoria and The University of Melbourne Grant, T.R., Griffiths, M. and Temple-Smith, P.D. (2004). Breeding in a free-ranging population of platypuses, Ornithorhynchus anatinus, in the upper Shoalhaven River, New South Wales - a 27 year study. Proceedings of the Linnean Society of New South Wales 125, 227-234. A total of 150 captures of lactating platypuses (97 individuals) were made over a period of 27 years in the study area. The proportion of lactating females from December samples ranged from 18 to 80% (mean 43.4+17.7%; n = 21 breeding seasons). The percentage of juveniles in samples taken at the seasons when young were leaving the nesting burrows varied from 0-63% (mean 34.4+17.9%; n = 22 breeding seasons). Only 8.8% percent of captured juvenile females went on to breed in the area; one bred in its second breeding season after emergence but two others did not breed until at least their 4" breeding season. Some females bred during at least 2-3 consecutive breeding seasons but others failed to breed in consecutive years. The percentages of females lactating in the months of September to April indicated a spread in the breeding season. Lactation in the wild was apparently shorter than reported in captivity, lasting more than 3 but less than 4 months. The majority of variation in breeding activity and recruitment could not be explained in terms of drought or observed riverine and riparian changes during the study. Manuscript received 2 September 2003, accepted for publication 8 January 2004. KEYWORDS: Breeding, Drought, Lactation, New South Wales, Ornithorhynchus anatinus, Platypus, Recruitment, Sedimentation INTRODUCTION Platypuses (Ornithorhynchus anatinus) mate in late winter or early spring. Eggs are laid and the developing young are nourished on milk in the nesting burrows for several months, after which juveniles leave these burrows, become independent and most disperse from natal sites. There is a north-south cline in the timing of the breeding season, which begins earliest in north Queensland and latest in Tasmania (Temple- Smith and Grant 2001). The current study was carried out near the centre of this cline, on the southern tablelands of New South Wales in the upper Shoalhaven River. It began with the investigation of the nature of lactation and the composition of the milk of the platypus (Griffiths et al. 1973; Grant et al. 1983; Messer et al. 1983; Parodi and Griffiths 1983; Griffiths et al. 1984; Griffiths et al. 1985; Gibson et al. 1988; Teahan et al. 1991; Grant and Griffiths 1992; Joseph and Griffiths 1992). However, in the mid-1980s there was considerable change to the habitat of the platypus within the study area, with sand slugs encroaching into many of the pools and considerable bank erosion occurring as a result of poor past and present riparian and catchment land management practices. On completion of the initial studies early in the 1990s, the investigation continued by sampling in December, when females captured would be most likely to be lactating, and in February or March when juveniles had left the nesting burrows but had not yet dispersed (see also Grant, 2004 this volume). While the study has permitted general aspects of lactation to be further considered since the work of Grant and Griffiths (1992), it has also investigated the effects of stream degradation and drought on the platypus population in the upper Shoalhaven River. With regard to this latter aspect of the study, the hypothesis being tested was that successful breeding, as indicated by females breeding and young being recruited to the population each year, would be adversely affected by stream degradation and/or by droughts. BREEDING IN FREE-RANGING PLATYPUSES MATERIALS AND METHODS Study Site The study area consisted of a series of 16 pools in agricultural land, separated by riffle areas along 12.5 kilometres of the upper Shoalhaven River and 3.9 kilometres of an adjacent tributary stream, near Braidwood on the southern tablelands of New South Wales. A narrow discontinuous strip of riparian vegetation, consisting of both introduced and indigenous species of trees and shrubs, interrupted by numerous gaps, which were normally eroded as a result of access by sheep and/or cattle from the surrounding pasture land to the river. During the period of the study (late 1977 to early 2004), some of these pools suffered in-filling by sand slugs. For some pools, the effects on the habitat from sand in-filling was so severe that they were deleted from the sampling program. During the study period, three significant droughts occurred. Sampling Periods Two to four pools representative of the area (core area) were sampled during December, then again in February and/or March of 21 and 22 breeding seasons respectively over the 27 years of the study. Other pools within and outside these core area pools were sampled intermittently at various times during research in associated projects (Grant, 2004 this volume). Capture, Marking and Possessing Animals were captured using the unweighted “gill” net methods outlined in Grant and Carrick (1974). Until 1987, individuals were marked using stainless steel leg bands (Grant and Carrick 1974) but these were phased out after trials on the use of Passive Integrated Transponder (PIT) tags proved to be successful (Life Chip tags; Destron Fearing Corporation Scanner; Grant and Whittington 1991; Grant 2004; this volume). After removal from the nets, animals were weighed, measured and age and sex were determined (Temple-Smith 1973; Grant 2003 this volume). Females were injected intramuscularly with 0.1-0.2 mL of synthetic oxytocin (1-2 International Units; Syntocinon, Novartis) to induce milk “let down” (Griffiths et al. 1972, 1973, 1984; Grant and Griffiths 1992). In females that were lactating, milk could be expressed from the mammary gland, using gentle pressure along the flanks towards the areolae, 5 minutes after injection. Data Collection The percentage of lactating females captured in each December sample and the percentage of juveniles caught in relation to the total numbers of animals captured at each sampling in February and/or March were calculated. These provided indices of breeding and recruitment success for each breeding season. The timing and duration of lactation were determined from these data and from the capture and recapture of females in other pools of the study area. RESULTS Timing and duration of lactation During the 27 years of the study, captures of 150 lactating platypuses were made. A total of 97 individuals were lactating at least once during the study (Table 1). Only a single individual was found lactating in late September, with the highest proportions of lactating animals being captured in December and January. Sequential recaptures of three individuals ~ within the same breeding season showed lactation in the field lasted at least 70-98 days (2.3-3.3 months) (Table 2). Other sequential data showed that 97% (30 from 31) of females found lactating in December or January, had ceased lactation when recaptured in March. Of five individuals lactating in December, three were no longer lactating when recaptured in February (Table 2). Breeding ages of juvenile platypuses Of 137 female platypuses captured as juveniles, only 12 were later recaptured as breeding females in the study area (8.8%, Table 3). One of these individuals was lactating in its second breeding season after emergence from the nesting burrow, but three others did not breed until at least their 3“ or 4" breeding season. The individual (FJ222) which bred in the second breeding season (1983/84) failed to breed the following year (1984/85). This animal was not captured Sept. Oct. Nov. Dec. Jan Feb. Mar. __Apr Total 26 5 25 256 60 59 179 ah Lact. 1 1 7 106 24 10 1 0 Jo 3.8 20.0 28.0 41.3 40.0 17.6 0.6 0 Table 1. Numbers and percentages of individual female platypuses lactating in all samples in the upper Shoalhaven River study area 228 Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT, M. GRIFFITHS AND P.D. TEMPLE-SMITH Animal December. January FAO15 FAO19 FA046 FA126 FA133 FA158 FA158 FA161 FA161 FA185 FA185 FA185 FA209 FA212 FA214 FA276 FA335 FA368 FA368 v FA370 FA391 FA462 FA514 FA535 FA530 v FA547 FJ157 FJ222 FJ248 - FJ248 i FJ272 F272 FJ272 FJ436 FJ469 SE NENEN NE NNR NRE SG NES NR NENA o iN I SON February Ne Ne a Lactation duration March April yao - SZ, days x >98 days MMM - >70 days PA PS PS PS PS OPS PS OPM PS PM PS PS OPS PS OS OP OS! PS Pd PS OP PS OS Table 2. Recaptures of lactating platypuses within given breeding seasons V Lactating; X Not lactating; - not recaptured. in the breeding season of the next year (1985/86) but was lactating again in the subsequent breeding season. Breeding success and recruitment Twenty-eight females were captured in successive breeding seasons. While some females were captured lactating in up to three consecutive breeding seasons, many failed to breed in consecutive seasons, with 39% not lactating in a season immediately following one in which they did breed. (Table 4). The mean percentage of lactating (breeding) animals in December of 21 breeding seasons in the core section of the study area was 43.4+17.7% but the numbers and proportions fluctuated considerably between breeding seasons from 80% down to 18% of the numbers of females captured (Figure 1). Proc. Linn. Soc. N.S.W., 125, 2004 The data showed a general relationship between the numbers of juvenile platypuses captured in February and/or March in the core area of the study site and the total numbers of lactating animals captured in each of the breeding seasons sampled (Figure 1). In some years recruitment of juveniles was low after reasonable numbers of lactating females had been captured in December and in other years higher than expected recruitment levels were observed in February/ March after relatively low numbers of lactating females had been captured in the previous December. However, in general higher percentages of juveniles were captured at the end of breeding seasons when the percentage of lactating females sampled was also high (Figure 1). 229 BREEDING IN FREE-RANGING PLATYPUSES Animal 79/ 80/ 81/ 82/ 83/ 84/ 85/ 86/ 87/ 88/ 89/ 90/ 91/ 92/ 93/ 94/ 95/ 96/ 97/ 98/ 99/ O0/ O1/ 02/ 03/ 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 00 01 02 03 04 FJ157 LEV FOX PIA SP ek AX SS 2 AAA OS Se A eee Fo SS 2 ee PS Sn SS xt FJ217 ee er ae ik, eS ee Veena Ree ie as, Ne Ba de a ee te eee oe ge, Pe FJ222+ BE YG Ve KK Vor ee eee, a rs a oe ee ee ee x™ FJ230 sed Ay ME on So) ae. Sean oe ee Re a OR eS oe ee ee ee ee ee i eS FJ235# EI GR oo ee eve ea ee we we ge ee fe a ee ee ee FJ248 ABS ce XM ee” Boe eK Xe XS OX ETS OR Oo eee Ee KO eS eee ee ee yee ye FJ272# A KR BX ea ee VY Ve AK ee ee SS Se ee ee = x™ 2 x™ vi FJ273 BOR ee NS SNR Aas SNe Ae cee VAP Lemme Ca MR ego ae elo a aes kc! ee eee FJ409 i I oS ae i ae ee Lae RR Be ee Ba ee FJ436 . Be Te A Ne EN Se ee ee me Se Bd x” ; FJ469# RR NEY ee Se PS yt eo Sea ss 2 xe FJ496 a 2 Xoo Eee a ef ee eS ye Table 3. Juvenile platypuses originally captured in the study which later bred within the study area. ** emerged; X not lactating; V lactating ‘ month of capture (eg November); - not captured/sampled; + First breeding in second breeding season after emergence from nesting burrow; # First breeding at 3-4 breeding seasons after emergence. Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT, M. GRIFFITHS AND P.D. TEMPLE-SMITH Animal 77/ 78/ 79/ 80/ 81/ 82/ 83/ 84/ 85/ 86/ 87/ 88/ 89/ 79 80 81 FA019 FA124 FA126 FA133 FA147 FA158 FA161 78 a v 29-90 0 pe pd ARS ww NK! 82 Ss SEAN 83 v \ NK MK gM: 84 | Bie Sdn 85 v 86 MK 87 838 v wo NSS: $9 90 KS SN Wye 90/ 91/ 92/ 93/ 94/ 95/ 96/ 97/ 98/ 99/ 96 «97 91 \ SNK! NK KN SKE 92 93 94 95 wr! 98 99 Table 4. Female platypuses captured in November to January in consecutive breeding seasons in the whole study area during the study. ** newly emerged juveniles; V lactating; X not lactating; ? caught during February, March or April (could have bred but have finished lactating; see Table 2). 00/ O1/ 02/ 03/ 00. O01 02 03 04 231 Proc. Linn. Soc. N.S.W., 125, 2004 BREEDING IN FREE-RANGING PLATYPUSES 100 7 % Lactating Females or Juveniles in Samples iS) uo _~ Nn oO ~~ oo oO oO Sy oS) So j=) oO oS) 1 i tt 1 i 1 1977/78 SS a ES ST ee 0 ————————SS ICQ? =e ee 1983/84 aS See+Do a EC aS] 1984/85 Ee i SP 1986/87 ays ea ae 1987/88 RS ee eee Se 1988/89 Bi eo ee 1982/83 has See 1985/86 (epee —_—$__— 1978/79 1980/81 | 1991/92 a ee ee oo a a4 + ra) \o > =) is) = NX cl NN SS ONT = SN DN SS) a es et) nN oO tT al \o co f=) So = nN fen) oO’ Ov ()) fo) fo) fo)) S 4 4 — = = = — AQ os] aQ Breeding Seasons Figure 1. Percentages of lactating females captured in relation to total adult females caught in each December sample (n = 21) and the percentages of juveniles in each February and/or March sample (n = 22) in the core study area. Open bars = lactating females (5 seasons not sampled); solid bars = juveniles (4 seasons not sampled; 1988/89 and 1991/92 seasons no juveniles were caught). DISCUSSION The low overall percentages of lactating animals in samples caught in September (3.8%), October (20%), November (28%) and in February (17.6%), March (0.6%), April (0%), contrasted with higher percentages in December (41.3%) and January (40%). This indicated a spread in the breeding season. However, it appears that the majority of animals were breeding around the same time, with a few individuals breeding earlier (eg. one already lactating by the end of September) and a few later (eg. one still lactating in March; Table 1). The sequential recaptures of lactating females within the same breeding seasons provided evidence that lactation in the wild can last at least 98 days (3.3 months) but is unlikely to exceed four months. This suggestion is supported by the distribution of lactating _ females in the various months (Table 1), combined with the observation that all but one of 31 females lactating when captured in December or January (97%) had ceased lactation on their subsequent recapture in March (Table 2). Nestlings in the wild may be weaned more rapidly than those bred in captivity. Lactation in captive animals has been reported to continue as long as 145 days (4.8 months; Holland and Jackson 2002; Healesville Sanctuary and Taronga Zoo, unpublished) and requires lactating females to consume up to 100% of their body weight in food during peak lactation (Holland and Jackson 2002). It may be that the young are weaned more quickly in the wild depending on the local availability of macroinvertebrate food items (Faragher et al. 1979) for the breeding females. Certainly in some years of this study, lactating females were in poorer body condition, based on observations of tail fat reserves (Temple-Smith 1973; Grant and Carrick 1978) and general body condition. Interestingly, both the lactating females and the captured juvenile animals in the 2002/03 breeding season, at the end of a very severe drought, were judged to still be in good body condition. However, four lactating females captured at the beginning of January 2004 appeared to be in poorer condition, despite the river flows being an improvement on those of the previous breeding season, when surface flows in the Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT, M. GRIFFITHS AND P.D. TEMPLE-SMITH study area stopped for several weeks (pers. comm. from local residents). Recapture rates of juvenile platypuses during the various studies in the area was quite low. Thirty- two percent of female platypuses first captured as juveniles were recaptured compared to only 14% for males (Grant 2004; this volume). In spite of this, only 12 female juveniles were recruited into the breeding population. Although the breeding season was predictable, the breeding of individuals in any one season was much less predictable, with varying numbers of non-breeding females in any sample, individuals not breeding until later in life and breeding animals failing to breed in consecutive seasons (Table 4). Similar observations have been made with platypuses in captive conditions, where no individuals have so far bred in successive breeding seasons (Holland and Jackson 2002; Healesville Sanctuary and Taronga Zoo, unpublished). Temple-Smith and Grant (2001) have speculated whether resource availability, social organisation or genetic factors are involved in this uncertain breeding in the species but little is known of any of these aspects of platypus biology. A decline was expected in the number of platypuses breeding and/or the number of juveniles recruited to the population after the mid 1980s, when sand slugs began to reduce the pools available for foraging and the provision of refuge areas during drought. Surprisingly, no such overall trend occurred in either numbers breeding or in recruitment data and there is no ready explanation for the considerable variation in the numbers breeding between the seasons covered by the study. Such variations must be attributable to more subtle changes: occurring in the environment and/or to unexplained sampling effects. Both breeding success and recruitment fell sharply in the 1982/83 breeding season, corresponding to the end of a long and severe drought, which lasted from October 1978-February 1983. The effects of the drought provided an explanation for the observation that three females lactating in the 1981/82 season did not breed in the 1982/83 season. However, there were no similar trends recorded in the 1993-95 or 2001-03 droughts, although during the latter, lactation and recruitment percentages were slightly below the mean values for each (Fig. 1). As discussed above, all lactating females and juveniles captured in the 2002/ 03 sampling were considered to be in good body condition in spite of the severe drought conditions which existed at the time. During the 1988/89 and 1991/92 breeding seasons no juvenile platypuses were captured. There is no obvious reason for the observed lack of recruitment in 1988/89, but two local over-bank flood Proc. Linn. Soc. N.S.W., 125, 2004 events in late December/early January (pers. comm. from local residents) of 1991/92 may have drowned many nestlings confined to burrows during that season. This would explain the failure to capture juveniles in a year when the percentage of lactating females in the previous December had been slightly higher the mean value (Fig. 1). Prior to enactment of legislation protecting platypuses in all states of Australia (1892 in Victoria to 1912 in South Australia; Grant and Denny 1991), thousands were hunted for their fur. Their numbers are reputed to have declined dramatically, although rebounding since protection has been enforced (Grant and Denny 1991; Grant and Temple-Smith 1978). The species is currently listed as protected, but is either regarded as ‘common’ or not threatened, in all states (except for South Australia, where it is probably now extinct, except for an introduced population on Kangaroo Island). In spite of this, there is concern at the fragmentation of populations in some river systems and in small local populations as a result of habitat degradation, illegal and recreational fishing and encroaching effects of urban and regional development (Grant and Temple-Smith 2003). While this study demonstrates that the species has continued to survive and reproduce in the upper Shoalhaven River in spite of considerable riparian and riverine degradation, the effects of drought and the combination of both of these perturbations, further investigation leading to a complete understanding of the factors determining the uncertain breeding in the species is critical to its conservation. Many questions regarding the population biology and reproduction of Ornithorhynchus anatinus still remain unanswered but the long-term studies reported here and in Grant (2004, this volume) have gone some way to providing a greater understanding of some aspects of the species’ field biology, which could not have been achieved by a study of shorter duration. ACKNOWLEDGMENTS Merv Griffiths, a friend, colleague and our co- author died on 06 May 2003. The many other friends and colleagues who were instrumental in the success of field work, in often severely inclement conditions, over the years are too numerous to name individually but the Heath family, Paul Anink, Marie-Loiuse Lissone, David Read and Gina Grant deserve special mention. All are gratefully acknowledged. Some of the work reported was done while in receipt of funding from the Environment Australia (then Australian National Parks and Wildlife Service) and the Australian Research Council (then Australian Research Grants Committee). The late Athol MacDonald and the Izzard and Laurie families are acknowledged for their permission to access the river and creek on the properties managed or belonging to them, and for their friendship and i) Oo oS) BREEDING IN FREE-RANGING PLATYPUSES assistance in various aspects of the field work. The work was carried out under NSW National Parks and Wildlife Service Scientific Investigations Licence A184, New South Wales Fisheries Scientific Research Permit F84/1245 and University of New South Wales Animal Care and Ethics Approvals 94/91, 97/46 and 00/45. REFERENCES Faragher, R.A., Grant, T.R. and Carrick, F.N. (1979). Food of the platypus, Ornithorhynchus . anatinus, with notes on the food of the brown trout, Salmo trutta, in the Shoalhaven River, New South Wales. Australian Journal of Ecology 4: 171-179. Gibson, R.A., Neumann, M., Grant, T.R. and Griffiths, M. (1988). Fatty acids of the milk and food of the platypus (Ornithorhynchus anatinus). Lipids 23, 377-379. Grant, T.R. (2004). Captures, capture mortality, age and sex ratios of platypuses, Ornithorhynchus anatinus, during studies over 30 years in the upper Shoalhaven River in New South Wales. Proceedings of the Linnean Society of New South Wales 125, 217-226. Grant, T.R. and Carrick, F.N. (1974). Capture and marking of the platypus, Ornithorhynchus anatinus, in the wild. Australian Zoologist 18: 133-135. Grant, T.R. and Carrick, F.N. (1978). Some aspects of the ecology of the platypus, Ornithorhynchus anatinus in the upper Shoalhaven River, New South Wales. Australian Zoologist 20: 181-199. Grant, T.R. and Denny, M.J.S. (1991). Historical and Current Distribution of the Platypus in Australia, with Guidelines for the Management and Conservation of the Species. Unpublished Report to Australian National Parks and Wildlife Service by Mt. King Ecological Surveys. : Grant, T.R., Griffiths, M. and Leckie, R.M.C. 1983. Aspects of lactation in the platypus, Ornithorhynchus anatinus (Monotremata), in waters of eastern New South Wales. Australian Journal of Zoology 31, 881-889. Grant, T.R. and Griffiths, M. (1992). Aspects of lactation and determination of sex ratios and longevity in a free-ranging population of platypuses, Ornithorhynchus anatinus, in the Shoalhaven River, New South Wales. In ‘Platypus and Echidnas’. (Ed M.L.Augee). pp. 80-89. (Royal Zoological Society of NSW: Sydney). Grant, T.R. and Temple-Smith, P.D. (1998). Field biology of the platypus (Ornithorhynchus anatinus) - historical and current perspectives. Transactions Royal Society London Series B 353, 1081-1091. Grant, T.R. and Temple-Smith, P.D. (2003). Conservation of the platypus, Ornithorhynchus anatinus: Threats and challenges. Aquatic Health and Management 6, 1-15. 234 Grant, T.R. and Whittington, R.J. (1991). The use of freeze-branding and implanted transponder tags as a permanent marking method for platypuses, Ornithorhynchus anatinus (Monotremata: Omithorhynchidae). Australian Mammalogy 14: 147-150. Griffiths, M., Elliott, M.A., Leckie, R.M.C. and Schoefl, G.I. (1973). Observations on the comparative anatomy and ultrastructure of mammary glands and on the fatty acids of the triglycerides in platypus and echidna milk fats. Journal of Zoology [London] 169, 255-275. Griffiths, M., Green, B., Leckie, R.M.C., Messer, M. and Newgrain, K.W. (1984). Constituents of platypus and echidna milk with particular reference to the fatty acid complement of the triglycerides. Australian Journal of Biological Science 37, 323-329. Gniffiths, M., McIntosh, D.L. and Leckie, R.M.C. (1972). The mammary glands of the red kangaroo with observations on the fatty acid components of the milk triglycerides. Journal of Zoology [London] 166, 265-275. Griffiths, M., McKenzie, H.A., Shaw, D.C. and Teahan, C.G. (1985). Monotreme milk proteins: echidna and platypus “whey” proteins. Proceedings of the Australian Biochemical Society. 17, 25. Holland, N. and Jackson, S.M. (2002). Reproductive behaviour and food consumption associated with captive breeding of platypus (Ornithorhynchus anatinus). Journal of Zoology [London] 256, 279-288. Joseph, R and Griffiths, M. (1992). Whey proteins in early and late milks of monotremes (Monotremata; Tachyglossidae, Ornithorhynchidae) and of the tammar wallaby (Macropus eugenit)(Marsupialia; Macropodidae). Australian Mammalogy 15, 125-127. Messer, M., Gadiel, P.A., Ralston, G.B. and Griffiths, M. (1983). Carbohydrates of the milk of the platypus (Ornithorhynchus anatinus). Australian Journal of Biological Science. 36, 129-138. Parodi, P.W. and Griffiths, M. (1983). A comparison of the positional distribution of fatty acids in milk triglycerides of the extant monotremes platypus (Ornithorhynchus anatinus) and echidna (Tachyglossus aculeatus). Lipids 18, 845-847. Teahan, C.G., McKenzie, H.A. and Griffiths, M. (1991). Some monotreme milk “whey” and blood proteins. Comparative Biochemistry and Physiology. 91B, 99-118. Temple-Smith, P.D. (1973). Seasonal breeding biology of the platypus, Ornithorhynchus anatinus (Shaw 1799) with special reference to the male. PhD thesis, Australian National University, Canberra. Temple-Smith, P.D. and Grant, T.R. (2001). Uncertain breeding: A short history of reproduction in monotremes. Reproduction, Fertility and Development 13, 487-497. Proc. Linn. Soc. N.S.W., 125, 2004 Depth and Substrate Selection by Platypuses, Ornithorhynchus anatinus, in the Lower Hastings River, New South Wales Tom GRANT School of Biological, Earth and Environmental Sciences, University of New South Wales, NSW 2052 t.grant@unsw.edu.au Grant, T. (2004). Depth and substrate selection by platypuses, Ornithorhynchus anatinus, in the lower Hastings River, New South Wales. Proceedings of the Linnean Society of New South Wales 125, 235- 241. Platypuses were observed foraging most frequently in water >1 metre in depth during normal (91.3%) and drought (82.1%) conditions. Mean water depth in the study pools was 1.08+0.66 and 0.86+0.61 metres during normal and drought conditions respectively. The distribution of depths in the study area was significantly different from the distribution of depths where platypuses were observed during normal (Chi? = 90.2; p < 0.01) and drought conditions (Chi? = 37.35; p < 0.01). Platypuses were apparently not simply utilising depths in relation to their occurrence but preferring to forage in water deeper than 1.5 metres and avoided depths < 1 metre. Overall distribution in numbers of platypuses observed foraging over different benthic substrate types was not significantly different (Chi? = 12.9; p > 0.05) from the distribution of these substrate categories in the study area. However, when the substrates were considered separately, significant preference was shown for cobbled substrate (Chi? = 18.4; p < 0.01) and avoidance of gravel (Chi? = 9.7; p <0.01). These observations have implications for catchment, stream and riparian management, where activities leading to sedimentation and reduced flushing flows may reduce depths and/or alter the distribution of preferred foraging substrates. Manuscript received 2 September 2003, accepted for publication 7 January 2004. KEYWORDS: depth, foraging, Hastings River, Ornithorynchus anatinus, platypus, substrate. INTRODUCTION During foraging in the wild, platypuses dive to feed almost exclusively on small benthic invertebrate animals (Faragher et al. 1979; Grant 1982), which are normally unevenly and often sparsely distributed in a variety of substrates and depth zones (Boulton and Brock 1999; Elliott 1977; Young 2001). The platypus is small, has a high metabolic demand to regulate its body temperature in water and has an estimated maximum aerobic capacity for diving of only 40-60 seconds (Bethge 2002; Bethge et al. 2001; Evans et al. 1994; Grant and Dawson, 1978). Consequently its foraging is restricted to relatively shallow depths and the species is seldom reported occurring in deep lakes or impoundments (Bryant 1993; Grant 1991; McLeod 1993: Ellem et al. 1998; Ellem and McLeod 1998). The current study reports on observations of depths of diving and foraging over different substrates by platypuses in a coastal river in New South Wales during drought and normal flow conditions. Grant and Bishop (1998) discussed the importance of the measurement of physical habitat variables associated with platypus occurrence as a means of assessing possible impacts of stream use and management activities. As part of the monitoring and detection of possible environmental effects of the Hastings District Water Supply Augmentation Scheme, the utilisation of depth and substrate categories by foraging platypuses was investigated. METHODS Study area The study was undertaken in two separate 1.5 kilometre sections of the lower Hastings River near Wauchope in New South Wales. Immediately above a large riffle separating the riverine section from the upper estuary tidal influence, the study area consisted of a series of pools, riffles and runs, with the banks predominantly consisting of earth consolidated by the roots of riparian vegetation, but with a number of gravel/cobble bars and sections of bedrock also present . Predominantly surrounded by agricultural land, especially pastures for dairy cattle, much of the stream bank supported a narrow strip of vegetation consisting of river oaks (Casuarina sp), rainforest species (e.g. Waterhousea floribunda, Ficus coronata) and DEPTH AND SUBSTRATE SELECTION BY PLATYPUSES introduced weed species (e.g. willows, Salix sp; Lantana camara; privet, Ligustrum spp, wild tobacco, Solanum mauritianum). A range of macrophyte species also occurred in the stream (especially Myriophyllum verrucosum), although these were reduced to low incidences after several flood events. The aquatic grass, Potamophila parviflora, was also common along several sections of bank and occurred in island clumps within several sections of the stream. Sampling _ The 3 kilometres of river surveyed consisted of four riffle areas and five pools. Each section was surveyed in both directions during the two hours prior to darkness and immediately after first light in winter (late May to early July) and spring (September to October) over six years from 1998-2003 (88 longitudinal transects x 2 river sections = 176 longitudinal transects; i.e. the whole 3 kilometres was surveyed 88 times). During 1998-2000 the same number of longitudinal transects (16) was surveyed in both winter and spring but from 2001 to July 2003 fewer were surveyed in winter (8) and more in spring (24), as lower numbers of platypuses were normally observed during the winter period. Depth and the predominant substrate type were recorded at the point where each platypus was first seen foraging. It should be noted that visibility, due to turbidity and/or poor light conditions, often meant that a determination of substrate could not be made at all of these. points. During the 2001 and 2002 sampling periods, visibility was so low (probably due to the high abundance of phytoplankton) that the substrate could be observed only in few instances where platypuses were foraging. Physical habitat analysis The stream was paced out into 60 x 50 metre sections (3 km) and marked at each point with brightly- coloured flagging tape. At each of these points depth measurements were made at both edges (approximately 2 metres from bank) and in the middle of the river (using a weighted line or the kayak paddle graduated in 25 cm units). These depth measurements (n = 174) provided a measure of the distribution of depth categories in the study area (Figs 1 and 2). The occurrence of benthic substrates (mud, sand, gravel, cobble and bedrock) was scored on a scale of 1-5 (using the following estimated percentage cover of each substrate type; 0 = 0%; 1 = 0-5%; 2 = 5-25%; 3 = 25- 50%; 4 = 50-75%; 5 = >75%) along three transects . parallel to the stream bank between corresponding depth measurement points at the beginning and end of each 50 metre section. Substrates along these transects were not homogeneous, often with some of each type within a single transect. However, the predominant 236 substrate types (score of 4 or 5; i.e. >50% estimated coverage) for each transect (n = 174) were used as a measure of the distribution of the occurrence of substrates (Fig. 3). Data on depths and substrate distribution were collected once during July 2000 but depth measurements were repeated in October 2002 when the river was under severe drought conditions and was barely flowing. Data analysis The null hypothesis being tested was that the occurrence of platypuses across depth and substrate categories was the same as the occurrence of these categories in the study area. The overall distribution of numbers of platypuses observed foraging within the various depth and substrate categories and the recorded numbers of occurrence of these physical attributes in the stream (n = 174 samples), were compared using Chi? analysis (Statistica, StatSoft Inc.) with expected values calculated using 2 x 5 contingency tables (Bailey 1969). Comparisons between numbers of platypuses observed foraging at specific depths or on specific substrates compared to those not foraging at these specific depths or substrates (i.e. all other depths or substrate categories) were made using Chi? for 2 x 2 contingency tables. Comparisons between drought and non-drought measurements of depth were made using Student’s t-tests for unpaired samples (Bailey 1969; Statistica; StatSoft Inc). Indices of selection/avoidance (Response Index) of depth or substrate categories by platypuses were calculated as: Response Index = % Occurrence of platypuses in a depth or substrate category % Occurrence of that depth or substrate category in the stream An index of greater than unity suggested a selection response and less than one an avoidance response to a depth or substrate category. All means given are + Standard Deviation. RESULTS Depth selection The mean depths of the stream during non- drought and drought were significantly different, being 1.08+0.66 and 0.86+0.61 metres respectively (t = 3.48; p < 0.001). The maximum water depth recorded by these transect-based measurements was 2.75 metres but platypuses were recorded foraging at depths of up to 3.2 metres and the maximum depth recorded opportunistically (not along transects or at foraging sites) was just under 4 metres. Figure | shows the numbers of observations Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT oy ( fo) oO recorded/Depth category N WwW oO oO % Occurrence of platypuses or Depths 0.0-0.5 0.6-1.0 1.1-1.5 1.5-2.0 >2.0 Depth Category (metres) Figure 1. Percent occurrence of platypuses observed foraging in various depth categories (n = 127 observations; white bars) and the percentage of occurrence of these depth categories (n = 174 observations; black bars) in the Hastings River study area during non-drought conditions. Hah 0.0-0.5 0.6-1.0 1.1-1.5 1.5-2.0 >2.0 Depth Category (metres) s (a (oy) o Oo oO N oO % Occurrence of platypuses or Depth: recorded/Depth category Ww oS Figure 2. Percent occurrence of platypuses observed foraging in various depth categories (n = 28 observations; white bars) and the percentage of occurrence of these depth categories (n = 174 observations; black bars) in the Hastings River study area during drought conditions. of platypuses foraging in the various depth categories during a range of non-drought conditions. These data show that 91.3% of the platypuses (total n = 127) were observed foraging in depths greater than 1 metre, despite this depth category occurring in only 39.1% of the study area. The distribution of numbers of platypuses foraging within the depth categories and the recorded numbers of occurrence of these categories were significantly different (Chi? = 90.2; p < 0.01). Platypuses showed significant preferences for - water deeper than 1.5 metre (Response Indices 3.1- Proc. Linn. Soc. N.S.W., 125, 2004 60 50 40 30 20 10 % Occurrence of platypuses or substrates/categoi Mud Sand Gravel Cobbles Bedrock Substrate Category Figure 3. Percent occurrence of platypuses observed foraging in various substrate categories (n = 56 observations; white bars) and the percentage of occurrence of these substrate categories (n = 174 observations; black bars) in the Hastings River study area during non-drought and drought conditions. 3.2) and avoidance of depths of less than 1 metre (Response Indices 0.2-0.7). Foraging within the 1.1- 1.5 metre category showed no significant preference or avoidance by platypuses (Table 1a). During severe drought conditions (July and October 2002) there was a significant difference between the distribution of foraging platypuses (n = 28) and the distribution of recorded depth categories (Chi? = 37.35; p = < 0.01; Fig. 2). Response Indices showed a similar, but apparently more marked pattern towards preference for depths > 1 metre (Response Indices 2.6-6.2) and avoidance of shallower depths (Response Indices from 0-0.7). Considering the small sample sizes of platypuses foraging in specific depth categories (n = 0-10) no Chi? analyses were attempted on these data collected during the drought. Substrate selection The numbers of platypuses observed foraging on particular benthic substrate types (n = 56) are shown in Figure 3. The overall distribution of numbers of platypuses foraging over the various substrates was not significantly different from the distribution of these substrates (Chi? = 12.9; p > 0.05). However, only 26.8% of platypuses were found foraging over gravel substrates, while this substrate type was the most abundant in the study area (50.9%). While cobbles made up only 12.7% of the available substrate, 28.6% of the platypuses observed were foraging over this substrate. Response Indices of 0.5 and 2.3 respectively 2357 DEPTH AND SUBSTRATE SELECTION BY PLATYPUSES Table 1. Chi? and probability values (2x2 contingency tables; Statistica; StatSoft Inc.) for comparisons of: a. platypuses foraging at specific depths and those not foraging at each of these depths Depth Category 0.0-0.5 m 0.6-1.0 m Chi * De 38.8 <0) (0 1* <0 1 1.1-1.5 m 1.6-2.0 m >2m 43 18.8 43 > 0.05 < 0.01* < 0.01* b. platypuses foraging on specific substrates and those not foraging on each of these substrates Substrate Category Mud Sand Gravel Cobbles Rock Chi ” 0.01 0.96 9.70 18.70 0.26 > 0.05 > 0.05 < 0.01* < 0.01* > 0.05 * indicates statistical significance. for gravel and cobbles, suggested avoidance of the former and preference for the latter. Individual comparisons between platypuses foraging over specific substrates compared to all other substrates showed significant differences from expected for gravel and cobbles but not for the other substrate types (Table 1b). DISCUSSION The study suggested that platypuses observed in the early morning and late afternoon/evening were selecting the deeper sections of their habitat, with 91.3% of the observed individuals foraging in water > 1 metre in depth and 33.1% foraging in water of greater than 2 metres, which constituted only 39.1% and 10.3% of the area respectively during normal flow conditions. Even during the severe drought conditions, 82.1% of platypuses were still observed foraging in water deeper than 1 metre, despite the fact that the proportion of recorded depths > 1 metre had decreased by 11.5%. During the drought observations, 48.3% of the area had a depth of 0.0-0.5 metres but no platypuses were observed forging in this depth category. Thus, platypuses appeared to show preference for foraging in deeper areas and an avoidance of shallower depths within the area during the study in both drought and non-drought conditions. These observations were similar to those reported for a study in a small alpine lake in Tasmania (Lake Lea). While reporting a maximum dive depth of 8.77 metres, Bethge (2002) and Bethge et al. (2003) found that the majority of dives recorded for platypuses fitted with data loggers were to depths of less than three metres (98% of dives), with a mean diving depth of 1.28 metres. These workers also found a large 238 proportion of the foraging dives were to depths of less than one metre (48%), although, during winter (when the lake level was higher than in summer), most dives were to depths greater than | metre. Platypuses were monitored foraging in the same area, rather than moving to the shallower parts of the lake during the winter, suggesting that foraging was determined by factors other than depth preference, possibly including substrate type and/or availability of benthic food species. However, the workers in this study did not report on these possibilities. Rohweder (1992), Bryant (1993) and McLeod (1993) also reported platypuses foraging in water less than 5 metres in depth. While Ellem et al. (1998) found increasing depth of pools (up to 2 metres) to be positively related to the observed presence of platypuses in 36 pools on the Macquarie River system in the Bathurst area of the central tablelands of New South Wales, Ellem and McLeod (1998) found radio- tacked platypuses using shallower parts of some sections of a weir pool in the Duckmaloi River near Oberon in New South Wales. Bethge (2002) also reported platypuses foraging in deeper areas and spending less time on the surface of the water in Lake Lea during daylight hours than at night. He speculated that these behavioural changes may have been related to avoidance of predators. Little is known regarding predation by indigenous predators which could take foraging platypuses from the water. Predation by the introduced red fox (Vulpes vulpes) on platypuses moving through or foraging in shallow water, such as riffle areas, has been reported (Serena 1994; Grant 1993; Anon. 2002) and the species has been included as a possible food item of wedge-tailed eagles (Aquila audax)( Rakick et al. 2001; Marchant and Higgins 1993). Several white-breasted sea eagles (Haliaeetus Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT leucogaster) and several ospreys (Pandion haliaetus) were observed at the Hastings River study site. Both of these species were seen taking fish from the surface of the water. Although a grey goshawk (Accipiter novaehollandiae) has been reported attacking a juvenile platypus on land (Richards 1986), it seems unlikely that either this species or the osprey would be large enough to take even a juvenile platypus (which are around 65-70% adult weight when they first leave the nesting burrows; Grant and Temple-Smith 1998) from the water. It is possible however, that the sea eagle may represent a potential predator of the platypus. During the study a sea eagle was observed retrieving a large dead Australian bass (Macquaria novemaculeata) of 485 mm in length, and estimated to weigh 2.5 kg (Harris 1987), from the bottom of a pool. The eagle was unable to fly with the fish and dragged it to a nearby gravel bar, where part of the flesh was eaten before darkness fell. Soon after first light the following morning the eagle was seen carrying off the remaining carcass of the bass. Interestingly, the platypus does not seem to have been recorded as a food item of this species of large eagle (Marchant and Higgins 1993; Olsen 1999). Substrate selection Higher invertebrate productivity is often associated with areas where logs, roots and vegetation provide a range of habitats for an array of types of benthic invertebrate species and coarse substrates (gravel, cobbles, rocks) provide fixed habitat, rather than a shifting substrate, such as sand and fine sediment (Young 2001; Boulton and Brock 1999; Smith and Pollard 1998). Data in the study were restricted by a small sample size due to the inability to observe the type of substrate over which platypuses were seen foraging in times of high turbidity and/or poor light conditions. However, there was some indication that platypuses were avoiding sections of the study area which consisted mainly of mud or sand (Fig 3) but this was not statistically significant (Table 1b). There appeared to be marked avoidance of gravel (the most abundant substrate; 50.9% of the area) and preference shown for areas where cobbles were the predominant substrate (12.7% of the area). Both of these trends were Statistically significant (Table 1b). The complexity of benthic habitat has been previously identified as being positively related to the occurrence of platypuses (Rohweder 1992) and Serena et al. (2001) found a positive relationship between numbers of radio-tracked platypuses and the occurrence of coarser substrates, including gravel, pebbles, cobbles, large rocks and coarse particulate organic matter. These observations may be related to the distribution of benthic food organisms but this was Proc. Linn. Soc. N.S.W., 125, 2004 not investigated in the present study. No explanation of the apparent avoidance of gravel substrate in the present study is suggested as the species has been observed by the author foraging on gravel substrates in other areas. Implications for stream management The development of adaptive management strategies for streams, particularly with regard to water extraction and the operation of impoundments, should consider flows which maintain pool depth and benthic habitat diversity by preventing the accumulation of sand and fine sediments. The removal of riparian vegetation, erosion as a result of unrestricted stock access to stream banks and poor catchment management practices have also resulted in the infilling of pools by sand ‘slugs’ in many streams in eastern Australia (Brooks and Brierley 1996; Boulton and Brock 1999; Brierley et al. 1999; Grant et al. 2003). Grant and Bishop (1998) encouraged the use of physical habitat analysis, considering broad habitat variables normally associated with platypus occurrence, in any attempts to monitor and/or predict effects of human activities impinging on streams and their catchments. More recently a habitat simulation model was used by Davies and Cook (2001) to generate weighted useable habitat area estimates for the platypus at various proposed discharge regimes in a regulated river in Tasmania. This model used more specific habitat requirements of the species in terms of depth, velocity and substrate, calculating habitat preference curves based on available information from the literature and from experts in the field. These authors observed that: “Platypus[es] are known to feed in very shallow water and up to ca 1-3 m” and “foraging is optimal at depths of < 2 m” and “platypus[es] actively feed in silt, sands, finer gravel substrates, and are known to forage on coarse gravel to smaller cobble substrate. Feeding activity is not deemed to be efficient or to frequently occur on coarse cobble, boulder or bedrock substrates” While the important modelling work of Davies and Cook (2001) drew upon the information available to the authors at the time, the data from the current study and that from Serena et al. (2001) do not totally support the information used to generate their habitat preference curves for the species in mainland sites. It is vitally important that studies seeking to predict possible impacts of human activities on the platypus (or any other species) must consider the widest range and the most currently available information on which to base assumptions. Too often, assessment of possible environmental impact is based on ‘conventional wisdom’ which may be enshrined in publications 239 DEPTH AND SUBSTRATE SELECTION BY PLATYPUSES which are either not current or poorly researched. It is not acceptable, for example for one Environmental Impact Statement to become the main reference for statements or predictions made in another such document, without reference to the wider and most current scientific literature. The following example from the assessment of dam development on the Burnett River in Queensland sharply illustrates these concerns. Arthington (2000) suggested that “platypus[es] feed by scooping prey items and mud into cheek pouches in the mouth and grinding the mixture to a sludge before digesting it’. Based on this suggestion, the resultant Environmental Impact Statement concluded that “the deposition of sediments in the shallower areas of the dam would provide extra foraging area for Platypus[es]” as “an increase of available muddy substrate would provide more foraging area. Hard substrates offer less feeding opportunities because prey cannot be as easily scooped and ground up if they are on hard substrates or if the scooped material contains large pebble material” (Anon. 2003). Neither the literature nor the current study support the original suggestion by Arthington (2000) which consequently has led to a very equivocal prediction regarding the possible impact of dams on platypus foraging. It is crucial that such equivocal predictions do not become established in the literature consulted by those carrying out environmental impact assessment studies. ACKNOWLEDGEMENTS This work was carried out during monitoring studies associated with the Hastings District Water Supply Augmentation Scheme and was funded by Hastings Council and the New South Wales Department of Commerce Offices of Government Procurement and Government Business (formerly Department of Works and Services). Keith Bishop provided valuable advice during this study and comments on drafts of this paper. Michael Augee and another anonymous referee are thanked for their comments and recommendations on the paper. REFERENCES Anonymous. (2002). Foxes kill four platypus. Hastings Gazette, 19 December, 2002 p. 5. Anonymous. (2003) “Burnett Catchment Water Infrastructure - Burnett River Dam. Environmental Impact Assessment. 11. Aquatic Flora and Fauna’. (Burnett Water Pty. Ltd.: South Brisbane). 240 Arthington, A.H. (2000). “Burnett Basin WAMP Current Environmental Conditions and Impacts of Existing Water Resource Development’. Appendix H. “Reptiles, Frogs, Rats, Platypus and Birds’. (Queensland Department of Natural Resources: Brisbane). Bailey, N.T.J. (1969). ‘Statistical Methods in Biology’. (The English Universities Press, London). Bethge, P. (2002). Energetics and foraging behaviour of the platypus Ornithorhynchus anatinus. PhD thesis. University of Tasmania, Hobart. Bethge, P., Munks, S. and Nicol, S. (2001). Energetics and locomotion in the platypus, Ornithorhynchus anatinus. Journal of Comparative Physiology - B, Biochemical, Systematic and Environmental Physiology 171, 497-506. Bethge, P., Munks, S., Otley, H. and Nicol, S. (2003). Diving behaviour, dive cycles and aerobic dive limit in the platypus, Ornithorhynchus anatinus. Journal of Comparative Physiology A 136, 799- 809. Boulton, A.J. and Brock, M. (1999). “Australian Freshwater Ecology. Processes and Management’. (Gleneagles Publishing: Glen Osmond, South Australia). Brooks, A.P. and Brierley, G.L. (1996). Geomorphic responses of the lower Bega River to catchment disturbance 1851-1926. Geomorphology 18, 291-304. Brierley, G.J., Cohen, T., Fryirs, K., Brooks, A. 1999. Post-European changes to the fluvial geomorphology of Bega catchment, Australia: implications for river ecology. Freshwater Biology 41, 839-848. Bryant, A.G. (1993). An evaluation of the habitat characteristics of pools used by platypuses (Ornithorhynchus anatinus) in the upper Macquarie River system. Bachelor of Applied Science (Hons) thesis. Charles Sturt University: Bathurst. Davies, P.E. and Cook, L.S.J. (2001). “Basslink Integrated Impact Assessment Statement. Potential Effects of changes to Hydro Power Generation’. Appendix 7: Gordon River Macroinvertebrate and Aquatic Mammal Assessment’. Ellem, B.A., Bryant, A. and O’Connor, A., (1998). Statistical modelling of platypus (Ornithorhynchus anatinus) habitat preferences using generalised linear models. Australian Mammalogy 20, 281-285. Ellem, B.A. and McLeod, A. 1998. Platypus (Ornithorhynchus anatinus) movement data from the Duckmaloi weir pool: Poisson regression models. Australian Mammalogy 20, 287-292. Elliott, J.M. (1977). “Some Methods for the Statistical Analysis of Benthic Invertebrates’. Freshwater Biological Association Scientific Publication 25, 1-150. Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT Evans, B.K. , D.R., Baldwin, J. and Gabbott, G.R.T. 1994. Diving ability in the platypus. Australian Journal of Zoology 42, 17-27. Faragher, R.A., Grant, T.R. and Carrick, F.N. 1979. Food of the platypus, Ornithorhynchus anatinus, with notes on the food of the brown trout, Salmo trutta, in the Shoalhaven River, New South Wales. Australian Journal of Ecology 4, 171- 179. Grant, T.R. (1982). Food of the platypus, Ornithorhynchus anatinus (Ornithorhynchidae: Monotremata) from various water bodies in New South Wales. Australian Mammalogy 5, 135-136. Grant, T.R. (1993). ‘The Bellinger River Water Supply Project Aquatic Studies - The Platypus’. (Report to Mitchell McCotter on behalf of the Coffs Harbour City Council and Department of Public Works by Mount King Ecological Surveys, Oberon). Grant, T.R. (1991). The biology and management of the platypus (Ornithorhynchus anatinus) in New South Wales. Species Management Report No. 5. NSW National Parks and Wildlife Service: Hurstville. Grant, T.R. and Bishop, K.A. (1998). Instream flow requirements for the platypus (Ornithorhynchus anatinus). An assessment strategy. Australian Mammalogy 20, 267-280. Grant, T.R. and Dawson, T.J. (1978). Temperature regulation in the platypus, Ornithorhynchus anatinus, production and loss of metabolic heat in air and water. Physiological Zoology 51: 315- 332. Grant, T.R., Griffiths, M. and Temple-Smith, P.D. (2004). Breeding in a Free-ranging Population of Platypuses, Ornithorhynchus anatinus, in the upper Shoalhaven River, New South Wales - a 27 Year Study. Proceedings of the Linnean society of New South Wales 125, 227-236. Grant, T.R. and Temple-Smith, P.D. 1998. Growth of nestling and juvenile platypuses (Ornithorhynchus anatinus). Australian Mammalogy 20, 221-230. Proc. Linn. Soc. N.S.W., 125, 2004 Harris, J. (1987). Growth of Australian bass, Macquaria novemaculeata (Perciformes; Perichthyidae), in the Sydney Basin, Australian Journal of Marine and Freshwater Research 38, 351-61 McLeod, A. L. (1993). Movement, home range, burrow usage, diel activity and juvenile dispersal of platypus, Ornithorhynchus anatinus, on the Duckmaloi Weir, New South Wales’ Bachelor of Applied Science (Hons) thesis. Charles Sturt University: Bathurst. Marchant, S. and Higgins, P.J. (eds)(1993). “Handbook of Australian, New Zealand and Antarctic Birds’. (Oxford University Press, Melbourne). Olsen, P. (1999). Winged pirates. Nature Australia 26, 30- 37. Rakick, R., Rakick, B., Cook, L. and Munks, S. (2001). Observations of a platypus foraging in the sea and hunting by a wedge-tailed eagle. Tasmanian Naturalist 123, 3-4. Richards, G.C. 1986. predation on a platypus, Ornithorhynchus anatinus (Monotremata: Ornithorhynchidae), by a goshawk. Australian Mammalogy 9, 67. Rohweder, D. (1992). Management of platypus in the Richmond River catchment, northern New South Wales. Bachelor of Applied Science (Hons) thesis. University of New England Northern Rivers: Lismore. Serena, M. (1994). Use of time and space by the platypus (Ornithorhynchus anatinus) along a Victorian stream. Journal of Zoology (London) 232, 117- 131. Serena, M., Worley, M., Swinnerton, M. and Williams, G.A. (2001). Effect of food availability and habitat on the distribution of platypus (Ornithorhynchus anatinus) foraging activity. Australian Journal of Zoology 49, 263-277. Smith, A.K. and Pollard, D.A. (1998). Policy guidelines. Aquatic habitat management and fish conservation. NSW Fisheries: Sydney. Young, W.J. (Ed) (2001). ‘Rivers as Ecological Systems: The Murray-Darling Basin’. (Murray-Darling Basin Commission: Canberra). 241 x oy pi s et : : i. ROE tr Ay a) BS" 2 em BV aN De YP ei x Space 2S ts - s : tye iy ‘ ne . 1 . 7 hak We ti aa y toma) : : " ) ¥e Pte | a ‘ * ieee N RO oP el Hi eas a " | Ae fn st ga Ce y val a Salam dante lacaikepitiainevih he seen “lent i vital ‘space Sa ileal ie : 9 ~~. 2 pt seabbutlgeo§ om iit pamabaectiur exe, ne iat cana opi akg ial : many Ls aera yh ty ZL REL marks. “lt nee phe. the aE Su at WE SPATE 6 SOU ( (eh 8 Saee pA, vety aia: AL Ly “ big ote Pare a me: Sir word : cana 5 Serient gusraseol Gee) oh A, are Ait 7 1G Teceaeitl Aieyp 70 Chae yt is fit, oa aa a ie ae a CUTTS Sys OeM icant Etnies ra rR Sua Ihig * ’ Awe SIE Ai Po PENS ere Heuwe hisieniad saat (ani Palbedisss i sane 3 = eee eae Bi ose “ier ian 3 u terneat : 4. te abr octal} HP Ab pe ehe | Ae deiyaiht dere erage ‘ _ecidonvoeo} ama ie Co. eae ih pasha bay. binidges eh gs secsagahess sic yt oiled Bilwiladd) ambos ae ae Sa Me eed tia Bet atid S30! 383) die oh sneer cape 8.1907 we Bis "ae t ars aT i ; cue ra Rew sh, At ; i tv +o?) 2 res e ee ee ee f es BST ciethabe sana eS A 2 iat sie sys ) vate era “4 geome: a a de a o Oe “ a a 4 pana ints iat s OF: thi nes ihe a ie ; sel ara a ba ict seosgerpi i sosieootns fea Od elebwbrs om ferit. i the pale wel aanaenonttr guampady inhonetiod ina 1 secre ey seeere sth Peach gers ts Lashueehodsa® . ace - : F ‘ot . preee itis ie if ¢ pet | Lasslsenh, 5, ie ss a ee 7 aa: te sive meee ais hay aaheayig Firs Moe Oe porate pail praises “g = | ¥ A eee eps Lipid. 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We > oe Kiet oh eats in vertcintiee, ) i ee Te f a Pe City iiical dbvatiaty oa Siebel F x a t - oe ® + «) ab Al 4 an a - é ’ lon iE ae nm, bark setiy ee ' oe 5 = | 242 m ; ; ~~. aay / a aie F ’ a < d ~ ay o- a tS Distribution of the Platypus in the Bellinger Catchment from Community Knowledge and Field Survey and its Relationship to River Disturbance DanteL Lunney!, ToM GRANT? AND ALISON MATTHEWws! ‘Department of Environment and Conservation (NSW), PO Box 1967, Hurstville, New South Wales 2220, Australia; "School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia. Lunney, D., Grant, T. and Matthews, A. (2004). Distribution of the platypus in the Bellinger catchment from community knowledge and field survey and its relationship to river disturbance, Proceedings of the Linnean Society of New South Wales 125, 243-258. Platypus distribution in the Bellinger catchment was investigated using a combination of field and community surveys. The field survey in 1996 consisted of netting and observations from the river bank and a canoe. The community-based wildlife survey consisted of a questionnaire and colour maps on which respondents were asked to mark the locations of sightings. Platypuses were observed or caught in 36 locations from all three rivers of the catchment. Two of the three platypuses captured were lactating females. The community recorded 123 locations of platypuses. The fact that the wildlife survey yielded similar results to the field surveys in identifying the location of individuals highlights the value of community records for platypus surveys. There were major floods in 2001, after which we contacted respondents who had reported seeing platypuses three years before. Of the 21 respondents who had been near the river since the flood, 7 had seen platypuses, principally in the tributaries of the Bellinger River. The habitat quality of the rivers was evaluated for platypuses and records were related to disturbance and rehabilitation. The species has survived in this system, but its future can only be assured by strategies which prevent further degradation of its habitat and institute proactive rehabilitation of the damaged sections of these streams. Manuscript received 18 August 2003, accepted for publication 8 January 2004. KEYWORDS: Catchment management, community wildlife survey, distribution, platypus, river management, wildlife. INTRODUCTION ; Grant (1992) and Grant et al. (2000) reported the distribution of the platypus Ornithorhynchus anatinus in New South Wales as having changed very little since the occupation of Australia by Europeans. Platypuses are considered common in the river systems of the coastal, tableland and western slopes (Grant 1991, 1992) and are frequently reported from streams flowing through agricultural land in these areas. In three separate surveys in New South Wales, 52-76% of recorded platypus sightings were from agricultural land (Grant 1991; Lunney et al. 1998; Rohweder and Baverstock 1999). The current study investigated the distribution of the platypus in a typical north coast river system, the Bellinger catchment, where the highland headwater streams arise in forested areas but grade into predominantly agricultural land (especially cattle grazing) towards the coast. The distribution of platypuses in the Bellinger-Kalang river system was investigated using both field and community surveys. Community surveys have been successful in identifying locations of platypuses (Lunney et al. 1998; Turnbull 1998; Rohweder and Baverstock 1999; Otley 2001). This survey was part of a wider community-based survey of the distribution of a number of key native wildlife species in the Bellinger and Kalang valleys adjacent to Bellinger River National Park. This study sought to assess the co-existence of typical rural community activities and wildlife species, including the platypus. The field observations and capture of platypuses were compared with the questionnaire reports for this and other native species in the catchment with the aim of testing the hypothesis that information gained from survey data provided by the community would be a reliable indicator of the presence of wildlife species in the area. Since our field and questionnaire study, Cohen et al. (1998) assessed the Bellinger-Kalang catchment using the River Styles framework (Brierley et al. 2002) and assigned conservation and PLATYPUS IN THE BELLINGER CATCHMENT rehabilitation priorities to various stream reaches. Using analysis of aerial photography and field observations, Cohen et al. (1998) assigned various sections of the Bellinger-Kalang catchment to a number of River Styles which have particular geomorphic attributes. The authors of the study stress that these categories are a “record of the character and behaviour of sections of river” and “are not a direct measure of river condition”. A separate set of procedures has been developed to appraise geomorphic river condition, building on attributes of river character and behaviour that are pertinent to any given River Style (Fryirs 2003). The attributes used in discerning the River Styles are shown in Table 1 and include channel planform and stability, morphology and geometry (depth and width), as well as descriptions of geomorphic units (e.g. pools, riffles, point bars), bed character (e.g. sand, gravel, cobbles) and vegetation character (including riparian vegetation and woody debris in the stream). Some of these attributes have been identified with the occurrence and foraging activity of platypuses and include: Channel geometry: pool depth has been positively related to the occurrence of the species (Ellem et al. 1998; Grant 2004), with platypuses often being observed foraging in water of greater than one metre depth and less than 5 metres. It has been suggested that foraging in shallow water can expose individuals to predation, especially from the introduced fox Vulpes vulpes (Grant and Denny 1993; Serena 1994; Anon. 2002). Geomorphic units: pool/riffle sequences have also been found to be associated with the presence (Rohweder 1992; Bryant 1993) and foraging (Serena et al. 2001) of platypuses, this probably being related to the benthic productivity of such geomorphic units (Hynes 1970; Logan and Brooker 1983; Boulton and Brock 1999). Bed character: the complexity of the bed substrate, including large particle sizes (rocks, cobbles, pebbles and gravel), has been positively related to both occurrence (Rohweder 1992) and foraging activities (Serena et al. 2001; Grant 2004) of platypuses, again probably resulting from greater benthic productivity (Hynes 1970; Marchant et al. 1984; Boulton and Brock 1999). Vegetation character: medium-to-large trees, especially indigenous species, are associated with the use of river reaches by foraging platypuses (Serena et al. 2001) and overhanging 244 vegetation has also been identified as a variable found in areas where platypuses are found (Rohweder 1992; Bryant 1993; Serena et al. 1998). This association between riparian vegetation and platypus occurrence is related to a number of important functions of such vegetation, including stabilisation of the bank, provision of cover from predators, supply of organic material to the food chains of the stream and shade moderating temperature variations, especially in summer (Riding and Carter 1992; Boulton and Brock 1999). The abundance of woody debris, included in the “vegetation character” attribute of Cohen et al. (1998), is also known to be positively associated with platypus occurrence (Rohweder 1992) and foraging (Serena et al. 2001), again probably being related to the complexity of habitats available for macroinvertebrates (Benke et al. 1985; Anon. 1998; Anon. 2000a). Cohen et al. (1998) also sorted sites in the Bellinger-Kalang catchment, grouping river reaches into five categories based on procedures outlined in Brierley and Fryirs (2000). These are summarised in Table 2 and are generally ranked from the least (conservation) to the most disturbed sites (degraded), although the “strategic” priority #2 sites were identified as being more disturbed than the priority #3 sites and were given a higher priority as they may impact on other sites downstream. This paper analyses these Rivers Styles and conservation/rehabilitation categories in relation to the data on occurrence of the platypus. We propose priorities for conservation and rehabilitation of the river system for the future survival of platypuses in the Bellinger and Kalang river system and in rural areas in general. Major floods in the Bellinger catchment in early 2001 provided an opportunity to assess the impact of floods on a known population of platypuses. A follow-up community survey was undertaken to determine the survival of platypuses post-flood. METHODS Study area The Bellinger is a fertile river valley on the north coast of New South Wales just south of Coffs Harbour, and includes the main townships of Bellingen and Urunga (Figure 1). The valley extends approximately 50 km inland from the coast at Urunga, and is approximately 20 km wide from Tucker’s Nob range in the north to the Bellbucca ridge in the south. Proc. Linn. Soc. N.S.W., 125, 2004 D. LUNNEY, T. GRANT AND A. MATTHEWS (F PIGEL :S66T ‘Te 19 UsYyoD) JUSUIYD}e9 SuLley-JOSUTT[Ig 94} UT 9[A}S IBANY Yee Jo saynqiajje d1yYd10UI0ey “[ a[qe], JSOLOJ Jeavis daap a[qeisun SATE IOUT “spose ‘oInjseg IOUNM “IIIs ‘pues seq jela}e] ‘sieq julod ‘sjoog WI ¢-Z ‘epiM WI OT - OF JeoHyouTUIAS Ayyeroues ‘peaiy) [301g TePLL seq [e1o}e] daep wi ¢-7 ajqeis poyejesea ‘sjood nods “SpA Ul CI-¢€ yeonjourmAsy Ajjeroues “peony ajsurs snonuljuoosip sugep Apoom rourur : *‘spooM “JSOIO} SANeU ‘oInjseg qJIS 0} pues : : Spuryst daep ul €-T aiqejsun peq- suqep Apoom 1ouro ‘seg jutod ‘saTyzu “sjoog “Opi WI 0S-0Z jeoWjourmAsy ATjeinjeu ‘peelyy ejsurg foaeid Sunopuem ‘SpooM “JSOIOJ SATIVUM ‘OINJseg — [AVIS 0} Se|qqoD poq-[earid suqep Apoom 1ourm pues Journ sieq [e1oje] doap WI OT-Z giqeisun SULOpueout- ‘spoom “jSOIO} SAIJeU ‘oINJseq “[OAPIS 0} SoTqGoD ‘seq julod ‘soyjz ‘sjoog ‘OpLM WH OZT-0€ Jeomjouussy Ajyeroues ‘pesry) e[su1Sg JOA [RIATITY yeomjaururAs Q} yeowjourmAse sure;dpooyy SUIpeOT “Te[NSaiIi 0} yeranyye sugep Apoom 9je1epour 0} pues rourox “SpURJSI ‘sIeq [e19}e] deep WOI-s 1e[nser wor SuisueI aiqeisun snonuljuoosip MOT ‘oINjsed pue JSoIOJ OATIVNT “TOABIS 0} SaTqqoD ‘sreq julod ‘sarzyi “sjoog ePrM Ul OOT-Or “aiqeuea Ayysiy Ayjeo0] “pesiy} epsurs IOAU yoIpeg jeAeis sIeq Jopinoq : SUIPeOT SiIqep Jour ‘ssqqqod — ‘seyqzu “sjood ‘sopijs ‘spidez doop mi ¢-Z } APOOM Je[NSoIIt “jsor0j AEN 0} Slopjnog ‘sopeoseo ‘sda}s yooipeg SPIM WU Op-0Z sJe[nsort 0} 1eynsoy 9[geIS ‘pealy} a]suIS 9310H SpUe]sI jouueYO - -plum ‘soyjyu ‘stood ‘spider SUIpPeO] SlIqop seyqqoo ‘sopl|s ‘sapeoseo yoorpeq doop WH Z-T yeomjeumuAse Apoom ysty ‘jsor0y oaTEN O} SIopinog ‘sdajs yoorpeq ‘syjeyIoieAA OpIA WI 0Z-O1 0} Je[nsery] Q1quIS “‘pealy} o[suIg wWeel}s IojeMpeopy Soy ‘sjood doep W ¢T ayqeisun YMoIder pue spoom ‘ainjseg ]OAeId 0} pues ‘sure[dpooyy pouters ony SpIM WH OT Jenga] Ayyeooy “peeryy opsurs tealjs pueydy, : = : Asojoqdiom AWyIqeys 1djJIBAVYD W0HL}959A 19798.1bY) pog S}lan I1qd.10010e5) ANouIossS jouUeyD jeuueyqsD pue aLi0jyueld jouueqD afA1g JAR wy N Proc. Linn. Soc. N.S.W., 125, 2004 PLATYPUS IN THE BELLINGER CATCHMENT Priority 1. Conservation sites 2. Strategic sites 3. High recovery potential 4. Moderate recovery potential 5. Degraded Nature of Sites least disturbed; river structure and vegetation relatively intact may be sensitive to disturbance or may affect sites downstream may show signs of natural recovery moderately degraded with reasonable potential for recovery highly degraded reaches with little natural recovery potential Table 2. Priority ranking of sites for river rehabilitation in the Bellinger-Kalang catchment (Cohen et al. 1998). Forested lands rise steeply from the valley, forming the extremely rugged fringe of the New England Plateau. The Bellinger Valley comprises the catchment areas of the Bellinger and Kalang Rivers, referred to in earlier maps as the North and South Arms of the Bellinger River. The third major river of the valley, the Never Never River, is a tributary of the Bellinger River, which it joins near Gordonville, about 10 kilometres upstream of Bellingen township. Tidal influence extends to Bellingen on the Bellinger River and as far as Spicketts Creek on the Kalang River (Cohen et al. 1998). The valley and floodplain was = DORRIGO Ji 2) rapidly cleared by the cedar-getters and during settlement in the mid to late 1800s (Anon. 1978; Lunney and Moon 1997). Now, the land is used primarily for dairying and beef cattle grazing, with small areas being planted for crops. The population of the valley was 12,253 in 1996, representing a population growth of 21 per cent over the previous 10 years (Anon. 2001a). The upper reaches of the streams of the Bellinger and Kalang valleys flow through steep forested areas in their headwaters, but degradation due to human activities, particularly clearing for /\_/ River [_] Bellingen LGA [|] National Parks <<--| State Forests wo *as* Canoe transects > Netting sites 4 Observation sites MN Figure 1. Location map of the Bellinger catchment, showing main features of the study area and the sections surveyed in the field work. 246 Proc. Linn. Soc. N.S.W., 125, 2004 D. LUNNEY, T. GRANT AND A. MATTHEWS Location Kalang River Jamisons Creek Date 14.12.96 Sex Female Age adult Length (cm) 39.0 Bill length (cm) 49 Bill width (cm) 42 Weight (g) 730 Spur 0 Milk (oxytocin) No -Kalang River Bellinger River Jamisons Creek Justins Bridge 14.12.96 15.12.96 Female Female adult adult 43.5 43.0 4.9 sell 44 43 905 900 0 0 Yes Yes Table 3. Details of platypuses captured during field survey in December 1996. agriculture, increases from the middle reaches to lower reaches upstream of the tidal limits at Bellingen and Spicketts Creek (Figure 1). Access of cattle to river banks has resulted in bank damage, especially in the lower Kalang River and the Bellinger River downstream of Thora. Parts of these sections of the rivers and their tributaries have good bank habitat for platypuses, but other sections are of lower quality due to the occurrence of natural gravel bars, to the effects of past gravel extraction, and to earth banks being cleared and/or damaged by cattle. The upper reaches of both these rivers and the Never Never River provide good platypus bank habitat, although some cattle damage to banks in parts of the upper sections of the Bellinger River and lower Never Never River was present at the time of the survey. Riparian vegetation is generally continuous on both banks of the upper reaches of all the streams in the system but becomes less continuous in the lower reaches of most streams. The Bellinger River between the Never Never River junction and Bellingen was considered to be the most degraded section of the system. River oak Casuarina cunninghamiana was the main native riparian species found, while exotic species — willows Salix sp., camphor laurel Cinnamomum camphora, privet Ligustrum sp. and lantana Lantana camara — were widely distributed in the riparian zones of most streams at the time of the study. Field sampling Field sampling was carried out over 10 days during December 1996. The sections of the system surveyed by canoe, bank observation and netting are shown in Figure 1. The dates of platypus captured are given in Table 3. Canoe and bank observations were made either in the two hours prior to darkness and/or the Proc. Linn. Soc. N.S.W., 125, 2004 two hours after dawn. Most of the Bellinger River from the mid-catchment gorge to Bellingen was surveyed by canoe either in the late afternoon or early morning (Figure 1). The Kalang River was unsuitable for canoe transects along much of its length due to its smaller size and the presence of obstacles in the channel. The section of river downstream from Duffys Bridge was suitable for use of the canoe and was surveyed a number of times. Live trapping of platypuses was carried out at four sites, one on the Never Never River, one on the upper Bellinger River and two on the Kalang River (Figure 1) using the methods of Grant and Carrick (1974). All captured females were injected with 0.2 ml of synthetic oxytocin (Syntocinon) to indicate the presence of lactation (Grant and Griffiths 1992). The Kalang River was less intensively sampled than the Bellinger section of the catchment due to its unsuitability for canoe transects and difficulty of access for observation at sites which appeared to represent good platypus habitat. However, netting was carried out at one downstream and one upstream site on the Kalang River to assess the accuracy of reports obtained from local residents during the survey and to compare with community reports from this part of the river system. Habitat assessment At a number of accessible sites (mainly at road crossings) on the Bellinger (15), Never Never (8) and Kalang (15) rivers the following data or rank scores were collected to provide an assessment of habitat characteristics known to be associated with the occurrence of platypuses and their use of an area (Rohweder 1992; Bryant 1993; Ellem et al. 1998; Grant and Bishop 1998; Serena et al. 1998, 2001). This scoring procedure was based on both published and PLATYPUS IN THE BELLINGER CATCHMENT unpublished field observations of platypus habitat: Habitat Category - this was a broad scoring of habitat suitability (1 best to 5 worst). Note: in the following categories, shade/shelter is usually provided by overhanging vegetation, but shade did not have to be present at time of observation as long as vegetation would provide shade/shelter at some times of the day. This is important not only to the platypus itself but to benthic invertebrate prey species: Category 1. EXCELLENT HABITAT - pools and/or riffle areas with >75% earth banks consolidated by roots of vegetation and providing significant shade/shelter, on both sides of river. Category 2. GOOD HABITAT - pools and/or riffle areas with 50-75% earth banks consolidated by roots of vegetation and providing significant shade/shelter, on at least one side of river or evenly distributed on both sides. Category 3. MODERATE HABITAT - pools and/or riffles with 25-50% earth bank consolidated by vegetation and providing a little shade/shelter. Category 4. POOR HABITAT - pools and/or riffles with 5-25% earth banks consolidated by roots of vegetation and providing little or no shade/shelter. Category 5. MARGINAL - pools and/or riffles with < 5% earth banks consolidated by roots of vegetation and providing no shade/shelter Riparian characteristics - these were features of banks that had been associated with platypus occurrence in other studies and were expressed as a percentage of sites at which they were present: - bank damage attributable to stock access; - bank damage attributable to floods; - presence of riparian vegetation; - presence of C. cunninghamiana (the most predominant native riparian tree species); - presence of introduced plant species in the riparian zone (especially willows, lantana, privet and camphor laurel). Community-based survey A community-based wildlife survey, in which the platypus was one of the target species, was posted to residents of the Bellinger-Kalang valley in December 1997. A total of 3000 survey forms was distributed by post to every household. There was a free-post return. The survey consisted of a questionnaire and colour maps on A3 size paper. The first colour map was a user-friendly map of the area where respondents to the survey could mark on it the locations of fauna, including the platypus, they had seen in the area. A grid was included on this map so that grid references could be determined with ease. These locations were then transferred to the geographical information system, ArcView, for analysis. The survey form, including the maps of the catchment, appear in Figure 2A&B. Relationship to River Styles To investigate the possibility that analysis of River Styles may be a useful method of predicting platypus occurrence or relative abundance in sections of a river system, platypus records from the field and community surveys were allocated by one of the authors (TRG, who has Provisional River Styler accreditation) to the various River Styles identified in the Bellinger System by Cohen et al. (1998). Stream reaches representing various River Styles from Cohen et al. (1998: Figures 1A and 9) were transposed onto the relevant 1:25000 topographical maps and the distances calculated using a manual map measure (Uchida Curvimeter). As only two platypus records were obtained from the mountain headwater streams and upland stream River Styles, these stream categories and observations were not included in the analysis. Observations of platypuses in streams which were not classified by Cohen et al. (1998) or were in the tidal sections of the rivers (one observation only) were also not considered. Relationship to river disturbance As was carried out for the River Styles categories, conservation/rehabilitation sections from Cohen et al. (1998: Figures 1A and 9) were transposed onto the relevant 1:25000 topographical maps and the distances calculated using a manual map measure (Uchida Curvimeter). Platypus records from the field and community surveys were allocated to the various conservation/rehabilitation categories. Post-flood survey In September 2001 we contacted those community members who had reported platypuses in the wildlife survey conducted three years previously. A letter was individually addressed to each respondent and contained a covering note, a questionnaire to gather information on post-flood platypus sightings and a map showing the results of the community and field locations of platypuses on which each respondent could mark recent sightings. Proc. Linn. Soc. N.S.W., 125, 2004 D. LUNNEY, T. GRANT AND A. MATTHEWS No Postage stamp required if posted in Australia Bellinger Valley Wildlife Survey Please fold and return to: Reply Paid 100 Bellinger Valley Wildlife Survey Segre AN c\- Dan Lunney > tO suE oN Biodiversity Survey and Research Division HOUSEHOLDER NSW National Parks and Wildlife Service a G PO Box 1967 One aS HURSTVILLE NSW 2220 Dear Shire Resident or Visitor, We are seeking your co-operation in conducting a wildlife survey of the Bellinger Valley. Its purpose is to locate wildlife populations as well as the habitats that are important for them. The long-term aim is to improve wildlife management of the valley by knowing which animals inhabit the area, where they occur, and the possible threats to their survival. This survey has the endorsement of Bellingen Shire Council and is supported by grants from the Heritage Assistance Program and the Foundation for National Parks and Wildlife. We would like you to fill out this survey even if you have only one wildlife sighting to record or you can only complete a part of the form. Also, if you have any historical information, this would help us understand the changes that have occurred to local wildlife populations over time in the Bellinger Valley. Please post your completed survey form (no stamp required) by 16 February 1998. Thank you for taking the time to assist us in compiling this community-based survey. If you would like a souvenir copy of this form, please tick the box on page 4. Dan Lunney Alison Matthews Dionne Coburn (02) 9585 6489 (02) 9585 6559 (02) 9585 6558 NSW NATIONAL New South Wales National Parks and Wildlife Service PARKS AND December 1997 WILDLIFE SERVICE Figure 2A. The Bellinger Wildlife Survey form. Proc. Linn. Soc. N.S.W., 125, 2004 249 PLATYPUS IN THE BELLINGER CATCHMENT vonal Parks and Witdife Service and its employees desclaim lability for any ack May 1997, © NSW Natfonal Parts and Wildlife Service - Map produced by K. Wala! This map Is not guaranteed (o be free of error or omission. Therefore, tht done or omission made Sn the information in the map and any conxeques Hesse stow fusing a aces &) cn the wep above theee pleces where you hve sen ayy of the wildlife listed helo. Beee wite tte imtials dom nest bo the anss to idrrify the gecies, Tf pile, plex alan pt te yer cf tte sigtting nee to te intials €o. ¥=S1 1563) Fleece also merck the location cf whee yo: ine oc Wolicey (tial). Green Tree Fing GF W ehetailed Sole W Enshtailed Hesample =P Soanna S Playps 1D flying Rox Fs Blue-tonged Lizard BT Echids E Fox FX Rastem Water Dream D Keala K Rabbit R Carpet or Diarond Pythn CP Bandicoot 2 Real Big PG L Q ‘This map is not guaranteed to be free of error or omission. Thertlore, the NSW Navonal Parks snd Wildlife Service and ts employes disclaim liability for any act done or © NSW National Parks and Wildlife Service. Printed by GIS Division December 1996 LANDSAT data supplied by Australian Centre far Remote Sensing, Image date 4 the information In the map and any consequences of such acts or omissions Benplles may include: Line around those areas on the photo above thet you Jrcw to be cod sites for wildlife. with a cood variety cf wildlife © ples vhee you see wildlife regularly heme you take visiting fiends to see wildlife © places with Icts of fog calls es Were yo. ee umn wildlife in the Line clearly using a dark pen oc percil thet will rt ante. Tr would aleo help us if you give the meseow for yor doioss in the space below: Blue-tongued Lizard Munvecon © NPWS Dern P. Sverre No, 11770206 Figure 2B. The Bellinger catchment map provided with the Wildlife Survey form. 250 Proc. Linn. Soc. N.S.W., 125, 2004 D. LUNNEY, T. GRANT AND A. MATTHEWS RESULTS Field survey Platypuses were observed or caught in all three rivers at a total of 36 locations (Figure 3). Two platypuses were captured at one of the two sites (the upstream site) on the Kalang River but none was observed in the limited sampling of this river by foot or by canoe. None was caught at the Never Never River netting site, but one was captured at the site on the upper Bellinger River. All the individuals captured were female, two of which were lactating, indicating the occurrence of breeding populations in both the Bellinger and Kalang rivers. All animals captured were within expected dimensions and body condition (Table 3). Platypuses were found to be common and continuously distributed along the Bellinger and Never Never Rivers, being captured or observed at 35 sites (Figure 3). The canoe transect survey method was most successful, yielding 2.2 animals per hour of observation, compared with 0.17 for both netting and observations by foot from river banks (Table 4). Community reporting of the occurrence of platypuses A total of 522 replies (17.4% return) was received to the Bellinger valley wildlife survey. Platypuses were recorded at 123 sites by the community-based survey. Only two platypus records were obtained from the headwater streams of the catchment and the field survey did not sample these streams. These data showed a much lower number of sightings (13) in the Kalang River and its tributaries than in the Bellinger River (110) and its tributary streams. The field and community-based data showed that the platypus is commonly found throughout the Bellinger River catchment, including the Never Never and Rosewood Rivers, and its distribution is probably continuous above the tidal limit at Bellingen to the headwater streams, which were not surveyed in this study. There was one report of a platypus downstream of the tidal limit on the Bellinger River. As well as being reported less often along the Kalang River eo | * Pass WK. DORRIGO- - a oS BGs we R L ge * 10 Kilometres 4 /\/ River (EJ Bellingen LGA [4] National Parks State Forests 4 Platypus field records * Platypus community records All community records \ y} ) } h 3 Figure 3. The location of field and community-based records of platypuses in the Bellinger catchment. Proc. Linn. Soc. N.S.W., 125, 2004 251 PLATYPUS IN THE BELLINGER CATCHMENT Method Kalang River Bellinger River Never Never River Total Hrs. No: “CPU Mrsag No, - CRU; e Eis 7 Now.) CPU CPU * Canoe Sey eal!) 0 95 29 MIB) Ja P0) 2D 2.20 observations | Bank 40 0 0 VAS ~ il 40 15 0 0 0.17 observations Netting 8.5 Z 0.23 45 1 0225725 0 0 0.17 Total 1625 "2 0.12 1425 31 Ped ls) SiS) = iS 0.65 0.98 *CPU: Catch/Obsetvation per Unit Effort Canoe = individuals seen/hour observation in each observation period Observation = individuals seen/hour in each observation period Netting = individuals captured per net hour (1x50m net in water for 1 hour) Total = division of total individuals seen/caught by total hours of observation or net hours Table 4. Success of various field survey methods used for recording platypuses in December 1996. system, platypuses seemed to be more discontinuous in their distribution in this part of the river system (Figure 3). Reliability of the community-based data set The distribution of community-based reports of platypuses in the Kalang and Bellinger components of the river system showed considerable overlap with the field records (Figure 3). There were few observations by the community outside the areas in which the field work identified the occurrence of the species. One exception to this was the section of the Kalang River between Moodys Bridge and Sunny Corner, where no sightings or captures were made during the field work, but where platypuses were reported by the community. Habitat assessment There was little difference among the 15 sites sampled on the Bellinger and Kalang rivers in terms © of habitat suitability (Figure 4), although on the Kalang River, 13% of the sites sampled were classified as 3 90 80 70 E 60 7/ Sp ) | Kalang = 50- j 'S) CO) Bellinger x 40-4 o Y YY Never Never 20 | ] ]/ 7 j 1 2 3 4 Bank Categories Figure 4. Percentage of sites at which bank suitability categories were recorded on the Kalang, Bellinger and Never Never Rivers. pay Proc. Linn. Soc. N.S.W., 125, 2004 D. LUNNEY, T. GRANT AND A. MATTHEWS 120 100 Yy, ]/ £80 2 _ i Kalang } 60 O Bellinger oy Never Never we 405 7 - 20 - _ Cattle Flood Riparian Casuarinas Introduced Damage Damage Vegetation. Plants Bank Characteristics Figure 5. Percentage of sites exhibiting various bank characteristics on the Kalang, Bellinger and Never Never Rivers. category 1, whereas none of the sites on either the Bellinger River or its major tributary, the Never Never River, fell into this category. All sites sampled had some riparian vegetation present but fewer sites on the Bellinger River had introduced species of riparian plants and more had C. cunninghamiana trees on the riverbank. More sites on the Kalang and Never Never Rivers than on the Bellinger River exhibited cattle and flood damage (Figure 5). Relationship to River Styles Table 5 details the lengths of each River Style, the total numbers of platypuses recorded in the field and community surveys and the numbers of platypus records per kilometre of each River Style. The River Styles of the Bellinger and Kalang sections of the catchment differ, with almost all (97%) of the Kalang River catchment being classified as confined bedrock with discontinuous alluvial floodplains, while the Bellinger River catchment contained a variety of Rivers Styles, ranging from 57% confined bedrock with discontinuous alluvial flood plains, through 24% alluvial with a meandering gravel bed to 10% and 9% of alluvial stream with a wandering gravel or discontinuous bed (Table 5). In the Bellinger River, there were Proc. Linn. Soc. N.S.W., 125, 2004 significantly more platypus records in the alluvial meandering gravel bed sections of river and fewer in the discontinuous alluvial stream than expected if platypuses were distributed uniformly across River Styles (77=26.64, 3d.f., P<0.01). In the Kalang River, platypus records were distributed evenly across River Styles. Expressed on the basis of platypus records per kilometre of river represented by each River Style, the Bellinger River had 0.95 records/km in the confined bedrock with discontinuous flood plains River Style, while the Kalang River (where this River Style made up 97% of the river downstream of the mountainous headwater reaches) had only 0.23 records/km (less than 25% of the value for the Bellinger). In the only other River Style represented in the Kalang River, alluvial river with meandering gravel bed (3% of the river), there were no platypus records and yet this was the River Style on the Bellinger River which had most records (2.1/km). Relationship to river disturbance The lengths of each conservation and rehabilitation priority reaches proposed by Cohen et al. (1998), along with the total numbers of platypus records from the field and community-based data, as well as the number of records per kilometre for each priority category in the Bellinger and Kalang 253 PLATYPUS IN THE BELLINGER CATCHMENT River Style Distance Platypus (km) Confined bedrock 56 with discontinuous floodplain Alluvial meandering gravel bed river Alluvial wandering gravel bed Discontinuous alluvial stream Total Bellinger River Kalang River Platypus | Distance Platypus Platypus per km (Km) per km See) 14 0.23 PA 0 0 0 0 0 0 0 0 Table 5. River Style distances, numbers of platypus records and numbers of platypuses reported per kilometre of river in the Bellinger and Kalang rivers and their tributaries in December 1996. catchments, were compared (Table 6). The number of platypus records per kilometre of river in the Bellinger River increased from the “strategic” category (0.74/ km) through the “high recovery potential” (0.97/km) and “moderate recovery potential” (1.80/km) categories to be highest in the most “degraded” (2.25/ km) section of the river. There were significantly more platypus records than expected in the degraded and moderate recovery potential categories (y7=17.99, 3d.f., p<0.01). In addition, the number of records of platypuses in the Bellinger River was much higher (1.22/km) than in the Kalang River (0.23/km), in spite of the fact that the latter system appears to be less disturbed than the Bellinger River. River Style Distance Platypus ~ Conservation Strategic High recovery potential Moderate recovery potential Degraded Total Bellinger River Post-flood survey A total of 43 replies was received from respondents who had reported platypuses in the 1997 survey. Twenty-one respondents had been near the river since the 2001 floods and of these, 7 had seen platypuses. Sightings of platypuses post-flood were in Hydes Creek (6 sightings), Boggy Creek (1 sighting), the Never Never River (1 sighting), the Kalang River between Duffys and Moodys bridges (1 sighting) and the upper Bellinger River between Diehappy and Bishops Creeks (2 sightings). Kalang River Distance Platypus Platypus per km Platypus per km 20.5 5 0.24 13.9 4 0.29 NS) 5 0.22 25h 0 0 Table 6. Conservation/rehabilitation priority distances, numbers of platypus records and numbers of platypuses reported per kilometre of river in the Bellinger and Kalang rivers and their tributaries in December 1996. 254 Proc. Linn. Soc. N.S.W., 125, 2004 D. LUNNEY, T. GRANT AND A. MATTHEWS DISCUSSION Data from the field and community surveys The results strongly support the hypothesis that the community-based data from this study are reliable. The field observations and captures of platypuses closely corresponded to community records. As a result of this correspondence, the field and community data were combined in the analysis of platypus occurrence in relation to River Styles and conservation and rehabilitation priorities. Further, a post-flood survey was able to be conducted because of the reliability of community records. The lack of sightings from the headwater streams of the whole catchment suggest a lack of observation, rather than the species not occurring in them. This is supported by the low reporting of other wildlife species in the community wildlife survey in these areas (Figure 3). It is assumed that platypuses would almost certainly occupy these sections of the catchment streams, although this was not determined by the field survey. The field and community-based data rank the Bellinger River part of the system as being more suitable for occupation by platypuses than the Never Never and Kalang rivers. The limited habitat data collected during the survey point to the Kalang River having less suitable platypus habitat than either the Never Never or Bellinger Rivers. The discontinuous distribution of the platypus in the Kalang River, especially between Moodys Bridge and Rosewood Creek, almost certainly identified poorer habitat conditions. As community reports of other wildlife Species were made along the Kalang River, the lack of platypus records from these sections means that the Species is not found or is uncommon in these sections of the river. Further, any temporary loss of individuals from an area, such as from a flood, should not affect an accurate determination of distribution if the community had observed them in these sections of the river at other times. This is one of the values of the community survey, namely it was not restricted to one point in time and it also considered historical records. Qualitative field observations of the Kalang River between Moodys Bridge and Rosewood Creek confirmed that this section had poorer habitat quality than other sections of the river, with considerable disturbance of the river banks due to depletion of the riparian vegetation and cattle access, as well as accumulation of sand in the river bed. The field survey did not allow an adequate explanation to be made of the differences between the Bellinger and Kalang sections of the river system in terms of the observed platypus distribution. While parts Proc. Linn. Soc. N.S.W., 125, 2004 of the Kalang were more degraded than sections of the Bellinger and Never Never rivers, platypus reports and field sightings were common in the Bellinger River between the Never Never River junction and the town of Bellingen, a river section which was also highly altered by bank clearing, stock damage to banks and past gravel extraction. One report was obtained of a platypus in the tidal section of the Bellinger River, close to the entrance of Connells Creek. Three reports of platypuses in the tidal section of the river were also made to the authors during the field study; one at Fernmount in the 1940s, another at the mouth of Hydes Creek in 1996 and one near the Old Butter Factory in Bellingen (which was said to be “recent’). Platypuses have occasionally been found in the sea (Fleay 1980; Connolly and Obendorf 1998) and in estuarine habitats, but such occurrences are irregularly reported and are considered unusual (Stone 1983; Grant 1991, 1999: Rohweder 1992; Hird 1993; Menkhorst 1995; Connolly and Obendorf 1998; Rakick et al. 2001). It seems unlikely that the species regularly occupies the brackish or saline waters of estuarine environments. Nothing is known of its abilities to osmoregulate under marine or brackish conditions or any need by the species to have access to fresh water to groom salt from the fur, as occurs in several species of otters (Kruuk 1995). Platypuses are known to consume a range of benthic invertebrates as food but insect larvae are the most common prey items (Faragher et al. 1979; Grant 1982). In a number of rivers along the coast of New South Wales, tidal influences and/or saline intrusion into the lower reaches results in the diversity of benthic macroinvertebrates beginning to change at the tidal limit from being numerically dominated by insect fauna to being dominated by Crustacea, including amphipods and isopods, with oligochaetes worms and gastropod molluscs also having greater representation (Anon.1993; Simon Williams, then of Australian Water Technologies, pers. comm.). This could affect platypus distribution in the lower reaches of rivers of coastal New South Wales. It is also known that increased conductivity impairs the ability of the platypus to locate moving prey items, particularly small invertebrates, using the electrosensory mechanisms in its bill (Pettigrew et al. 1998). Competition with benthic- feeding fish species, which do not enter the freshwater sections of rivers, and possible predation by larger fish species, may also be involved in the occurrence of platypuses being unusual in tidal areas. Relationship to River Styles The differences in distribution and numbers of platypus records between the two rivers were not 255: PLATYPUS IN THE BELLINGER CATCHMENT found to be related to the differences in River Styles between the two parts of the system. On the basis of our findings in the Bellinger catchment we consider that analyses using the River Styles framework (Brierley et al. 2002) will not successfully predict the occurrence of platypuses. However, methods integrating geomorphic and biological considerations could lead to a framework which may be capable of predicting the suitability of streams for occupation by the platypus. Such integration could also provide a better basis for river management and rehabilitation than arises from the consideration of either geomorphic or biological considerations in isolation. This approach has been called the “landscape ecology approach” by Tockner et al. (2002). Relationship to river disturbance This study has shown platypuses to be present in degraded habitat of the Bellinger catchment. However, it would be a mistake to be complacent about these observations and regard disturbances of rivers to be benign with respect to the platypus. Despite the common occurrence of platypuses in agricultural areas, there are strong indications that platypus distribution has been fragmented and/or their numbers reduced in the streams of the Eden area (Lunney et al. 1998) and in the Bega (Brooks and Brierley 1997), Thredbo (Goldney 1998) and Richmond (Rohweder and Baverstock 1999) rivers of New South Wales and in the Wimmera River system in Victoria (Anon. 1999, 2000b, 2001b). In each of these instances the changes have been mainly attributed to the effects of agricultural practices. Lack of reports of platypuses from the Kalang River in the disturbed section between Moodys Bridge and Rosewood Creek also point to a fragmentation of platypus distribution within this part of the Bellinger catchment. Lunney et al. (1998) attributed fragmentation of platypus populations in the Eden region (Bega Valley Shire) of New. South Wales to the effects of farming, particularly cattle grazing and the clearing of the riparian vegetation since 1830. Brooks and Brierley (1997) and Brierley et al. (1999) have detailed the effects of early agricultural practices in the Bega River valley of New South Wales, confirming that these practices were almost certainly responsible for the irreversible changes to that river system. However, Turnbull (1998) recorded the occurrence of platypuses in most of the rivers around Bombala, in the tableland headwater streams of the Bega and Snowy Rivers in 256 New South Wales, in spite of the area having been utilised for both cattle and sheep grazing for the past 160 years. Of the 11 platypus sightings made by respondents to the post-flood questionnaire in the Bellinger catchment, 8 were in tributary streams. This suggests that the tributaries act as refuge areas during extreme floods. The tributaries could also be important for this population if the main streams of the Bellinger River system experience further degradation. This latter suggestion is based on our data from the Bega River (Lunney et al. 1998) where historically platypuses were found in the lower reaches, but it is now so degraded, shallow, sandy and exposed, that it no longer supports viable platypus populations. Instead platypuses occur only in the more protected and less developed tributary streams of the Bega River system. Conservation and rehabilitation Considering that the distribution of this unique Australian species overlaps extensively with activities of rural communities, its conservation depends on the adaptive management of these activities. The species has survived the current environmental disturbances so far, but its future conservation can only be assured by strategies aimed at preventing any further degradation of its habitat in — these areas and by proactive rehabilitation of damaged sections of streams and a recognition of the possible importance of the tributary streams in retaining refuge populations of platypuses. ACKNOWLEDGEMENTS The authors are indebted to many people, particularly those who took the time to respond to the survey. We wish to acknowledge the contribution of P. Sherratt for the design of the community survey form; K. Weinman for production of the map and extra digitising of road and river systems that were not previously available on the National Parks and Wildlife Service geographic information system; and I. Dunn and G. Brierley for critical comments on the manuscript. Funding was provided by the Foundation for National Parks and Wildlife and a National Estate Grant. Platypuses were captured under licences issued by the NSW _National Parks and Wildlife Service (A184) and NSW Fisheries (Scientific Research Permit F84/1245) and under Ethics Approval from the University of NSW Animal Care and Ethics Committee (Animal Research Authority ACE 94/ 91). Proc. Linn. Soc. N.S.W., 125, 2004 D. LUNNEY, T. GRANT AND A. MATTHEWS REFERENCES Anonymous. (1978). Pioneering in the Bellinger Valley. (The Bellinger Courier-Sun, Bellingen). Anonymous. (1993). ‘Investigation of the invertebrate benthic fauna of the Bellinger River’. (Unpublished report for Mitchell McCotter and Associates. June, 1993). Anonymous. (1998). “Riparian management #7. Managing snags in rivers’. (Land and Water Resources Research and Development Corporation, Canberra). Anonymous. (1999). Wimmera update. Newsletter of the Australian Platypus Conservancy Ripples 13, 2. Anonymous. (2000a). ‘Snags. A valuable but scarce resource’. (CRC for Freshwater Ecology, Canberra). Anonymous. (2000b). 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The use of a community-based survey to determine the distribution of the Platypus Ornithorhynchus anatinus in the Huon River catchment, southern Tasmania. Australian Zoologist 31, 632-641. Pettigrew, J.D., Manger, P.R. and Fine, S.L.B. (1998). The sensory world of the platypus. Philosophical Transactions of the Royal Society of London B. 353, 1199-1210. Rakick, R., Rakick, B., Cook, L. and Munks, S. (2001). Observations of a platypus foraging in the sea and hunting by a wedge-tailed eagle. Tasmanian Naturalist 123, 3-4. Riding, T. and Carter, R. (1992). ‘The importance of the riparian zone in water resource management - a literature review’. (Department of Water Resources, Sydney). Rohweder, D. (1992). “Management of platypus in the Richmond River catchment, northern New South Wales’. (Bachelor of Applied Science (Hons) Thesis, University of New England Northern Rivers, Lismore). Rohweder, D.A. and Baverstock, P.R. (1999). Distribution of platypus, Ornithorhynchus anatinus, in the Richmond River Catchment, northern New South Wales. Australian Zoologist 31, 30-37. Serena, M. (1994). Use of time and space by platypus (Ornithorhynchus anatinus: Monotremata) along a Victorian stream. Journal of Zoology (London) 232, 117-131. Serena, M., Thomas, J.L., Williams, G.A. and Officer, R.C.E. (1998). Use of stream and river habitats by the platypus, Ornithorhynchus anatinus, in an urban fringe habitat. Australian Journal of Zoology 46, 267-282. Serena, M., Worley, M., Swinnerton, M. and Williams, G.A. (2001). Effect of food availability and habitat on the distribution of platypus (Ornithorhynchus anatinus) foraging activity. Australian Journal of Zoology 49, 263-277. Stone, G.C. (1983). ‘Distribution of the platypus, Ornithorhynchus anatinus, in Queensland’. (Queensland National Parks and Wildlife Service, Brisbane). Tockner, K. Ward, J.V., Edwards, P.J. and Kollman, J. (2002). Riverine landscapes: an introduction. Freshwater Biology 47, 497-501. -Turnbull, R.W. (1998). Distribution of the platypus (Ornithorhynchus anatinus) in the Bombala River catchment, South-Eastern New South Wales. Australian Mammalogy 20, 251-256. Proc. Linn. Soc. N.S.W., 125, 2004 Reducing the By-catch of Platypuses (Ornithorhynchus anatinus) in Commercial and Recreational Fishing Gear in New South Wales T.R. Grant!, M.B. Lowry’, Bruce PEASE”, T.R. WALFORD? AND K. GRAHAM? ' School of Biological, Earth and Environmental Sciences, University of New South Wales, Kensington NSW 2052. email: t.grant@unsw.edu.au * NSW Fisheries P.O. Box 21 Cronulla, NSW 2230. Grant, T.R., Lowry, M.B., Pease, B., Walford, T.R. and Graham, K. (2004). Reducing the by-catch of platypuses (Ornithorhynchus anatinus) in commercial and recreational fishing gear in New South Wales. Proceedings of the Linnean Society of New South Wales 125, 259-272. The problem of platypus by-catch mortality in the eel, yabby and carp trap fisheries in New South Wales is reviewed, and the results of several experiments to determine the effectiveness of gear modifications to reduce platypus by-catch are presented. Entrance screens with 50-60 mm openings prevented the entry of platypuses into eel or yabby traps. Larger screens were not effective as a deterrent to platypuses entering traps. By-catch of platypuses in the eel fishery can be minimised by restricting traps to estuarine areas, where platypuses seldom occur, and by providing air spaces in the cod ends of traps used in impoundments and farm dams. Prohibiting the use of yabby traps in areas where platypuses are known to occur provides the most practical protection against by-catch of platypuses in this fishery. Platypuses were unable to exit from prototype carp traps, designed to permit escape of air-breathing species, but the provision of appropriately-sized openings at the base of the entrance funnels in these drum traps permitted platypuses to escape. Manuscript received 4 September 2003, accepted for publication 24 November 2003. KEYWORDS: by-catch, carp, eel, fishing, Ornithorhynchus anatinus, platypus, yabby. INTRODUCTION By-catch mortality of air-breathing vertebrates, including several species of freshwater turtles and diving birds, water rats (Hydromys chrysogaster) and platypuses (Ornithorhynchus anatinus), has been recognised for some time as a significant problem in various inland fisheries in Australia (Jackson 1979; Beumer et al. 1981; Grant 1991, 1993; Grant and Denny 1991; Leadbitter 2001). Such by-catch mortality of platypuses is of particular concern in small streams, where multiple drownings of breeding individuals have the potential to impact severely on small local populations. For example, an abandoned fyke net in a tributary of the Gellibrand River in Victoria was found to contain the skeletons of 17 platypuses (Serena 2003). There has often been conflict between the desires of fishers to maximise catches of their target species, and the implementation of effective methods to reduce non-target by-catch. This has resulted in a diverse range of regulations enacted by fishery authorities and voluntary gear modifications by fishers aimed at reducing the mortality of non-target species (e.g. Leadbitter 2001). Unfortunately, little research or monitoring has been done to assess the effectiveness of voluntary and regulated gear modifications. An historical assessment of inland fishing in New South Wales showed that commercial fishing probably resulted in significant platypus mortality when small-mesh nets were used (Grant 1991, 1993; Grant and Denny 1991). No commercial or recreational fishery using nets or traps to capture native fish species or salmonids in freshwater sections of coastal rivers is now permitted in New South Wales (NSW), but there is a commercial eel fishery based on the use of baited traps in estuaries, farm dams and a few large impoundments. West of the Great Dividing Range, the commercial fishery for native fin-fish species was phased out in 2001. Fishers previously involved in that industry have been encouraged to fish for yabbies, mainly (Cherax destructor), using “Opera house” traps (Rankin 2000). The introduced carp (Cyprinus carpio) is also targeted by commercial fishers using a variety of gear, including traps, mesh and haul nets and electrofishing. There are a number of options to prevent or minimise mortality of air-breathing wildlife species in traps. The most direct way is to ban fishing in areas where these potentially vulnerable species occur. REDUCING BY-CATCH OF PLATYPUS UNREGULATED FISHERY PREVENT BY-CATCH REGULATE TO CLOSE WATERS TRAP MODIFICATION NECESSARY BY-CATCH REDUCTION DEVICE MINIMISES BY-CATCH MORTALITY MODIFY TRAP TO PERMIT NON-TARGET SPECIES TO ESCAPE MINIMISE BY-CATCH REGULATE FISHING METHODS STANDARD TRAP MINIMISES BY-CATCH MORTALITY BY-CATCH REDUCTION DEVICE NOT APPROPRIATE OR IS INEFFECTIVE MODIFY TRAP TO PROVIDE NON-TARGET SPECIES WITH AIRSPACE Figure 1. Schematic diagram of possible options available to achieve by-catch reduction of air-breathing species in fisheries operations. However, maintaining a commercial fishery, while still. addressing the issue of by-catch mortality, is to adopt capture methods which minimise by-catch. Mortality of air-breathing non-target species can be reduced or prevented by trap modifications, such as fitting devices 260 to keep non-target species out (By-catch Reduction Device - BRD), providing a route to let them escape or permitting access to an airspace once they have entered a trap. Figure 1 summarises these possible options, which need to be explored in relation to the following issues: Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM Fishery requirements. The practicalities and economics of the fishery, in terms of trap design and cost, catch per unit effort, size of target species, and even the necessity to hide traps from possible interference and/ or vandalism must be considered. For instance, a device which reduces by-catch but unduly restricts the entry of the target species into a trap may be economically unviable. Behaviour of target species. It is necessary to know the reactions of the target species to trap modifications provided for non-target species. For example, the target species may escape via holes provided for the non- target species, or its behaviour could prevent the non- target species from utilising air spaces or escape routes provided. Behaviour of non-target species. In fishing areas where a number of potential by-catch species occur, escape holes, BRDs or air spaces in traps may not be suitable for all potential non-target species. For example one Species may use an escape hole in a trap which will not be used by another species. This paper reviews past efforts to reduce the mortality of platypuses in the eel, yabby and carp fisheries and reports on a number of recent studies carried out to assess the effectiveness of trap modifications designed to reduce by-catch mortality of this species in these fisheries. The three fisheries are reviewed in separate sections of the paper and the experiments pertinent to each are discussed within these sections. THE EEL FISHERY IN NEW SOUTH WALES Freshwater eels were initially captured in upper estuarine waters of NSW as a by-catch of other fisheries. A fledgling industry targeting eels, based on the use of traps, was established in the early 1980s. At that time prices for eels were low but in the late 1980s and early 1990s a high-value export market to Asia was established. This increased interest in the fishery and the adoption of potentially more productive fishing methods. Requests were made by fishers to extend their operations into freshwaters using fyke nets (Figure 2a), which were known to be involved in the mortality of air-breathing wildlife species in the eel fisheries both in Tasmania and in Victoria (Jackson 1979; Beumer et al. 1981; Grant 1991). The potential fishers drew attention to a brief experiment in Lake Crescent and Dee Lagoon in Tasmania, where two fyke nets screened with 100 mm square mesh grids, and two unscreened control nets, were deployed in those lakes for six days. During that time, two platypuses were captured in the unscreened nets but none were captured Proc. Linn. Soc. N.S.W., 125, 2004 in the ones with the screens in place (Grant 1991). While it appeared from this very limited experiment that a 100 mm mesh screen may have been effective in reducing platypus by-catch in Tasmania, an experiment done in the upper Shoalhaven River did not support this contention (Grant, unpublished data). Six platypuses (two female and four male) were placed separately between the river bank and the wing of a fyke net with a 100 mm mesh entrance screen in place. Two of these animals moved off after bumping the mesh and did not enter the fyke net but the other four either passed straight through into the net, or did so after first investigating the screen. At the time it was also known that elevating the cod end of fyke nets above the surface was effective in permitting platypuses to breathe and survive capture (Jackson 1979; Beumer et al. 1981; Grant 1991; Figure 2b). Unfortunately professional fishers were unprepared to do this, as they feared their catch could be stolen and/or their equipment vandalised if it was visible above the surface. As a result of the brief experiment with the Shoalhaven River platypuses described above, and advice from experts in the other states regarding the poor compliance of fishers to fit BRDs and/or to raise the cod-ends of their nets above the water level, the request by fishers to use fyke nets for eels, and to extend the fishery to freshwater streams was denied by NSW Fisheries. Instead, the fishery was restricted to estuarine waters, a limited number of impoundments and private farm dams, using baited traps without wings to direct animals~into the traps (NSW Fisheries Eel Policy Document, May 1992). The standard eel traps used in the fishery are shown in Figure 2c. They consist of a metal rod frame 50 cm wide by 40 cm high by 90 cm long covered with 30 mm mesh polyethylene netting. The single entrance funnel (or ‘valve’) is located in one end of the trap. The opening in the funnel consists of a hole in the netting stretched firmly into a 100 mm wide slot, and pulled approximately 20 cm into the trap. The traps used in estuaries have a 1.5 m long cod end (bag with a draw-string) on the opposite end of the trap from the entrance funnel. Those used in freshwater impoundments and farm dams are similar to the estuary trap, but have a 5 m long cod end. A 150-200 mm diameter float is fastened inside the cod end near the draw-string and from one to three 50 cm diameter aluminium hoops are fastened to the inside of the cod end to keep the passage to the surface open. These traps are normally baited with frozen pilchards or mullet to attract eels. In the late 1990s anecdotal reports to the National Parks and Wildlife Service, NSW Fisheries and one of the authors (TRG) indicated that platypuses 261 REDUCING BY-CATCH OF PLATYPUS \) \) Wy! NIN INNING DINU PHN \) ( cod end entrance funnel Figure 2. (a) Fyke net used in eel fisheries in Tasmania and Victoria. (b) Commercial eel trap used in the impoundment or farm dam eel fishery, showing the elevated cod-end creating an air space. (c) Typical eel trap used in the tidal estuary fishery in New South Wales. Note the entrance funnel or ‘valve’ which permits animals to enter the traps in one direction (wide outside to narrow inside). Animals are unable to locate the narrow inside entrance to escape. In Experiment 1 grids were placed at the narrow end of the funnel and in Experiment 2 at the wide end. 262 Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM were being drowned in eel traps, not only in the upper reaches of some estuaries (where tidal influence changed with river discharges) but also in farm dams and impoundments (where air spaces were not consistently being maintained in the cod ends of traps). As aresult, the following experiments were undertaken to determine if it was possible to reduce this mortality of platypuses by trap modification. EXPERIMENT 1 - Investigation of grid sizes for a platypus exclusion device The objective of this experiment was to determine the optimum grid size for excluding most platypuses from eel traps. The experiment was conducted in two pools on the Wingecarribee River in New South Wales from 17-19 February 2000. Methods The entrance funnels in eight standard eel traps were fitted with grids of different sizes. Each grid was a square divided into four equal openings; the openings in these grids ranged from 55 to 90 mm, in 5 mm increments. The plastic material used to make the grids was reinforced with lengths of 3 mm wire. The traps were fastened end to end (in order of decreasing grid size) and placed on a flat sandy area in the pools where platypuses were to be captured (Figure 3a). Water depth varied between traps but all had an airspace to allow the platypuses to breathe during the experiment. Trials were done in different pools on two days. Platypuses were captured using unweighted gill nets (Grant and Carrick 1974) during the evening or morning. Once the required numbers of platypuses were captured, each individual was measured and weighed, then tested individually in the experiment. Platypuses were placed through an access door into the first trap leading into an entrance funnel with the 90 mm grid in place (Fig. 3a). Red-filtered lights were used to observe the animals at night, as observations in captivity indicated that platypuses are less responsive to disturbance under red light illumination (Grant, personal observation). The time that animals remained in each trap before passing through each grid was recorded, along with the number of attempts that each animal made to pass through the entrance funnel into the next trap in the series. Animals were removed from the experiment and released immediately if they remained in any trap for more than 15 minutes. Results A total of ten platypuses were used in the trials, comprising two adult males (1190 and 1760 g), Proc. Linn. Soc. N.S.W., 125, 2004 six adult females (890-1060 g) and two juvenile females (700 and 760 g). Data are summarised in Table iL. Trial 1: Animals tested at night were reluctant to pass through the 85 mm grid and none passed through the 75 mm grid, while a single female captured in the morning, and tested in daylight readily, passed through all grid sizes, although exhibiting some delay at the 80 and 70 mm grids. However, it was noted that the traps with 85-70 mm grids, which were apparently difficult for the animals to negotiate, were located in slightly shallower water than the rest of the traps. The water level in these traps was located at or just above the top of the grid, whereas the water level in the other traps was well above the top of the entrance grids. It was thought that this difference in water depth may have influenced platypus behaviour. Subsequently, all traps were placed in deeper water (well over the top of the grid) during the second trial. Trial 2: The largest male (1760 g) could not pass through the 65 mm grid, but the smallest female (700 g) passed through each grid in less than | minute. The two slightly larger females did not initially pass through the 55 mm grid. However, it was found that, due to some unevenness on the bottom of the pool, the trap with this grid was in slightly shallower water than the preceding traps in the series. After moving this last trap to a position in slightly deeper water, animals passed through the 55 mm grid almost immediately. The data from Trial | indicated that there was a greater reluctance for platypuses to negotiate the grids when the traps were less submerged. However, Trial 2 confirmed that female platypuses of up to 1 kilogram in weight could pass through a 55 mm grid. Animals smaller than 1 kg passed through easily, while the 1 kg female had a tighter squeeze. Only one male platypus was captured for use in Trial 2. This was the largest animal tested (1760 g) and was stopped by the 65 mm grid. A grid between 55 and 65 mm would apparently be required to exclude most adult male platypuses. EXPERIMENT 2 - Investigation of possible avoidance of entrance grids by free-swimming platypuses In Experiment 1, each platypus was closely confined inside the traps so there was an imperative to find an escape route. However, two of the four animals in Trial 2 hesitated, and made more than one attempt to pass through the 70 mm grid, indicating possible deterrent effect of this grid size. Experiment 2 was designed to test whether grids across the outer end of the entrance funnel (Figure 3b) deterred foraging REDUCING BY-CATCH OF PLATYPUS entrance funnels 95. RRS WY netting enclosure \) \) i wy water level Ra FM MK Figure 3. Set up used in Experiments 1 and 2 to test the effectiveness of by-catch reduction devices (BRDs) on entry of platypuses into eel traps. (Top) Experiment 1. Traps were attached together in a line with grids of different sizes at the narrow end of each entrance funnel or ‘valve’. (Bottom) Experiment 2. Mesh enclosure in a river pool with trap entrance attached. Note the position of the replaceable rectangular grid across the outer (wide) entrance of the funnel. platypuses from entering traps. The experiment was done in a pool on the upper Shoalhaven River in the southern tablelands of New South Wales from 17-19 March 2000. } Methods A circular enclosure, 1.5 m high x 3 m diameter, made from 10 mm mesh monofilament gill net material, was constructed in a pool between the two netting sites where platypuses were captured for the experiment. The enclosure was designed so that the only possible escape for a platypus was through the grid of the entrance funnel of a trap inserted in the enclosure wall. Square grids, made from 4 mm steel rods, with 50, 60, 70 and 80 mm openings were used in this experiment. Each trial was done by attaching a 264 replaceable grid to the entrance funnel of the trap, then placing a platypus into the enclosure (Figure 3b). At night, red-filtered lights were used to observe the animals. The time each animal remained in the enclosure before passing through the grid was recorded, along with the number of attempts that each made to pass through the grid. If an animal did not pass through a particular grid in the test series, this was replaced by the next larger grid in the series and the observations repeated. After the first animal was obviously unable to exit the 50 mm grid, the trials on all others were begun with either the 60 or 70 mm grid. Results Eight relatively small platypuses (ranging in Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM Table 1. Details of platypuses exiting through the various grid sizes within the funnels of eel traps in the two trials of Experiment 1. + = animal exited specific grid size; X = platypus did not exit through specific grid size. Sex/ Weight 90mm 85 mm age (g) Grid Grid Grid Trial 1 Male Adult 1190 + x x Female Adult 1060 + + x Female Adult 1030 + x XxX Female Adult 1020 + + a Female Adult 920 + + + Female Adult 890 + + + Exited 6/6 4/6 3/6 Trial 2 Male Adult 1760 + + + Female Adult 1000 + + + Female Juvenile 760 + + + Female Juvenile 700 + + + Exited 4/4 4/4 As4 size from 500 to 940 g) were tested in the enclosure at night. Results of the grid-deterrent trials are shown in Table 2. The first platypus was initially placed in the enclosure with the 50 mm grid. After six attempts to go through the grid it was apparent that the animal would not fit through the spaces. After several tentative attempts at the 60 mm grid it appeared to stop trying to escape through the subsequent grids and remained in the enclosure even after the largest grid was completely removed. The test with the second platypus was started with the 60 mm grid in place, but this platypus was less active than the first animal and made only one tentative attempt to pass through this grid. It then readily passed through the 70 mm grid after only one attempt. Trials with the next three platypuses were all started with the 60 mm grid. All three of these animals swam past the grid at least once before escaping through it. The last three animals were initially trialed with the 70 mm grid, and all passed Proc. Linn. Soc. N.S.W., 125, 2004 80mm 75 mm 70mm 65mm 60mm 55 mm Grid Grid Grid Grid Grid x x x x x x x x x x x x x x x x x x x x + + + + + x x x x x 1/6 1/6 1/6 1/6 1/6 + + x x x + + + + + + + AP + x + + + + + 4/4 4/4 3/4 3/4 2/4 through it at the first attempt. Overall, two animals out of five appeared to be deterred by a 60 mm grid (40%) and only a single animal was deterred by a 70 mm grid (Table 2). EXPERIMENT 3 - Platypus behaviour in the elevated cod ends of traps modified for use in farm dams and impoundments The objective of this experiment was to record the behaviour of platypuses in modified eel traps used in impoundments and farm dams (Figure 2c) and to investigate their ability to negotiate the long cod end extension to the air space. The experiment was done in a pool on the upper Shoalhaven River from 17-19 March 2000. Method Two impoundment eel traps, with 5 m cod ends (Figure 2c) were placed in a pool of 0.5 m depth. REDUCING BY-CATCH OF PLATYPUS Table 2. Details of platypuses deterred from entering the various grid sizes across the entrances of eel traps in Experiment 2. Animals are arranged in the order in which they were used in the experiment. + = animal passed through specific grid size; X = platypus did not pass through specific grid size i.e. deterred; - no data; Sex/Age Weight 50 mm 60 mm 70 mm 80 mm (g) Grid Grid Grid Grid Female Adult - 800 xX x x x Female Juvenile 500 - x + - Male Juvenile 800s - + = = Male Juvenile 740 - + - - Male Juvenile 640 - + - - Female Adult 940 —- - + - Female Adult 900 =- - + - Female Juvenile 690i; - + - One trap had three evenly spaced hoops in the cod end and the other had only one hoop near the airspace. The cod end of each trap was stretched and tied off above the surface of the water to a star-picket. Three platypuses (one male and two females) were placed consecutively in the trap with three hoops, and one female platypus was placed in the trap with one hoop. Each platypus was observed for 15 to 20 minutes before being released. Results ' In each case the platypus spent several minutes searching the inside of the trap before travelling up the cod end to the airspace. Each took several breaths then travelled to the trap where it again searched around or ‘wedged’ itself under the entrance funnel. Within five to eight minutes each would again travel up to the airspace for several breaths before returning to the trap. Platypuses travelled back and forth from the trap to the airspace 2-3 times during the 15-20 minutes they were confined in the trap. ; DISCUSSION - Eel Trap Experiments The results of Experiments | and 2 indicated that a grid of 50-55 mm would be necessary to exclude platypuses from entry into eel traps. Such a by-catch reduction device (BRD) would almost certainly affect the catch rates and sizes of eels (Koed and Dieperink 1999). This would be unacceptable 266 to commercial fishers, particularly those fishing for adults of the long-finned species (Anguilla reinhardtii). Free-ranging platypuses may be deterred from entering traps fitted with external grids of 70 mm or less across the entrance funnels but such screening would be unlikely to significantly reduce platypus by-catch in eel traps. Raising the cod end to provide an air space would facilitate the survival of platypuses captured in eel traps fitted with elongated cod ends. Platypuses captured in these traps were reluctant to stay at the surface and preferred to remain submerged in the trap between taking breaths. This behaviour, which minimises the time spent at the surface, may be a mechanism to avoid natural predation. Because platypuses must breathe at least every 2-10 minutes (Bethge 2002), captured individuals would need to travel back and forth to the airspace many times during any extended period of confinement after capture. This would be stressful and energetically demanding. It is essential that captured animals be released as soon as possible after capture. Studies using fyke nets (with elevated cod ends) to capture fish have shown that platypuses can survive for periods of up to 24 hours (Grant and NSW Fisheries, unpublished data). However, hypothermia has been reported in platypuses restrained in fyke nets after a few hours in cold conditions (Serena, personal communication). The current regulations in New South Wales demand that eel traps be inspected at least every 24 hours. The platypus forages aerobically for short periods by holding its breath, following a comparatively large inspiration of air after each dive (Evans et al. 1994; Bethge 2002). The behaviour observed in this study of ‘wedging’ themselves under an object, and reducing energetic demands by remaining stationary, has been reported in captivity to last up to 11 minutes (Evans et al. 1994; Bethge et al. 2001; Bethge 2002). The function of this behaviour and its occurrence in the wild has not been determined. However, from the perspective of by-catch mortality this behaviour would not prevent platypuses from being drowned in completely submerged traps during normal fishing operations, which demand a period of trap submergence of hours rather Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM than minutes. Observation of traps with airspaces maintained only by the use of a float has shown that the cod end can easily become twisted or bunched. This situation would undoubtedly prevent a captured air-breathing species from reaching the airspace. This can be avoided by stretching the cod end tightly to a fixed point, either on the bank or a star picket driven into the bottom of the water body. It should be noted however, where traps are set with elevated cod ends attached to a fixed point, allowance needs to be made for anticipated rises in water level as a result of rainfall and/or tidal influences. Attachment of the cod ends of eel traps to a fixed point is mandatory under regulations for the use of eel traps in impoundments and farm dams in NSW. THE COMMERCIAL AND RECREATIONAL YABBY FISHERY IN NEW SOUTH WALES The results of the experiments done to evaluate the effectiveness of devices to prevent or deter platypuses from entering eel traps are also directly applicable to both the commercial and recreational “yabby’ [freshwater crayfish] fisheries. Based on the lack of adverse reports and on the assumption that the traps used to capture yabbies were small and did not have mesh wings to direct foraging platypuses into them, Grant (1993) suggested that “yabby fishing poses little threat to platypuses”. This conclusion is now thought to be incorrect, as anecdotal reports from a number of states suggest that yabby traps were affecting some local platypus populations. These traps have also been implicated in the mortality of other non- target species, especially freshwater turtles. The drowning of as many as five platypuses in a single yabby trap has been reported, although the species’ attraction to these traps is not fully understood. Platypuses are known to locate their prey by sensing the electrical fields generated by muscular activity of the prey species, especially large food items such as yabbies (Pettigrew et al. 1998). A trap containing live yabbies may therefore attract platypuses during their normal foraging activities. Once there is a dead platypus in a trap, more yabbies may feed on the decomposing carcass, which could in turn attract other platypuses into the trap. Rankin (2000) suggested that a fixed ring 60- 70 mm in diameter may prevent platypuses from entering traps and also facilitate their escape. Some commercially available yabby traps are fitted with 90 mm entrance rings, which are effective in excluding larger turtles but which are still reported to have Proc. Linn. Soc. N.S.W., 125, 2004 drowned platypuses. The experiments described above for eel traps indicate that a 90 mm diameter ring is too large to exclude platypuses. Similarly, neither the experiments reported here nor anecdotal observations support Rankin’s (2000) suggestion that platypuses could escape by returning through a fixed entrance ring. Allanson and Thurstan (1999) evaluated the effect of entrance rings of different diameters in yabby traps using relatively small captive-bred yabbies (Cherax destructor). These trials showed that the smallest ring tested (63 mm) still permitted yabbies of the same size to enter the experimental traps as were entering the control traps with no rings fitted. However, the experimental traps caught substantially fewer yabbies. When the results of Allanson and Thurston’s (1999) experiments were discussed with commercial fishers, it was concluded that the use of such a small entrance ring was not a viable option for the commercial yabby fishery. Current regulations in New South Wales exclude the use of traps in commercial and recreational yabby fishing from known platypus waters and 90 mm rings are required in all yabby traps to exclude most turtles. Closed waters are located east of the Newell Highway, from the Victorian border (Murray River) to the Queensland border (Macintyre River), along with local closures around Deniliquin on the Edward River, Echuca on the Murray River and between Narrandera and Darlington Point on the Murrumbidgee River, where platypuses are also know to occur. THE CARP FISHERY IN NEW SOUTH WALES Carp (Cyprinus carpio) were probably first introduced into Australia around 1850 but did not spread until the introduction of the ‘Boolarra’ strain in the 1960s. Ecological effects of high densities of carp are poorly understood, but increased bank damage, disturbance of aquatic macrophytes and turbidity are all possible consequences. The overall disruption of riverine food webs by the large biomass of carp is thought to be detrimental to freshwater ecosystems (Schiller and Harris 2001). Carp are harvested in New South Wales using a variety of gear, including traps, haul and mesh nets, and electrofishing equipment. There is considerable overlap between the distribution of carp and platypuses (Boulton and Brock 1999), making the use of submerged traps a concern in this fishery. A drum trap was constructed by NSW Fisheries (Fig. 4), which was designed to permit the escape of air-breathing vertebrate species, including 267 platypus escape holes ~ &£ y SY 3 (3 err y) ERTS y E ZO: Neate Gs \ ye >, ” 0 Y | ») REDUCING BY-CATCH OF PLATYPUS turtle escape hole Y entrance funnel escape hole platform entrance funnel Figure 4. (Top) Modified drum trap showing escape hole in the roof above the mesh platform. Note the entrance funnel (or ‘valve’) on the left end of the drum. The entrance was sealed in the experiments and the triangular escape holes were made at the base of this funnel. (Bottom) Inside the trap showing the position of the steel mesh platform below the escape opening. vertebrate species could pass through the 8 cm gap between it and the roof of the trap and exit through the escape hole, while larger carp would not be able to escape. Carp are also inclined to congregate near the bottom of a trap. The design assumed that air- breathing species would tend to swim towards the surface and search along the roof of the trap for a means of escape (surface/ search behaviour). The objective of the following experiment was to test the effectiveness of the escape device for platypuses. EXPERIMENT 4 - Assessment of escape of platypuses from a prototype carp trap Platypuses close their eyes, ears and nostrils when under water, using the sensory mechanisms in their bills to find their way around (Pettigrew et al. 1998). It was expected that platypuses in the experiment would exhibit surface/search behaviour and be able to escape from the modified drum trap. The experiment was done in several pools on the Wingecarribee River, New South Wales from 25- 27 November 2002 to determine if this expectation was realised. Method The trap consisted of a 90 cm diameter x 170 cm long cylinder, covered with black plastic mesh (55 mm x 40 mm), except at the entrance end, where a conical funnel or ‘valve’ made from 3 mm diameter braided polyethylene trawl netting was strung tightly between the circular steel frame at one end of the trap and an oval ring rigidly suspended inside the trap (Figure 4). The trap was fully platypuses, water rats, turtles and diving birds, through a hole in the trap’s roof. A wire-mesh platform was positioned below the escape hole so that small 268 submerged in the pools from which the platypuses were captured. The trap was oriented with the escape hole uppermost. A remote lens for a video camera was Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM Table 3. Results of Experiment 4. Assessment of escape of platypuses from the carp trap in the Wingecarribee River. Sex/Age Weight (g) Length (cm) Female Adult 1080 48.5 Male Adult 1880 S52, Male Juvenile 1790 56.5 Male Adult 1880 57.5 Male Juvenile 1400 53.0 mounted inside the trap to record the behaviour of the animals and these images were stored for later analysis. Platypuses were captured using unweighted gill nets (Grant and Carrick 1974). Each animal was weighed and measured, then temporarily marked with a piece of brightly coloured tape attached to the tail, making the platypuses more visible to observers and to the video camera. Based on observations reported above and Bethge (2002), who reported a maximum foraging dive duration of 138 seconds, individuals were immersed for a maximum of 3 minutes before the trap was lifted to permit them to breathe. If they exited the trap prior to lifting, the elapsed time was recorded. The numbers of times each animal approached the platform below the escape hole was recorded. All animals were used only once in the experiment and remained in the trap for no more than 3 minutes. Results Table 3 shows the dimensions of the platypuses used, the time in the trap, the number of approaches to the platform below the escape hole, and whether or not individuals escaped. Only one juvenile male platypus managed to find the escape hole (after 30 seconds in the trap), but showed reluctance to leave the steel ring around the hole. It re-entered the body of the trap three more times before finally leaving the trap completely. This animal repeatedly relocated the escape hole after re-entering the trap, taking 30, 50 and 50 seconds respectively, before finally escaping. The other four trial animals failed to find the escape hole and were released after 2-3 minutes. Contrary to expectation, platypuses (including the one which escaped) spent most of the time investigating the bottom or ends of the trap, rather than exhibiting surface/search behaviour. In fact, they seemed to actively avoid the platform area below the Proc. Linn. Soc. N.S.W., 125, 2004 Time in Approaches Escape trap (sec) to platform 150 0 No 165 3 No 180 0 No 180 0 No 30 0 Yes escape hole. All animals searched with their bills around the corners of the trap between the sides and ends. The video showed them frequently investigating the acute angled edge between the base of the entrance funnel and the sides of the trap. When released, all animals were observed to surface and appeared to be breathing heavily. EXPERIMENT 5. Assessment of escape of platypuses from the modified carp traps In Experiment 4, platypuses were observed continually searching for an escape hole around the corners of the trap. It was therefore decided to test the effectiveness of escape holes positioned around the base of the entrance funnel. Because the gap between the funnel and the sides of the trap was quite narrow at the base of the funnel, it was considered that most carp would be too large to access openings in this position. Experiment 5 tested the effectiveness of these modifications. The experiment was done in one pool on the Wingecarribee River on 27 November 2002 and then in four pools on the upper Shoalhaven River from 21-23 December 2002. Methods Every third mesh attached to the trap frame at the base of the funnel was released and tied back to provide 90 x 90 x 90 mm triangular openings (Fig. 4, top). In the initial trial in the Wingecarribee River these openings were made only in the upper half of the trap, but in the later trials in the upper Shoalhaven River, openings were made in both the upper and lower halves of the trap. Fourteen platypuses were individually placed in the submerged trap as described in Experiment 4. Again observations were made of the number of times animals approached the platform below the escape 269 REDUCING BY-CATCH OF PLATYPUS Table 4. Results of Experiment 5. Assessment of escape of platypuses from the modified carp trap. * not observed escaping but were not present in trap when it was lifted after 3 minutes; - escape holes only available in upper part of this trap. Sex Weight Length Time in (g) (cm) trap (sec) Female 850 43.0 85 Female 690 41.0 15 Female 900 46.0 22 Female 940 43.5 15 Female 900 44.0 40 Female 790 41.0 4] Female 870 43.5 140 Female 930 44.0 33 Female 860 44.0 <180* Female 840 43.5 45 Female 790 43.0 156 Male 1850 55.2 35 Male 1740 52.0 <180* hole. Escapes through the triangular holes at the base of the entrance funnel were partitioned as being from the ‘upper’ or ‘lower’ openings in the trap. Some underwater video observations were made but the turbidity of the pools made viewing difficult. However, brightly coloured tape attached to the tails of the animals (see Experiment 4) usually permitted their movements in the trap to be observed. Again, if the platypus was not seen to escape, the trap was lifted from the water after a maximum of 3 minutes. Results Thirteen platypuses escaped from the openings around the base of the entrance funnel of the trap within 3 minutes (Table 4). As was observed in Experiment 4, all animals attempted to find an escape route around the bottom or ends of the trap. Another individual used in the initial trial located a hole inadvertently left at the bottom of the trap (which was sealed before subsequent trials). No preference was shown for escape location, with six animals exiting from the ‘upper’ and 5 from the ‘lower’ openings, where both were available (Table 4). One individual moved into the space between the platform and the escape hole but did not find the hole, submerging again and leaving the trap by one of the openings at the base of the entrance funnel. Only two individuals approached the platform at any time during their confinement in the trap. In two instances the platypuses could not be seen, but were no longer in the traps when they were lifted after 3 minutes. It was presumed that they had exited the lower holes, as they were not seen leaving the upper ones, which were visible to the observers. 270 Approaches Escape Escape to platform location 1 Yes - 0 Yes lower 0 Yes lower 0 Yes lower 0 Yes upper 0 Yes upper 0 Yes upper 0 Yes upper 0 Yes lower 0 Yes upper 0 Yes upper 1 Yes - 0 Yes lower DISCUSSION - Carp trap experiments Experiment 4 indicated that the unmodified carp trap would probably result in significant mortality of platypuses if deployed in areas where their distribution overlaps that of carp. However, experiment 5 indicated that carp traps with appropriate escape holes could be used to reduce by-catch of platypuses. Platypuses over a size range of 690-1880 grams were able to exit quite quickly (15-156 seconds) through the 90 mm triangular openings in the modified carp trap. It should be noted that the platypuses used in these experiments were not particularly large. There is considerable sexual dimorphism in the species, with the average male being around 75% heavier and 20% longer than females (Carrick 1995; Grant 1995; Connolly and Obendorf 1998). Individuals of up to twice the size of those used in current experiments are found in some mainland areas (especially west of the Great Dividing Ranges; Carrick 1995; Grant 1995) and in Tasmania males may reach up to three kg (Connolly and Obendorf 1998). Further experiments are required to determine the size of escape holes effective for larger platypuses. In the interim, the authors recommend triangular openings of 100 x 100 mm for east-flowing streams in New South Wales and openings of at least 120 x 120 mm for west-flowing streams in the state. Trials would also need to be carried out to assess the effectiveness of retaining captured carp in the presence larger escape holes. The unexpected lack of surface/search behaviour in platypuses during Experiments 4 and 5 Proc. Linn. Soc. N.S.W., 125, 2004 T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM indicates the importance of field trials of fishing equipment with regard to specific wildlife species. The reason for the unexpected lack of surface/search behaviour in water can only be speculated upon. Platypuses frequently forage among dense woody debris and under submerged overhanging banks (Grant 1995 and personal observation). It may be that a behavioural response of moving down and/or sideways away from an obstruction during foraging may be of greater survival value than attempting to rise directly to the surface when seeking an escape route. No ‘wedging’ behaviour (Evans et al. 1994; Bethge et al. 2001; Bethge 2002; Experiment 3) was exhibited by animals in the carp traps. Rather, all individuals searched constantly for an escape route. GENERAL CONCLUSIONS The results of the literature reviewed and experiments presented in this paper indicate that any fishery in freshwaters of New South Wales based on the use of traps should not be operated as an unregulated fishery (Figure 1) if reducing platypus mortality is a priority. By-catch minimisation has been possible in the eel fishery by a combination of closures of some inland waters and by modifications to provide an airspace in traps used in farm dams and impoundments. Exclusion devices (e.g. grids across the entrance funnels of traps) do not provide a commercially viable option for reducing the by-catch of platypuses in eel or yabby traps. Banning of yabby traps from areas where platypuses occur is currently the only available means of avoiding by-catch mortality in this fishery. The commercial and recreational yabby fisheries in New South Wales are currently restricted to waters where platypuses do not commonly occur or are very uncommonly reported. Trap modifications, which permit the escape of platypuses, appear to be the most feasible means of by-catch minimisation in the use of traps to capture carp. ACKNOWLEDGMENTS This work was conducted under Animal Research Authorities (ACEC 99/13 and ACEC 02/12) from the NSW Fisheries Animal Care and Ethics Committee and under NSW Fisheries Scientific Research Permit F84/1245 (TRG) and NSW National Parks and Wildlife Service Scientific Investigation Licence A184 (TRG). Frank Jordan constructed the grids used in Experiment 2 and Stuart Scott and the Izzard and Laurie families allowed us to work on their properties on the Wingecarribee and Shoalhaven Rivers. Dr Melody Proc. Linn. Soc. N.S.W., 125, 2004 Serena, of the Australian Platypus Conservancy is acknowledged for her personal communication regarding hypothermia in platypuses captured in fyke nets. Funding for much of the reported work was provided by NSW Fisheries. The authors thank Mike Augee and an anonymous referee for their valuable comments and suggestions. REFERENCES Allanson, M. and Thurstan, S. (1999). “Yabby (Cherax destructor) trap by-catch reduction.’ Unpublished report NSW Fisheries-Inland Management. August/September, 1999. Bethge, P. (2002). Energetics and foraging behaviour of the platypus Ornithorhynchus anatinus. PhD thesis. University of Tasmania, Hobart. Bethge, P., Munks, S. and Nicol, S. (2001). Energetics and locomotion in the platypus, Ornithorhynchus anatinus. Journal of Comparative Physiology - B, Biochemical, Systematic and Environmental Physiology 171, 497-506. Beumer, J.P., Burbury, M.E. and Harrington, D.J. (1981). The capture of fauna other than fishes in eel and mesh nets. Australian Wildlife Research 8, 673- 677 Boulton, A.J. and Brock, M.A. (1999). ‘Australian Freshwater Ecology. Processes and Management’. Gleneagles Publishing, Glen Osmond, South Australia. Carrick, F.N. (1995). Family Ornithorhynchidae. Platypus. In “The mammals of Australia’ (Ed R. Strahan) pp. 35-38. (Reed Books: Chatswood). Connolly, J.-H. and Obendorf, D.L. (1998). Distribution, captures and physical dimensions of the platypus (Ornithorhynchus anatinus) in Tasmania. Australian Mammalogy 20, 231-237. Evans, B.K., Jones, D.R., Baldwin, J. and Gabbott, G.R.J. (1994). Diving ability of the platypus. Australian Journal of Zoology 42, 17-27. Grant, T.R. (1991). The biology and management of the platypus (Ornithorhynchus anatinus) in New South Wales. Species Management Report # 5. NSW National Parks and Wildlife Service, Hurstville, New South Wales Grant, T.R. (1993). The past and present freshwater fishery in New South Wales and the distribution and status of the platypus, Ornithorhynchus anatinus. Australian Zoologist 29, 105-113. Grant, T. (1995). ‘The platypus. Unique mammal’. University of New South Wales Press. Sydney. Grant, T.R. and Carrick, F.N. (1974). Capture and marking of the platypus, Ornithorhynchus anatinus, in the wild. Australian Zoologist. 18: 133-135. Grant, T.R. and Denny, M.J.S. (1991). Distribution of the platypus in Australia with guidelines for management. Report to Australian National 271 REDUCING BY-CATCH OF PLATYPUS Parks and Wildlife Service. Mount King Ecological Surveys, Oberon, New South Wales. Jackson, P.D. (1979). Survey of fishes in the west branch of the Tarwin River above Berrys Creek. Victorian Naturalist 97, 11-14 Koed, A. and Dieperink C. (1999). Otter guards in river fyke-net fisheries: effects on catches of eels and salmonids. Fisheries Management and Research. 6: 63-69. Leadbitter, D. (2001). Bycatch Solutions. A Handbook for Fishers in Non-Trawl Fisheries. Oceanwatch and Fisheries Research and Development Corporation. Pyrmont, New South Wales, Australia. FRDC Project NO. 98201. Pettigrew, J.D., Manger, P.R., Fine, S.L.R. (1998). The sensory world of the platypus. Transactions of the Royal Society of London. Series B. 353, 1199-1210. Rankin, T. (2000). Status of the New South Wales commercial yabby fishery. NSW Fisheries Fishery Resource Assessment Series No. 9. Fisheries Research Institute, Cronulla. May 2000. Schiller, C.B., Harris, J.H. (2001). Native and alien fish. In ‘Rivers as Ecological Systems: the Murray- Darling Basin’ (Ed W.J. Young) pp. 229-258. (Murray-Darling Basin Commission: Canberra). Serena, M. (2003). Wanted. Platypus sightings. Australian Society for Limnology Newsletter March 41, 36. to ~~ tO Proc. Linn. Soc. N.S.W., 125, 2004 Platypus Burrow Temperatures at a Subalpine Tasmanian Lake Par Betuce!, SARAH Munks’, HELEN OTLEY’, STEwaART Nicov! ' Division of Anatomy and Physiology, University of Tasmania, GPO Box 252-24, Hobart TAS 7001, Australia (Address for correspondence: Brandstwiete 19, 20457 Hamburg, Germany; philip @bethge.org); * School of Zoology, University of Tasmania, Private Bag 4,GPO, Hobart TAS 7001, Australia Bethge, P., Munks, S., Otley, H. and Nicol, S. (2004). Platypus burrow temperatures at a subalpine Tasmanian lake. Proceedings of the Linnean Society of New South Wales 125, 273-276. When platypuses are in their burrows, microhabitat is of great importance for energy conservation, especially where air temperatures frequently fall below freezing in winter. In this study, we investigated burrow temperatures of platypuses (Ornithorhynchus anatinus) living at a sub-alpine Tasmanian lake. Nine individual platypuses were equipped with time-depth recorders with integrated temperature sensors measuring ambient temperature. Burrow temperatures were recorded in two minute intervals for a total of 61 resting periods (duration: 5.45 to 27.20 hours) and were averaged over the period of resting. Mean burrow temperatures were 17.5 and 14.2°C (SD=2.76 and 0.89, respectively, n=9) in summer and winter, respectively, and ranged between 12.2 and 22.8°C for individual resting periods. In winter, burrow temperatures were held fairly constant over the resting period while in summer larger variations were observed. Burrow temperature in winter was found to be up to 18°C higher than outside air temperature. Manuscript received 26 November 2003, accepted for publication 8 January 2004. Key words: Burrow temperature, Energetics, Ornithorhynchus anatinus, Platypus, Tasmania INTRODUCTION The platypus, Ornithorhynchus anatinus, inhabits the lakes, rivers and streams of eastern Australia from the Cooktown area in the north to Tasmania in the south. Over much of its range, the animal is found in alpine and tableland areas where, especially in winter, air temperatues fall well below freezing and water temperatures approach 0°C (Grant 1995). Grant (1983a) suggested that under such conditions, the microhabitat of platypus burrows is of great importance for energy conservation. Even in an unoccupied artificial burrow the insulation of layers of earth was found to provide significant buffering effect against outside ambient temperature changes both in winter and in summer (Grant 1976). (Grant 1983b) suggested a further modifying effect of the animal’s presence on the microhabitat temperature, elevating it several degrees above that of an unoccupied burrow. In this study ambient temperatures in occupied platypus burrows at a sub-alpine Tasmanian lake were investigated. The use of time-depth recorders with integrated temperature sensors made it possible to determine burrow temperatures during naturally occuring resting periods of the equipped animals. MATERIALS AND METHODS Field experiments were carried out at Lake Lea (41°30° S, 146°50° E), a sub-alpine lake in northwestern Tasmania. Information on burrow temperatures was obtained from nine individual platypuses (4 adult males, mass: 2.27 kg + 0.26 (SD), 5 adult females, mass 1.48 kg + 0.07 (SD)) between November 1998 and January 2000. Platypuses were captured and processed following the methods outlined in Otley et al. (2000) and Bethge et al. (2003). Individuals were equipped with combined data logger- transmitter packages (max 62 mm x 28 mm x 18 mm, weight 50 g, Fig. 1) consisting of a specially designed standard transmitter (Faunatech, Eltham, Victoria) and a time-depth recorder (LTD 10, Lotek Inc., Canada). The packages were attached with glue (5 min-Araldite, Selleys Inc., Australia) to the guard fur of the lower back of the animals, just above the tail, following the method outlined in Serena (1994). Animals were released at the site of capture. After approximately two weeks the animals were relocated by radiotracking and recaptured on emergence and the devices were removed. The data loggers allowed measurement of ambient temperature in the range from 2 to 25°C with PLATYPUS BURROW TEMPERATURES goed) 10 Temperature [°C] “5 Tbur Tw Ta epg 15/6 16/6 17/6 18/6 19/6 20/6 21/6 22/6 23/6 24/6 25/6 26/6 27/6 Figure 1: Winter sample data of water (Tw), air (Ta) and burrow temperatures (Tbur, derived from a time-depth recorder with integrated temperature sensor fitted to the back of a platypus; the temperature is only shown at times when the animal was in the burrow). an accuracy of 0.06°C. The devices were calibrated by the manufacturer (Equipment for temperature- calibration: Neslab RTE-2000 Bath/Circulator and Omega HH40 Thermistor/Thermometer). Temperature sensors were located at the back end of the devices and were facing backwards when the devices were fixed on the platypus’s lower backs. Ambient temperature was measured in two-minute intervals for 11 days each. While foraging, the sensors measured water temperatures. In resting platypuses, ambient temperatures close to the animals’ bodies (approx. 5 mm from above the fur) were recorded. The resting period was defined as the time span between the end of the last dive of a foraging trip (detected by the depth sensor of the time-depth recorders) and the beginning of the first dive of the following foraging trip. Burrow temperatures, i.e. ambient temperatures during resting periods, were recorded in two minute intervals for a total of 61 resting periods and were averaged over the period of resting. Resting periods ranged from 5.45 to 27.20 hours. All investigated platypuses occupied burrows in consolidated steep or gently sloping earth banks of the lake or along associated creeks. Water and air temperatures at Lake Lea were recorded in two-hour 274 intervals using archival tags (HOBO Thermocouple logger and Stowaway Temperature Logger, Onset Computer Corp., USA). Water temperature was measured in the lake while air temperature was taken in a wind shaded forest patch nearby. RESULTS Mean burrow temperatures were 17.5 and 14.2°C (SD=2.76 and 0.89, respectively, n=9) in summer and winter, respectively, and ranged between 12.2 and 22.8°C for individual resting periods. In winter, burrow temperature was held fairly constant over the resting period while in summer larger variations were observed. A low but significant correlation between air temperature and burrow temperature was found with higher air temperatures resulting in higher burrow temperatures (p=0.003, n=61). Ambient air temperatures ranged between -4°C and 31°C and water temperatures between 0°C and 29°C depending on season. Samples of measured burrow temperatures and corresponding air and water temperatures are shown in Fig. | and Fig. 2 for winter and summer, respectively. Proc. Linn. Soc. N.S.W., 125, 2004 P. BETHGE, S. MUNKS, H. OTLEY AND S. NICOL Temperatuie [°C] 7H 68 ON 10/4 11/1 12 13/1 14/1 151 161 17/1 184 19/1 20/1 Date Figure 2: Summer sample data of water (Tw), air (Ta) and burrow temperatures (Tbur, derived from a time-depth recorder with integrated temperature sensor fitted to the back of a platypus; the temperature is only shown at times when the animal was in the burrow). DISCUSSION Grant (1983b) suggested that platypus burrows act aS microenvironments, buffering the animals against the rigours of below-freezing air temperatures in winter, and modifying the effects of high summer temperatures. Accordingly, we found that in winter, burrow temperatures at Lake Lea were up to 18°C higher than outside air temperatures (Fig. 1). In summer, the burrows at Lake Lea clearly buffered high midday temperatures of over 25°C (Fig. 2). These findings are in line with results by Grant (1976) and Munks (personal communication). In winter, unoccupied artificial burrow temperatures in the upper Shoalhaven River, NSW, averaged around 14°C (this study: 14.2°C) despite the fact that ambient air temperatures dropped as low as -5°C. During summer the temperature of an unoccupied artificial burrow averaged around 18°C (this study: 17.5°C) with air and water temperatures being several degrees higher (Grant 1976, Grant 1995). Munks (personal Proc. Linn. Soc. N.S.W., 125, 2004 communication), while monitoring the burrow of a breeding platypus in lowland Tasmania, recorded a mean burrow temperature of 16.5°C (range 12.5 to 20°C) during late summer/early autumn. The consistency of these data from different sites suggests that platypus burrow temperatures are fairly constant regardless of habitat. Whether this is a consequence of the metabolic heat produced by the animals or mainly of physical characteristics of their burrows, remains unclear. Results of Grant (1976) from unoccupied burrows are in line with findings presented here from occupied burrows. This suggests that - at least in burrows located in consolidated earth banks - physical characteristics of the burrow are more important for burrow temperature than the absence or presence of the animal. This view is supported by the significant correlation between air temperature and burrow temperature found in this study. However, Munks (personal communication) reported peak burrow temperatures when the mother returned to the nest to suckle her young. Also, Grant (1983b) suggested that the animal’s presence further PLATYPUS BURROW TEMPERATURES elevates the microhabitat temperature of the burrow. In captivity, Grant (1976) observed, that the temperature in an uninsulated plywood nest box rose around | to 2°C above ambient temperature when an animal was inside. We suggest that these different observation are a consequence of different burrow characteristics. In this study, all investigated platypuses occupied burrows in consolidated earth banks. Under such conditions, the insulation properties of the surrounding earth and of the nesting material inside the burrow are most likely the main factors determining burrow temperature. A fairly constant burrow temperature may of course be more critical during the breeding period (Grant, personal communication), which makes deep earth burrows ideal for nesting. A different situation, however, might occur in burrows, which are closer to the surface or above ground. Otley et al. (2000) reported that 25 % of burrows at Lake Lea were located within dense vegetation, such as sphagnum and button grass. The insulation properties of such burrows would be expected to be poor compared to underground earth burrows. How animals cope with high thermal stress in vegetation burrows and if they use this sort of burrow site regardless of season or even during nesting requires further investigation. ACKNOWLEDGMENTS This work was supported by the Australian Research Council, an Overseas Postgraduate Research scholarship by the University of Tasmania and a doctoral scholarship by the DAAD (Deutscher Akademischer Austauschdienst, Germany ,Hochschulsonderprogramm III von Bund und Landern‘). The field work was carried out under permit from the Department of Parks, Wildlife and Heritage, Tasmania, the Inland Fisheries Commission, Tasmania and the University of Tasmania Ethics Committee. Thanks to all those who assisted with the field work and to Mr H.Burrows for access to private land. 276 REFERENCES Bethge, P., Munks, S., Otley, H.and Nicol, S. (2003) Diving behaviour, dive cycles and aerobic dive limit in-the platypus Ornithorhynchus anatinus. Comparative Biochemistry and Physiology A 136/4, 799-809. Grant, T.R. (1976). Thermoregulation in the Platypus, Omnithorhynchus anatinus. PhD thesis, University of New South Wales, Australia. Grant, T.R. (1983a). Body temperatures of free-ranging platypuses, Ornithorhynchus anatinus (Monotremata), with observations on their use of burrows. Australian Journal of Zoology 31, 117-122. Grant, T.R. (1983b). The behavioural Ecology of Monotremes. In ,Advances in the Study of Mammalian Behaviour’ (Eds J.F. Eisenberg and D.G. Klieman). The American Society of Mammalogists, Special Publication Vol. 7, pp 360-394. Grant, T.R. (1995). The platypus. A unique mammal. University of New South Wales Press, Sydney. Otley, H.M., Munks, S.A. and Hindell, M.A. (2000). Activity pattern, movements and burrows of platypuses (Ornithorhynchus anatinus) in a sub- alpine Tasmanian lake. Australian Journal of Zoology 48, 701-713. Serena, M. (1994). Use of time and space by platypus (Ornithorhynchus anatinus: Monotremata) along a Victorian stream. Journal of Zoology 232, 117-130. Proc. Linn. Soc. N.S.W., 125, 2004 Ultrasonography of the Reproductive Tract of the Short-beaked Echidna (Tachyglossus aculeatus) D. P. Hiccins Faculty of Veterinary Science, BO1, University of Sydney NSW 2006 (damienh @ vetp.usyd.edu.au) Higgins, D,P. (2004). Ultrasonography of the reproductive tract of the short-beaked echidna (Tachyglossus aculeatus). Proceedings of the Linnean Society of New South Wales 125, 277-278. We describe a brief investigation of ultrasonography as a tool to monitor reproductive activity and to determine the sex of short- beaked echidnas (Tachyglossus aculeatus). We found trans-abdominal ultrasound to be of limited use for monitoring ovum development but it appears to be useful for imaging the uterus. We also found ultrasonography to be a useful tool to confirm the sex of echidnas by visualizing the testis. Manuscript received 18 August 2003, accepted for publication 8 January 2004. KEYWORDS: abdomen, echidna, monotreme, reproduction, Tachyglossus, testis, ultrasound, uterus. Here we describe a brief investigation of ultrasonography as a tool to monitor reproductive activity and to determine the sex of short- beaked echidnas (Tachyglossus aculeatus). Griffiths (1968) described the gross anatomy of the reproductive tract of the female echidna. An ovum of 3-4 mm diameter is ovulated from one of the two flat, sauropsid-like ovaries, which lie ventrocaudal to the kidneys (Griffiths 1968). Although only one ovum is ovulated in the echidna, Flynn (1930) reported that up to three large ova and several much smaller ova may occur on the ovary. Hughes and Carrick (1978) concluded from Hill and Gatenby (1926), Caldwell (1887) and Flynn and Hill (1939) that the ovum has a vitelline membrane, a zona pellucida and a proalbumen which may be analogous to the liquor folliculi of the graafian follicle, but has no follicular antrum. During its passage down the fallopian tube, the ovum swells to 5 mm diameter. The shell membrane is first laid down in the fallopian tube and later thickens in the uterus. The egg absorbs fluid in-utero and expands from 6.5mm diameter to 15 mm x 13 mm. Ten short- beaked echidna carcasses were placed in dorsal recumbency. A portable ultrasound machine with a 7.5 Mhz linear transducer (SSD- 500, Aloka, Japan) was used to image the abdomen. Results were confirmed by dissection. An additional nine echidnas were then anaesthetized and examined in a similar fashion. Positioning the transducer on the ventral abdominal wall, lateral to the epipubic bones avoided the need to shave the hair of the pseudopouch and minimised interference by intestinal gas. Dissection confirmed that the gonads lie against the dorsal body wall, dorsal to the cranial ends of the epipubic bones. Ovaries of freshly dead echidnas lacked grossly visible developing ova. Of frozen and thawed bodies, which generally had poorer tissue contrast, ovaries and ova were not visible by ultrasonography. A structure in the expected location of the ovaries and comprising several 2- 3 mm diameter, thin walled, echolucent bodies was sometimes visible in living echidnas during the breeding season, however, the scarcity of surrounding interstitium made repeatable identification of individual putative ova very difficult. In addition, the small intestine frequently cast gas shadows over the gonads, reducing their visibility. It is likely that trans- rectal ultrasound would improve visualization of the ovaries but may be of limited use in serial observations, where the extent of manual or chemical restraint required may introduce variations to the reproductive cycle (Clarke and Doughton 1983, River and Rivest 1991). The entire oviducts of reproductively active live and dead animals were clearly visible, especially when adjacent to a full bladder. Ova were not seen in the oviducts of any of our animals. Testes appeared as 15 to 25mm long, ovoid, homogenous, soft tissue structures and, when present, were always visible caudal to the kidney and, on the left side, dorso-caudal to the mobile, spherical portion of the spleen. Due to their similar appearance on ultrasound, both the spleen and testis were sighted on the left before the left testis was identified. ULTRASONOGRAPHY IN ECHIDNA REPRODUCTIVE STUDIES In conclusion, we found trans-abdominal ultrasound to be of limited use for monitoring ovum development but it appears to be useful for imaging the uterus. In sexing echidnas, the inability to extrude or palpate a phallus does not confirm its absence, and other characteristics such as absence of a pseudo-pouch or presence of spurs may not be reliable indicators of sex, therefore the gender of echidnas in captive collections is sometimes mistaken. We found ultrasonography to be a useful tool to confirm the sex of echidnas in these circumstances. ACKNOWLEDGMENTS We thank Dr Marianne Offner for her assistance in interpretation of ultrasonographs, Medtel for the provision of the Aloka SSD-500 ultrasonography machine, and the Zoological Parks Board of NSW, Novartis Australia and the Winifred Scott Foundation for their financial support. REFERENCES Caldwell, W. H. (1887) The embryology of Monotremata and Marsupialia- Partl. Philosophical Transcripts of the Royal Society. 178(B), 463- 480. Clarke, I. J. and Doughton, B. W. (1983) Effect of various anaesthetics on the resting plasma concentrations of lutienising hormone, follicle stimulating hormone and prolactin in ovariectomised ewes. Journal of Endocrinology. 98, 79-89. Flynn, T. T. (1930). On the unsegmented ovum of the echidna (Tachyglossus) Quarterly Journal of Microscopical Science. 74, 119-131. Flynn, T. T. and Hill, J. P. (1939) The development of the Monotremata PartVI -Growth of the ovarian ovum, maturation, fertilisation and early cleavage. Philosophical Transcripts of the Royal Society (London) XXYV (6), 571-578. Griffiths, M. E. (1968) ‘The Echidna.’ (Pergamon Press: UK). Hill, J. P. and Gatenby, J. B. (1926). The corpus luteum of the Monotremata. Philosophical Transcripts of the Royal Society (London) Wl, 715-762. Hughes, R. L. and Carrick, F. N. (1978). Reproduction in female monotremes. Australian Zoologist 20, 233-254. River, C. and Rivest, S. (1991). Review Article. Effect of stress on the activity of the hypothalamic- pituitary- gonadal axis: peripheral and central mechanisms. Biology of Reproduction 45, 523- 532) 278 Proc. Linn. Soc. N.S.W., 125, 2004 Excretion Profiles of Some Reproductive Steroids in the Faeces of Captive Female Short-beaked Echidna (Tachyglossus aculeatus) And Long-beaked Echidna (Zaglossus sp.) D.P. Hiccins, G. Tostas, G.M. Stone Faculty of Veterinary Science, BO1, University of Sydney NSW 2006 (damienh@ vetp.usyd.edu.au) Higgins, D.P., Tobias, G. and Stone, G.M. (2004). Excretion profiles of some reproductive steroids in the faeces of captive female short-beaked echidna (Tachyglossus aculeatus) and long-beaked echidna (Zaglossus sp.). Proceedings of the Linnean Society of NSW 125, 279-286. We evaluated and applied an existing faecal reproductive steroid extraction and radio-immunoassay (RIA) procedure to samples from captive short-beaked (Tachyglossus aculeatus) and long- beaked (Zaglossus sp.) echidnas. Steroids were extracted from faeces with diethyl ether, resuspended in 80% methanol and lipids removed with petroleum ether. The methanol fraction was purified and assayed for progestins or oestrogens, results corrected for procedural losses and converted to ng/ g dry weight of faeces. One T. aculeatus was injected with radiolabelled and natural progesterone and faecal extracts were subjected to high- performance liquid chromatography (HPLC) to allow partial identification of radiolabelled and RIA- reactive metabolites. The major RIA-reactive substance and the major labelled ["*C] compound co-eluted with progesterone. An additional RIA-weak compound co-eluted with 208-dihydroxyprogesterone, and three additional RIA-weak, radio-labelled compounds eluted but were not identified. Increases in faecal progestin of echidnas occurred at 17 + 3 (n=5), 33 + 3 (n=4) and 48 (n = 1) day intervals, supporting a cycle length of approximately 17 or 33 days. However, further study micorporating more animals, behavioral observations and more frequent sampling of faecal oestrogens is required to produce more definitive results. Manuscript received, 18 August 2003, accepted for publication 8 January 2004. KEYWORDS: Faecal reproductive steroids, HPLC, monotreme, oestrogen, progestin, radioimmunoassay, Tachyglossus aculeatus, Zaglossus. INTRODUCTION The short- beaked echidna (Tachyglossus aculeatus) is widespread within Australia and New Guinea. The long- beaked echidna (Zaglossus bruijnii) is restricted to the highlands of New Guinea where it is endangered by human interference (Flannery 1990). Despite more than 100 years of captive husbandry, it is rare for these animals to breed in captivity (Augee et al. 1978; Boisvert and Grisham 1988) and knowledge of the timing and hormonal control of the reproductive cycles of monotremes is limited. The presence of a luteal phase is generally accepted, based on histological evidence (Hill and Gatenby 1926; Griffiths 1968; Hughes and Carrick 1978; Griffiths 1984) but its role and duration is unknown. In addition, observations of gestation range from greater than 10 days (Carrick 1977) to 28 days (Broom 1895) after mating. Griffiths (1984) speculated that, like some reptiles and bats (Racey and Potts 1970), female echidnas may store sperm, or that torpor may alter gestational length, as in pygmy possums (Cercartetus spp), brown antichinus (Antichinus stuartii), eastern quolls (Dasyuris viverrinus) (Tyndale- Biscoe 1973) and bent-winged bats (Miniopterus spp) (Wimsatt 1969). Longitudinal studies better define reproductive cycles and illustrate inter-individual variation than cross-sectional studies, which are more conveniently applied to wild animals (Lasley 1985). However, even captive echidnas are cryptic and curl into a tight ball when threatened, making difficult the frequent collection of blood, urine or urogenital swabs without anaesthesia or forceful restraint, which may cause variation of reproductive cycles and behaviour (Clarke and Doughton 1983; Rivier and Rivest 1991; Cleva et al 1994). Non- invasive faecal reproductive steroid assays have been used to describe the reproductive cycles of many species. This paper reports the initial assessment and application of a faecal reproductive steroid assay as a non- invasive technique for the first sequential study of female echidna FAECAL REPRODUCTIVE STEROIDS IN CAPTIVE ECHIDNAS reproductive endocrinology. MATERIALS AND METHODS Animals and housing Study animals were six female T. aculeatus, aged between 4 and 7 years and of 3 to 5 kg bodyweight, and three female Zaglossus bruijnii (probably Z. bartoni of Flannery and Groves 1998), aged between 20 and 32 years and of 6.5 to 14 kg bodyweight, all from the Taronga Zoo collection. The study was conducted in two phases: From May to September 1995, two T. aculeatus and all Zaglossus sp. were housed in two indoor enclosures with reverse cycle seasonal lighting and in the continual presence of a male of their species. From June to October 1997, four female 7. aculeatus were housed outdoors, in adjacent 5.1 x 6.3 m enclosures. These females were housed individually to accommodate the solitary nature of the animal (Augee et al. 1975; Abensperg-Traun 1991) and to facilitate identification of the source of faeces. A male T. aculeatus had access to all four enclosures through magnetically controlled doors until they failed, after which he was manually rotated, daily, between enclosures. Sixty-centimeter deep woodchip substrate, half pipes, tables, tree branches and logs were provided as shelter. Feeding and sample collection Animals were fed daily slurry of minced beef, egg, cereal, vitamin and mineral supplements and sufficient unprocessed bran to produce firm stools. Initially, 1mm x 1mm x 3mm food grade polyethylene pellets (Hoechst Industries, Australia) were added to the food of the indoor groups to identify the source of faeces. It appeared possible that not all pellets were being excreted, therefore the feeding of pellets was discontinued and for the course of the study each animal from the indoor groups was placed in a separate room for 24- 48 h once weekly and faeces were collected. Blue food dye (8 mg/day, Hexacol Brilliant Blue FCF Supra 75328, Pointing Hodgsons Pty Ltd, Australia), was added to the food of the outdoor female animals to make faeces more visible, and all visible faeces were collected daily. Samples were handled using latex gloves and stored in plastic zip- lock bags at -20°C for up to one year. Extraction and purification Due to the need to separate echidnas for sample collection in the first phase of the study, the sampling interval for progestin excretion profiles of echidnas housed indoors was 5 to 7 days. Sampling 280 interval for progestin and oestrogen excretion profiles of echidnas housed individually outdoors in the second phase of the study was 1 to 3 days. The steroid extraction technique was based on a procedure used by Hindle and Hodges (1990). Each stool was finely chopped and mixed, then duplicate 0.5g samples were transferred to new glass vials (Econo Glas Vial, Packard, USA). Pieces were broken up using a glass rod, 5 ml diethyl ether (APS, Ajax Finechem, Australia) was added and vials were rotated for 30 min then centrifuged at 1500 G for 15 min at 4°C. The faecal sediment and aqueous portion were frozen in liquid nitrogen. Supernatant was decanted, evaporated at 30°C under nitrogen gas and reconstituted in 5 ml 80%(v/v) methanol (80% MeOH) by rotation for 30 min. Solutes were partitioned by addition of 5ml of petroleum ether (B.P. 40°C to 60°C, APS, Ajax Finechem, Australia), rotation for 20 min, and centrifugation at 1500 G for 15 min. The 80% MeOH fraction was aspirated and then stored at -20°C. Following extraction, faecal residue was dried at 100°C for 4 hours and weighed to determine dry matter content. Aliquots of 500 ul faecal extract in 80% MeOH were dried at 80°C under vacuum, reconstituted in Iml 10% MeOH by agitation at 30°C for 30 min, and purified using Sep-Pak C18 Cartridges (Waters Scientific, Milford, USA) according to manufacturers recommendations. Eluants of 25%, 50%, 75%, 90% and 100% MeOH were collected and stored at -20°C. Radioimmunoassay (RIA) Duplicate 200 wl aliquots of eluates (unknowns) were dried and reconstituted in 2001 10% MeOH in 1P buffer (0.031M Na,HPO,, 0.019M NaH,PO,.2H,O, 0.154M NaCl and 0.1%w/v gelatin, pH 7.4) by agitation at 30°C for 60 min. Radiolabelled steroids ([1,2,6,7--H] progesterone in toluene ((7H]P, 96 Ci/mmol; Amersham Australia, Sydney, NSW) or [2,4,6,7-7H] oestradiol in toluene ([HJE, 104 Ci/mmol; Amersham Australia, Sydney NSW)) were dried and reconstituted in 1P buffer to approximately 15000 dpm/ 100 ul. Our ovine antiserum to progesterone-11a- hemisuccinate-BSA conjugate (1:55000 final dilution, #C-9817 Sirosera™, CSIRO Bioquest, Blacktown, Australia), cross-reacted with progesterone (100%), 11B- hydroxyprogesterone (32.5%), corticosterone (18.8%), 20a- hydroxy-4-pregnane-3-one (0.7%), 17a- hydroxyprogesterone (0.2%), 20B- hydroxy-4- pregnane-3-one (0.2%), pregnenolone (0.2%), oestradiol (<0.2%), testosterone (<0.2%), cortisol (<0.2%) (Curlewis, Axelson and Stone, 1985). Our Ovine antiserum to 17f- oestradiol-6- carboxymethyloxime-BSA (1:100000, #9757 Proc. Linn. Soc. N.S.W., 125, 2004 D.P. HIGGINS, G. TOBIAS AND G.M. STONE Sirosera™, CSIRO Bioquest, Blacktown, Australia), cross-reacted with oestradiol (100%), oestrone (10.8%), oestriol (2.3%), oestradiol- 17a (<0.1%), progesterone (<0.1%), testosterone (<0.1%), androstenedione (<0.1%), cortisol (<0.1%), corticosterone (<0.1%) (Curlewis 1983). Standards were generated from two overlapping doubling dilutions of progesterone (BDH Chemicals, Australia) from 500 - 7.84 pg/100 wl or oestradiol (BDH Chemicals, Australia) from 500 - 1.96 pg/100 ul. Reactions contained 100 ul radiolabelled steroid, 100 ul antiserum and either 200 ul of unknown in 10% MeOH in 1P buffer or 100 ul 20% MeOH in 1P buffer and 100 ul of standard. The resulting 400 pl was vortexed for 30 sec and incubated at 4°C for 18 h. Triplicate “total” (200 ul 1P buffer, 100 wl radiolabelled hormone in 1P buffer, 100 ul 20% MeOH in 1P buffer), “non-specific binding” (200 ul 1P buffer, 100 pl radiolabelled hormone in 1P buffer, 100 ul 20% MeOH in 1P buffer) and “B ” (100 ul 1P buffer, 100 wl antiserum in 1P buffer, 100 pl radiolabelled hormone in 1P buffer, 100 ul 20% MeOH in 1P buffer) standards were processed simultaneously with unknowns and standards. Free radiolabelled hormone was removed from all except “total” solutions by incubation for 10 min at 4°C with 500 ul of charcoal/ dextran solution (0.25% w/v Norit-A filtered activated charcoal powder, Matheson, Coleman and Bell, USA) and 0.025% w/v dextran I70 (Pharmacia Fine Chemicals, Sweden) suspended in 1P buffer). In place of the charcoal/ dextran solution, 500 ul of milli Q water was added to “total” solutions. After centrifugation at 1500 G for 10 min at 4°C, the supernatant was decanted and its radioactivity measured as counts per minute (cpm) on a Beckmann LS 6500 Liquid Scintillation Spectrometer (Beckmann Instuments Inc, CA, USA.), which then converted cpm to disintegrations per minute (dpm) using an external standard. High- performance liquid chromatography (HPLC) of excreted metabolites One female T. aculeatus was injected intra- peritoneally with 5 mCi of [4-'4C] progesterone ({'“C]P, 48.9 mCi/mmol, NEN Dupont, USA) and 2 mg of natural progesterone in 30% (v/v) propylene glycol in isotonic saline. Eight 0.5 g faecal samples were obtained two days after injection. Extracts from these samples were pooled into two samples and subjected to sep-pak chromatography. Eluates of 2252 dpm and 2440 dpm were dried under N, gas, reconstituted in 75% acetonitrile, filtered and subjected to HPLC (K65B HPLC system, ETP Kortec, Australia) at a flow rate of 0.5 ml per minute, using 61% acetonitrile at a Proc. Linn. Soc. N.S.W., 125, 2004. pressure of 2250 psi at room temperature. Fractions — were collected every 30 sec for 22 min, then every minute for 19 min, then every 10 min for 20 min. Absorbance at 240nm was measured, to monitor the separation of steroids with a 4-ene-3-ketone structure. Elution time of progesterone was identified using [H]P and a progesterone standard and the column was calibrated for testosterone, androstenedione, progesterone and 20a dihydroprogesterone. Assessment of extraction, purification and RIA procedures Three different solvents were tested for use in the extraction process. Faeces containing metabolites of injected radiolabelled and natural progesterone were agitated in 90% MeOH, 80% MeOH or diethyl ether, and partitioned with petroleum ether as described above. The three solvents and their petroleum ether portions were assayed for progestins as above. The Sep-pak chromatography elution profile for oestrogen calculated by Spanner et al. (1997) was assumed for this study. The elution profile for progestin was determined by Sep-pak chromatography of solutions containing 200 fmol [?H]P, using the series of MeOH dilutions described previously or the same series of dilutions of ethanol (EtOH). Co-elution of metabolites of faecal origin with progesterone was determined by adding 25000 dpm [?H]P to duplicate faecal extracts from five female T. aculeatus and subjecting these to Sep-pak chromatography. Sample steroid recovery was estimated by adding 25000 dpm [?H]P or 30000 dpm [?H]E to respective samples and then performing the extraction. Duplicate 50 ul aliquots of purified 80% MeOH extract were combined with 500 wl Milli-Q water and 5 ml of scintillation fluid. Triplicate “total” vials were prepared, each containing 100 ul of radiolabelled hormone solution, 50 ul 80% MeOH, 400 ul Milli-Q water and 5 ml of scintillation fluid. Triplicate “blank” vials were prepared, each containing 50 ul 80% MeOH, 500 ul Milli-Q water and 5 ml of scintillation fluid. Radioactivity was measured and percentage recovery was calculated by the formula: R = [400(d-B)/ (T-B)] x 100 where R = percentage recovery (%), d = sample dpm, B = mean “blank” dpm, T = mean “total” dpm. To assess parallelism, faecal extracts from three faecal samples were reconstituted and diluted twofold and fourfold in 10% MeOH in 1P buffer. Standards were similarly diluted and all dilutions were assayed for progestins as described above. Spanner et al. (1997) estimated parallelism of the oestradiol assay. 281 FAECAL REPRODUCTIVE STEROIDS IN CAPTIVE ECHIDNAS The inter- assay coefficients of variation for progesterone and oestradiol assays were taken as the mean of the coefficients of variation of repeated (n=2), duplicated extraction and assay of 6 and 4 randomly chosen samples, respectively. The intra- assay coefficient of variation was estimated from the mean of two coefficients of variation, each calculated from five concurrent replicate extractions and assays of two randomly chosen samples. The intra-assay coefficient of variation was estimated for two progesterone extraction methods to determine the homogeneity of steroid in the stool. In the first (unmixed) method, 5 samples were taken from an intact stool. In the second (mixed) method, the stool was finely chopped and mixed and each of the 5 samples consisted of at least 5 randomly chosen pieces from the mix. Spanner et al. (1997) estimated the intra-assay coefficient of variation of the oestradiol assay. Data analysis Standard curve generation and conversion of dpm to pg hormone’ scintillation vial were calculated with “Assayzap” (Biosoft, Cambridge). All other calculations and graphs were made using “Excel 5.0” (Microsoft, USA). Mean steroid recovery was calculated from the first 60 samples in each assay. Mean recovery was used to correct results of progestin assays for procedural losses. As steroid recovery was more variable in oestrogen assays, results were corrected using a recovery value calculated for each individual sample. Faecal steroid peaks were defined as those values greater than 1.5 standard deviations from the mean of all values from that animal (Graham et al 1995). RESULTS High pressure liquid chromatography The eluate with the highest RIA activity and moderate radioactivity was collected at 26 min, approximating the progesterone standard, which eluted at 25.5 min. [*H]P eluted at 24 min. A moderately radioactive eluate with poor RIA activity that was collected at 22 min coincided with a 20B- dihydroxyprogesterone standard, which has a low cross reactivity with the antiserum. Other [C]-labelled, moderately RIA-reactive compounds that eluted at 30, 36 and 38 min and one [“C]-labelled, weakly RIA- . reactive compound that eluted at 22 min were not identified. One RIA-reactive compound that eluted at 41 min did not co-elute with a ['*C]-labelled metabolite and this substance is yet to be identified. Assessment of progestin extraction, purification and RIA procedures As an initial solvent, ether extracted 37.0 + 4.3% (mean + S.E.) more [“C] labelled progestin than either 80% MeOH or 90% MeOH and was used in all subsequent extractions. Less than 10% of extracted steroid appeared in the petroleum ether fraction. Mean percentage recovery of [7H]P through extraction and Sep-pak chromatography was 52% + 7.17 (mean + SD). Mean percentage recovery of HJE was 30% + 13.7 (mean + SD). Almost all (7H]P was recovered during Sep- pak chromatography. MeOH was chosen as the chromatography solvent as EtOH eluted the [H]P across a greater range of EtOH concentrations. Of recovered [*H]P metabolites, 79.1% eluted in the 90% MeOH fraction and 93% eluted in the 75% MeOH and 90% MeOH fractions combined, with a mean 75% MeOH: 90% MeOH ratio (75:90 ratio) of 1:3.5. Of the progestin RIA- reactive faecal steroids recovered from the column, 85% was measured in the combined 75% MeOH and 90% MeOH fractions with a mean 75:90 ratio of 1:2.5. Correlation coefficients of parallelism curves for the progestin assay ranged from 0.988 to 1.000. Dose response curves for standards and extracts did not differ significantly (P>0.05) in slope. Sensitivity of the assay, as defined by 10% displacement from the Bo binding was 10 pg/assay tube. The intra- assay coefficients of variation were estimated to be 6.2% (mixed) and 25.7% (unmixed), therefore the mixed method was employed in all further extractions. The inter- assay coefficient of variation was estimated to be 14.9% for progesterone assays and 6.8% for oestradiol assays. Faecal progestins Maximum and minimum faecal progestin concentrations from each of the six T. aculeatus ranged from 480 to 1800 ng/g dry weight faeces (mean 860 ng/g) and 5 to 100 ng/g dry weight faeces (mean 71 ng/g), respectively. Intervals between samples that contained progestin peaks clustered at 17 + 3 (n =5), 33 + 3 (n = 4) and 48 (n = 1) days. Maximum and minimum faecal progestin concentrations from each of the three Zaglossus sp. ranged from 260 to 500 ng/ g dry weight faeces (mean 420 ng/g) and 10 to 70 ng/ g dry weight faeces (mean 40 ng/g), respectively. Two Zaglossus sp. produced two peaks each and the intervals between samples that contained these were 28 and 70 days. The third produced one peak. Faecal Oestrogens Maximum and minimum faecal oestrogen concentrations from the four animals ranged from 21 Proc. Linn. Soc. N.S.W., 125, 2004 D.P. HIGGINS, G. TOBIAS AND G.M. STONE to 45 ng/g dry weight faeces (mean 33 ng/g) and 3 to 14 ng/g dry weight faeces (mean 7 ng/g), respectively. Intervals between adjacent oestrogen peaks were 8, 19, 24 and 30 days apart. Fluctuations approaching 1.5 SD above the mean were common and made difficult the detection of any possible cyclic activity as intervals between these ranged from 4 to 16 days. Combined profiles Of the 8 oestrogen peaks, 7 were associated with progestin increases to concentrations less than 1.5 SD above the mean. Combined oestrogen and progestin profiles from two T. aculeatus are shown in Figures 1 and 2. DISCUSSION Feeding and sample collection Plastic pellets were less suitable as a faecal marker than the blue food dye. T. aculeatus consumed 95% of pellets placed in their food while Zaglossus sp. consumed less than 50%. Not all ingested pellets ~ were recovered, indicating that pellets were being retained or faeces were remaining undetected. Food containing the blue dye was readily eaten and faeces containing the dye were considerably more detectable than those without. Passage time of the plastic pellets ranged from 12 hours to greater than 48 hours. High pressure liquid chromatography The strong antiserum cross-reactivity of the substance which eluted at 26 min, and its proximity to the elution time of the progesterone standard, makes progesterone its likely identity, however further confirmation of this would be desirable as we are unable to explain the elution of (7H]P 2 min earlier. The cross-reactive metabolites that were less polar than progesterone were not identified. These substances may contribute to the difference in 75:90 ratio between the Sep-pak elution profiles of [7H]P and progestins of faecal origin as the 75%MeOH, or less polar, component was less in the ?H]P profile. As a priority, future studies should identify the RIA-reactive progestins (ng/g dwt) 56 64 65 68 69 70 72 75 76 77 79 81 83 86 89 92 94 96 _ day oestrogens (ng/g dwt) me Progestin —e—oesirogen Figure 1. Combined faecal oestrogen and progestin profiles of one T. aculeatus over a 40-day period showing alternating progestin and oestrogen peaks greater than 1.5 SD above the mean, suggesting an oestrous cycle of 29 days. Also visible are increases less than 1.5 SD above the mean, suggesting concurrent progestin and oestrogen rises at 65, 79 and 94 days with interceding raised progestin/ lowered oestrogen periods surrounding 70 and 89 days, suggesting two cycles of 15 days. = mean faecal progestin/ oestrogen concentration; ............00. = mean faecal progestin/ oestrogen concentration + 1.5 SD. Proc. Linn. Soc. N.S.W., 125, 2004 283 FAECAL REPRODUCTIVE STEROIDS IN CAPTIVE ECHIDNAS ge Progestins —¢— oestrogens progestins (ng/g dwt) 27 33 36 42 45 52 58 61 62 64 66 69 71 72 74 76 78 80 82 84 86 88 90 92 94 days oestrogens (ng/g dwt) Figure 2. Combined faecal oestrogen and progestin profiles of one T. aculeatus over a 67-day period showing alternating progestin and oestrogen peaks greater than 1.5 SD above their respective mean, suggesting an oestrous cycle of approximately 32 days. Also visible are additional fluctuations of oestrogen concentration at 52, 78, 82 and 86 days and progestin at 36 days, which hinder clear interpretation of oestrous cycles. progestin/ oestrogen concentration + 1.5 SD. compound that did not correspond to a radiolabelled metabolite. A similar examination of oestrogen metabolites would assist interpretation of oestradiol assays. Extraction and recovery As homogeneity of steroid in the faeces was poor, mixing of the stool before sampling was necessary to reduce intra-assay variance. Though diethyl ether extracted the most steroid, recovery through extraction was low and variable in this study, especially in oestradiol assays. We attempted to correct for this by correcting for procedural losses using individual recovery values for each sample in oestrogen assays but progestin assay data were corrected using a mean recovery value. Use of individual recovery values for progestin samples may have improved interpretation of data. MeOH concentrations exceeding 5% in the RIA incubations considerably reduced steroid- antibody binding. At each corresponding 284 = mean faecal progestin/ oestrogen concentration; .................00+ = mean faecal concentration, EtOH had a greater effect on steroid- antibody binding than MeOH. Reconstitution of eluates in 10% MeOH to produce a final concentration of 5% MeOH in the assay provided adequate steroid solubility and minimised interference with steroid-antibody binding. Progestin and oestradiol profiles Mean, maximum and minimum progesterone and oestrogen values varied among animals, indicating that this technique may be unsuitable for assessing the status of an animal from a single measurement. The small number of animals available for the study and the need for further work to identify antiserum-reactive metabolites limits the conclusions that can be drawn from the sequential data obtained in this study. However, the lack of knowledge in this area makes some trends worthy of comment for consideration in future work. The intervals between subsequent progestin peaks in this study suggest a progesterone periodicity Proc. Linn. Soc. N.S.W., 125, 2004 D.P. HIGGINS, G. TOBIAS AND G.M. STONE of 16-17 days. However, as there was no clear pattern in oestrogen excretion, we could not determine whether this reflects concurrent vitelline progesterone and oestrogen peaks at 32- 34 day intervals with an interspersed luteal peak at 16-17 days (see figs 1 and 2), or concurrent vitelline progesterone and oestrogen peaks at 16-17 day intervals with an interceding luteal phase with progestin increases below our arbitrary significance criterion. We expect that daily sampling and identification of potentially confounding RIA- reactive faecal steroids would be necessary to resolve this question. However, observations of fetal development add some support to the hypothesis of a 17-day progesterone cycle. Decreasing blood progesterone is a precursor to parturition in many species of eutheria (Rowlands and Wier 1984) and metatheria (Tyndall-Biscoe and Renfree 1987), and to oviposition in many reptilia (Licht 1984). At 17 days the tammar wallaby fetus consists of 17- 20 somite pairs (Griffiths 1984), similar to the 19-20 somite pairs possessed by the echidna at oviposition (Hill and Gatenby 1926; Luckett 1976; Hughes and Carrick 1978). Both young also exhibit similar stages of development at parturition or hatching 11 days later (Griffiths 1984). The similar rates of development in the last third of gestation and incubation suggests that the age of the echidna fetus at oviposition is approximately 17 days, consistent with a luteal phase of 16 to 17 days. The many irregular oestrogen fluctuations we measured could be inherent in the technique or indicative of follicular development and atresia. Hill and Gatenby (1926) described channels, from the vitellus to a well-developed lymphatic sinus in the ovarian medulla and histological features indicative of follicular regression in the platypus. The authors believe that this study provides a Starting point for further work and suggest the further identification of progesterone and oestrogen metabolites and the comparison of faecal steroid concentrations with blood hormone concentrations, urogenital cytology, ultrasonography of the reproductive tract or behaviour in a controlled study accounting for the potential confounding effects of repeated physical or chemical restraint. ACKNOWLEDGEMENTS We thank the staff of the faculty of Veterinary Science, University of Sydney, in particular Michael Lensen, Irene van Ekris, Margaret Byrne and Geoff Dutton; and Taronga Zoo, in particular Margaret Hawkins, Kerry Foster, and Debbie Pritchard for their assistance. We also thank Michele Thums and the journal referees for their comments Proc. Linn. Soc. N.S.W., 125, 2004 on the manuscript. The project would not have been possible without the help of the staff of Australian Mammals, Taronga Zoo and the Taronga Zoo Friends and without the financial assistance of Novartis, Australasia and the Winifred Scott Foundation. REFERENCES Abensperg-Traun, M. (1991) A study of home range, movements and shelter use in adult and juvenile echidnas (Tachyglossus aculeatus) in Western Australian wheatbelt reserves. Australian Mammalogy 14, 13-21. Augee, M.L., Bergin, T.J. and Morris, C. (1978). Observations on behaviour of echidnas at Taronga Zoo. Australian Zoologist 20, 121-129. Augee, M.L., Haley, E.H.M. and Price, I.P. (1975). Movements of echidnas Tachyglossus aculeatus determined by marking - recapture and radio tracking. Australian Wildlife Research 2, 93- 101. Boisvert, M. and Grisham, J. (1988). Reproduction of the short beaked echidna at the Oklahoma City Zoo. International Zoo Yearbook 27, 103-108. Broom, R. (1895). Note on the period of gestation in echidna. Proceedings of the Linnean Society of NSW 10, 576-577. Carrick, F.N. (1977). Studies in the reproductive physiology of male marsupials. PhD thesis, University of New South Wales, Australia. Clarke, I.J. and Doughton, B.W. (1983). Effect of various anaesthetics on the resting plasma concentrations of lutienising hormone, follicle stimulating hormone and prolactin in ovariectomised ewes. Journal of Endocrinology 98, 79-89. Cleva, G.M., Stone, G.M. and Dickens, R.K. (1994). Variation in reproductive parameters in the captive male koala. Reproduction, Fertility and Development 6, 713-719. Curlewis, J.D. (1983). Some interactions between gonadal steroid hormones and target organs in the male and female brush-tail possum (Trichosurus vulpecula). PhD thesis, University of Sydney, Australia. Curlewis, J.D., Axelson, M. and Stone, G.M. (1985). Identification of major steroids in the ovarian and adrenal venous plasma of the brush-tail possum (Trichosurus vulpecula) and changes in peripheral plasma levels of oestrogen and progesterone during the reproductive cycle. Journal of Endocrinology 105, 53-62. Flannery, T. (1990). ‘Mammals of New Guinea.’ (Robert Brown Associates: Australia). Flannery, T. F. and Groves, C. P. (1998). A revision of the genus Zaglossus (Monotremata, Tachyglossidae), with description of new species and subspecies. Mammalia. 62, 367- 396. FAECAL REPRODUCTIVE STEROIDS IN CAPTIVE ECHIDNAS Graham, L.H., Goodrowe, K.L., Raeside, J.I. and Liptrap, R.M. (1995). Non-invasive monitoring of ovarian function in several felid species by measurement of fecal estradiol-17b and progestins. Zoo Biology 14, 223-237. Griffiths, M.E. (1968). ‘The Echidna’ (Pergamon Press: UK). Griffiths, M. (1984). Mammals: Monotremes. In “Marshall’s Physiology of Reproduction Vol 1, 2nd ed’ (Ed G.E. Lamming) pp. 351-385. (Churchill Livingstone: Edinburgh). Hill, J.P. and Gatenby, J.B. (1926). The corpus luteum of the Monotremata. Proceedings of the Zoological Society of London Ml, 715-762. Hindle, J.E. and Hodges, J.K. (1990). Metabolism of oestradiol-17@ and progesterone in the white rhinoceros (Ceratotherium simum simum). Journal of Reproduction and Fertility 90, 571- 580. Hughes, R.L. and Carrick, F.N. (1978). Reproduction in female monotremes. Australian Zoologist 20, 233-254. Lasley, B.L. (1985). Methods for evaluating reproductive function in exotic species. Advances in Veterinary Science, Comparative Medicine 30, 209-228. Licht, P. (1984). Reptiles. In “Marshall’s Physiology of Reproduction Vol 1, 2nd ed’ (Ed G.E. Lamming) pp. 206-282 (Churchill Livingstone: Edinburgh). 286 Luckett, W.P. (1976). Fetal membranes of the Monotremata and the origin of mammalian Viviparity. Anatomical Record 184, 466. Racey, P.A. and Potts, D.M. (1970). Relationship between stored spermatozoa and uterine epithelium in the pipistrelle bat (Pipistrellus pipistrellus) Journal of Reproduction and Fertility 22, 57-63. River, C. and Rivest, S. (1991). Review article. Effect of stress on the activity of the hypothalamic- pituitary- gonadal axis: peripheral and central mechanisms. Biology Reproduction 45, 523- 332. Rowlands, I.W. and Weir, B.J. (1984). Mammals: non primate eutherians. In ‘Marshall’s Physiology of Reproduction Vol 1, 2nd ed’ (Ed GE. Lamming) pp. 455-658. (Churchill Livingstone: Edinburgh). Spanner, A., Stone, G. M.and Shultz, D. (1997). Excretion profiles of some reproductive steroids in the faeces of captive Nepalese red panda. Reproduction, Fertility and Development 9, 565-570. Tyndale-Biscoe, H. (1973). “Life of marsupials’ (Edward Arnold: Australia). Tyndale-Biscoe, H. and Renfree, M. (1987). ‘Reproductive physiology of marsupials; monographs on marsupial biology.’ (Cambridge University Press: Cambridge). Wimsatt, W.A. (1969). Some interrelations of reproduction and hibernation in mammals. Symposia of the Society for Experimental Biology 23, 511-549. Proc. Linn. Soc. N.S.W., 125, 2004 Anatomy of the Central Nervous System of the Australian Echidna M. Hassiotis!, G. PAxINos? AND K.W.S. ASHWELL!* ‘Department of Anatomy, School of Medical Sciences, The University of New South Wales, 2052, Sydney, NSW, Australia. Prince of Wales Medical Research Institute, The University of New South Wales, 2052, Sydney, NSW, Australia. *author to whom correspondence and proofs should be addressed. Postal address as above. Fax: 61 2 9385 8016, Phone: 61 2 9385 2482 Email: k.ashwell@unsw.edu.au Hassiotis, M., Paxinos, G. and Ashwell, K.W.S. (2004). Anatomy of the central nervous system of the Australian echidna. Proceedings of the Linnean Society of New South wales 125, 287-300. Even from their gross appearance, the brain and spinal cord of the Australian echidna show unusual features. The spinal cord is one of the shortest ever recorded for any mammal, ending at the mid-thoracic level, a feature which may be related to the defensive posture of the echidna. The pattern of termination of unmyelinated afferents in the spinal cord as revealed by lectin labelling with the B4 isolectin from Griffonia simplicifolia is also quite different from that seen in placental mammals, with termination in patches within deeper layers of the dorsal horn. Within the brainstem, specializations of the trigeminal system are apparent with great enlargement of all trigeminal nuclei. The mesencephalic trigeminal nucleus also shows an unusual aggregation of neurons in a central midline position quite unlike therian mammals. While the dorsal thalamus of therian mammals shows compartmentation related to function , the dorsal thalamus of the echidna is remarkable for its lack of cytoarchitecttural differentiation. Most of the high encephalization in this mammal is attributable to the highly gyrified cerebral cortex. This cortex is further distinguished by the positioning of the major functional areas (primary motor, somatosensory, visual and auditory areas) towards the caudal pole of the brain. Manuscript received 21 July 2003, accepted for publication 22 October 2003. KEYWORDS: cerebral cortex, echidna, monotreme, spinal cord, thalamus, trigeminal. INTRODUCTION In this paper we will be reviewing what is known about the anatomy of the central nervous system of the Australian echidna (Tachyglossus aculeatus), with special reference to those features with functional relevance. Even at the level of gross inspection, the central nervous system of the echidna is remarkable for the large size of the brain and the short relative length of the spinal cord. PERIPHERAL RECEPTORS AND ELECTRORECEPTION One of the most remarkable features of monotreme neurobiology, and one which touches on trigeminal nuclei development and cortical organization, is the reported presence of electroreception in two members of this subclass (short-beaked echidna and platypus) (Iggo et al. 1985; Scheich et al. 1986; Gregory et al. 1987, 1988, 1989; Proske et al. 1998). This sensory modality utilizes the trigeminal system in both monotremes studied. To date, physiological and anatomical studies of peripheral sensory systems in this animal have concentrated on peripheral receptors of the trigeminal system. The short-beaked echidna is known to use its sensitive snout as its major sensory tool. Anatomical studies of this snout have revealed a rich distribution of unusual receptors on the tip (Andres et al. 1991; Manger and Hughes 1992). One of these, the gland duct receptor system (Andres et al. 1991) or mucous sensory gland (Manger and Hughes 1992), is present in both platypus and echidna and is thought to be involved in electroreception (Iggo et al. 1985; Scheich et al. 1986; Gregory et al. 1987, 1988, 1989). Despite this attention to snout receptors, very little attention has been given to the structure or function of central trigeminal pathways in any monotreme. ECHIDNA CENTRAL NERVOUS SYSTEM SPINAL CORD ANATOMY The spinal cord of the Australian echidna was examined by Ashwell and Zhang (1997). Even at the gross level, the spinal cord is notable because of its relatively short length, terminating at the level of the seventh thoracic vertebra (Figure 1a)(cf human spinal cord which terminates at the intervertebral disc between lumbar vertebrae and 2). This may represent an adaptation to the pronounced vertebral flexure, which this mammal achieves when it adopts its defensive posture (Figure 1b). Since the spinal cord lies posterior to the vertebral column, extreme flexion would place the neural elements (spinal cord and cauda equina) under considerable tension, amounting to an increase of 15% in length or 6 cm in a large adult. The cauda equina in this animal is collectively as thick as the spinal cord, but consists of multiple nerve bundles aa Echidna - ambulatory posture Cauda equina and spinal nerves b Echidna - defensive posture Hig Cauda equina and spinal nerves Spinal Cord Figure 1. The spinal cord of the Australian echidna is very short, ending opposite the seventh thoracic vertebra when the animal is in the ambulatory posture (a). This may be an adaptation which allows pronounced flexing of the vertebral column in the defensive posture (b), since the spinal nerves of the cauda equina would be more tolerant of the stretching associated with flexing of the vertebral column than the much thicker and more vascular spinal cord. Figures c and d show features of the spinal cord reported in Ashwell and Zhang (1997). Please see that paper for ethical clearance details and tissue preparation methods. Figure 1c shows the large neurons of the median nuclear group (arrowhead) at the lower lumbar level of the spinal cord (L4). The small inset indicates the position of the larger image. CC — central canal of spinal cord: DC —- dorsal column; VC — ventral column; VH - ventral horn. Figure 1d shows unmyelinated afferent fibres labelled with a peroxidase conjugated B4 isolectin from Griffonia simplicifolia. The small inset indicates the position of the larger image. These afferents enter via Lissauer’s zone (LZ) and some descend to deep layers of the dorsal horn (arrowhead) terminating in the nucleus proprius (NP), unlike unmyelinated afferents to therian spinal cord, which are confined to the superficial layers; e.g. marginal zone — MZ; substantia gelatinosa — SG). 288 Proc. Linn. Soc. N.S.W., 125, 2004 M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL which are free to move independently of each other, unlike the spinal cord where individual axons are tightly bound together and are surrounded by delicate capillaries. Therefore the stretching of neural elements associated with extreme vertebral flexure, which is not only large of itself but also affects different segmental nerve roots to a greater or lesser extent, is perhaps more easily accommodated by shifting more of the nerve pathway length into the cauda equina. At a histological level, the spinal cord was found to have similar cytoarchitectural features characterising the laminar organization to that seen in the spinal cords of eutherian mammals (Ashwell and Zhang 1977). Spinal cord nuclei found in eutherians were also identified in the monotreme, except for the central cervical nucleus. In addition, a distinct group of large neurons, named the median nuclear group, was identified in the ventral part of Rexed’s lamina X and extending into the ventral funiculus at the lower lumbar level (Figure 1c). Fibre calibre in the dorsal and ventral roots of the echidna was similar to that reported in eutheria, suggesting similar proportions of afferent fibre classes and « and B motorneurons. The distribution of unmyelinated primary afferent fibres within the dorsal horn of the echidna spinal cord have been examined using lectin labelling with Griffonia simplicifolia isolectin B4. When conjugated with horseradish peroxidase, GSB4 is known to label unmyelinated primary afferents terminating in both the dorsal horn and cranial nerve sensory nuclei (Streit et al. 1985; Plenderleith et al. 1989; Ashwell and Zhang 1997). It was seen that the pattern of labelling with this lectin within the spinal cord differed significantly from that seen in eutheria in several respects. Firstly, while labelling was seen within layers I and II of the echidna dorsal horn (similar to eutheria, Streit et al. 1985; Plenderleith et al. 1989), labelled fibre bundles were also seen coursing around the lateral margin of the dorsal horn as well as through layers I and II to terminate in deeper layers of the echidna dorsal horn (Figure 1d). In eutheria, lectin labelled primary afferents terminate only in the superficial layers of the dorsal horn (Streit et al. 1985; Plenderleith et al. 1989). This deeper labelling in the echidna was found to consist of discrete patches in the central and lateral parts of layers III and V (corresponding to the nucleus proprius). Furthermore, in upper cervical segments of the echidna spinal cord, labelled axons were identified coursing around the margins of the dorsal columns to terminate in the internal basilar nucleus (Ashwell and Zhang 1997). These two aforementioned features reflect unusual primary afferent termination in the echidna, but the elucidation of the functional significance of these would require electrophysiological studies. Generally Proc. Linn. Soc. N.S.W., 125, 2004 however, spinal cord cytoarchitectural organization seems to be highly conserved across mammals. CORTICOSPINAL TRACT The echidna corticospinal tract (Figure 2a) differs from other mammals (Figure 2b, c, d) in both its position within the brainstem and in the level at which it decussates (Goldby 1939). The tract runs through the cerebral peduncle, decussates in the pons, and continues in the lateral medulla, dorsal to the spinal tract of the trigeminal nerve. At the spinomedullary junction it enters the most posterior part of the lateral column of the spinal cord and has been traced as far caudally as the 24th spinal segment, which corresponds to lower lumbar to upper sacral levels. No evidence has been found for the presence of a pyramidal tract close to the ventral midline of the medulla, nor for a decussation in the usual position at the caudal end of the medulla, as seen in most eutheria. In no other mammal is the pyramidal decussation as high as in the echidna, nor does the tract, after decussation, lie in such an extreme lateral position as in this monotreme. Itis of interest to note, however, that a high decussation of the pyramidal tract is particularly characteristic of a small number of highly specialised mammals, which probably developed these corticospinal specialisations at a very early period in mammalian evolution (Goldby 1939). For example, some bats and edentates have a decussation just caudal to the pons and there is a tendency in some of these mammals for fibres from this high decussation to take up a lateral position in the medulla, e.g. in an armadillo, Lysiurus unicinictus, and the pangolin, Manis tricuspis (Goldby 1939). Since both of these eutherians are capable of pronounced vertebral flexure, as is the echidna, one is tempted to speculate that a high pyramidal tract decussation may be advantageous for mammals which use this type of defensive posture, although the precise nature of the advantage which this may confer is not clear at present. In polyprotodontid metatheria e.g. the American opossum Didelphis virginiana, the corticobulbar and corticospinal tracts have been shown to be small and probably extend no further than the upper cervical segments of the spinal cord (Turner 1924, see also review by Heffner and Masterton 1983) and yet as noted above the pyramidal tract in the echidna is much more extensive. Among eutherians, both hedgehogs and tree shrews (Figure 2c) show termination of the corticospinal tract at higher segmental levels (upper cervical for the hedgehog and midthoracic for the tree shrew, for review see Heffner and Masterton 1983) than that seen in the echidna. These observations have made the extensive and 289 ECHIDNA CENTRAL NERVOUS SYSTEM unusual corticospinal pathway of the surviving monotremes of particular interest. Extension of the corticospinal tract down the greater length of the spinal a) Echidna b) Marsupial (e.g. Phascolartus or Pseudochirus) »s Ke : RY cord is usually regarded as a feature of s advanced neurological organization, as re) seen in primates (Figure 2d) and carnivores, because it allows direct control of the cerebral cortex over motor units within many levels of the spinal cord. ay : VAG BRAINSTEM AND HYPOTHALAMUS The gracile and cuneate nuclei are extraordinarily large in the echidna (Figure 3a), reflecting the well- developed somatosensory pathways for the limbs of the echidna. Furthermore, estimates of the proportion of white matter in the dorsal columns to total white matter in the cord gave an average result of 25%, suggesting well-developed trunk and appendicular somatosensory pathways comparable in development to carnivores and primates (Ashwell and Zhang 1997). In absolute terms the dorsal column pathway is as large as that in the domestic cat and Macaca fuscata - therians of similar body weight. This degree of development of the dorsal columns ranks the echidna among the most neurologically specialized primates with well-developed discriminative tactile sense. Perhaps the high level of somatosensory development can be attributed to dense innervation of the echidna’s forelimb (Mahns et al. 2003) Cc Oo r d c) Edentates and Chiroptera d) Human oh Cera ors, Diencephalon Figure 2 Diagrammatic summary of the course, size and extent of the corticospinal tract (bold) in representative mammals. Note that the corticospinal tract in the echidna is large, has a high decussation and extends to caudal levels of the spinal cord. Contrast this with the small size of the corticospinal tract in marsupials (b) and bats and edentates (c) and restriction of the tract to upper segmental levels of the spinal cord in those mammals, In size and extent, the echidna corticospinal tract is more like that seen in primates (d) and carnivores: mammals in whom a long and large corticospinal tract is believed to confer neurological advantages in the form of direct cortical control of motor units in the spinal cord. The corticospinal tract of the echidna also has a relatively high level of decussation (crossing over) compared to therian mammals, although some mammals (e.g. edentates and chiroptera - c) with the ability to flex their vertebral column also have a high level of decussation. No undecussated ventral corticospinal tract, as seen in primates (d) has been reported in the echidna. Data for the echidna is derived from Goldby (1939), while data for other mammals comes from Kappers, Huber and Crosby (1960). 290 Proc. Linn. Soc. N.S.W., 125, 2004 M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL Figure 3. Coronal cryostat section (40 um thickness) through the brainstem of an echidna stained for cytochrome oxidase by the Wong Riley technique (Wong-Riley 1979)(a, b) and Nissl substance (c, d). Please see Hassiotis and Ashwell (2003) for details of experimental ethics and animal acquisition. Strong cytochrome oxidase reactivity demonstrates the presence of high densities of mitochondria in axon terminals of major sensory pathways for limb and trunk somatosensory pathways (e.g. cuneate nucleus) and cranial somatosensory pathway (e.g. nucleus of the trigeminal spinal tract). The mesencephalic nucleus of the trigeminal nerve occupies a midline position dorsal to the cerebral aqueduct. The inset in c indicates the position of d. 3 — oculomotor nucleus; 4v — fourth ventricle; 12 — hypoglossal nucleus; Aq — cerebral aqueduct; Cu — cuneate nucleus; IO — inferior olivary nuclear complex; mcp- middle cerebral peduncle; Pn — pontine nuclei; SC — superior colliculus; t5 — trigeminal spinal tract; Vc —-caudal part of the nucleus of the trigeminal spinal tract ; Ve — vestibular nuclei; Vmes — mesencephalic nucleus of the trigeminal nerve; Vo -oralis part of the nucleus of the trigeminal spinal tract. or spines, although this has never been studied histologically. Alternatively, this specialization may have arisen because the echidna spends time in subterranean channels, where visual and auditory input are of little benefit, and the sense of smell and touch are of the most value. At present there are no morphological studies of echidna postcranial tactile Proc. Linn. Soc. N.S.W., 125, 2004 receptors available to shed light on this. The trigeminal nerve is also greatly enlarged in the echidna as are the nuclei of the trigeminal spinal tract (Figure 3a, b). This is consistent with the impression from behavioural and electroreception studies that the echidna’s snout is extremely sensitive (see above). The trigeminal system in the echidna 291 ECHIDNA CENTRAL NERVOUS SYSTEM displays a high degree of specialisation similar in kind to that seen in Ornithornychus, but not to such a large extent. In other words, it does not appear to be as sensitive an electroreceptive tool as the platypus bill (Proske et al. 1998). Another note-worthy feature is that the motor nucleus of the trigeminal nerve in the echidna brainstem is much larger than would be expected in an animal whose jaw musculature is so poorly developed (Abbie 1934). The mesencephalic nucleus of the fifth nerve in echidna is very like that seen in reptiles in that it adopts an almost exclusively mid-line distribution (Abbie 1934, Figure 3c, d). Metatheria exhibit a condition intermediate between that of the echidna and eutheria with more extensive development of the lateral mesencephalic V extensions. The mesencephalic nucleus and root of the fifth nerve are generally considered as being concerned with proprioceptive sensibility of jaw musculature. Since the echidna has very poor jaw musculature, such a pronounced development of mesencephalic V is inexplicable. The echidna auditory and vestibular apparatus are also notable. In Ornithorhynchus, the entire labyrinth has been described as being typically avian (Gray 1908). In the echidna, the inner ear shows dissimilarities to therians, in that the echidna cochlea is banana shaped and has only half a turn, hence is partially coiled, whereas in humans the cochlea has two and a half turns, and is fully coiled (Gray 1908). The cochlea also shows maximal response to sound of about 5kHz, substantially lower than in eutheria (Augee and Gooden 1993). It has been proposed that when the cochlea evolved from the primitive labyrinth, it employed the existing vestibular connections within the brain, and that when the cochlear apparatus attained a mammalian level of structural specialization, a trapezoid body appeared in the brainstem. The trapezoid body in the echidna is so rudimentary that it reveals its primitive vestibular and primarily trigeminal origin, because it consists almost entirely of vestibular parts and external arcuate fibres which include a large trigeminal element (Winkler 1921; Abbie 1934). In therians, the pronounced increase in auditory fibres almost completely obscures the original trigeminal and vestibular connection. Winkler (1921) has argued that the poor cochlear development in the echidna renders the vestibular fibres relatively conspicuous. While central auditory pathways have never been closely examined in the echidna, these observations suggest that those pathways are either organized differently or not as extensive as in theria. The hypothalamus in the echidna has been reported to have few striking features (Abbie 1934). The mammalian hypothalamus is very old 292 phylogenetically (Simerly 1995) and very conservative in structure throughout the vertebrate series. One peculiarity, which links the echidna hypothalamus with that of reptiles, and is in sharp contrast to the majority of mammals, is the extremely poor development or possible absence of the echidna mammillothalamic tract (Abbie 1934). Regidor and Divac (1987) for example found no evidence of the mammillothalamic tract in the echidna on examination of myelin-stained coronal sections. This pathway is a key link in the Papez circuit underlying memory and emotions in eutheria. Its poor development in the echidna may indicate that this mammal has an alternative circuit for these functions. THE ABSENCE OF A CLAUSTRUM IN THE FOREBRAIN The absence of a claustrum in the echidna was initially noted by Abbie (1940) and by Divac and co-workers (Divac et al. 1987a) and was further discussed more recently (Butler et al. 2002). Similarly, no claustrum has been identified in the platypus brain (Butler et al. 2002). This structure has been identified in all therian mammals so far examined (Johnson et al. 1994) and is believed to have a structural and chemical affinity with the neocortex, although its precise functional significance is uncertain. It engages in reciprocal connections with neocortex and receives projections from the non-specific intralaminar nuclei of the thalamus (see Butler et al. 2002 for review). The question remains open as to whether the claustrum was present in ancestral mammals and disappeared in the monotremes, or whether its evolution represents an exclusively therian brain development. THE DISTRIBUTION OF CHEMICALLY IDENTIFIED NEURONS Manger and co-workers have recently examined the distribution of cholinergic, catecholaminergic and serotonergic neurons in the brains of the platypus and echidna (Manger et al. 2002a, b, c). Those authors showed that while there are many similarities between monotremes and therians in the distribution of these neurons, there were also- some evolutionarily and potentially functionally significant differences. For example, cholinergic cells are present in the monotreme brain, but important cells groups identified in theria do not appear to be present in the platypus or echidna. These include cholinergic cells in the cerebral cortex, nuclei of the vertical and Proc. Linn. Soc. N.S.W., 125, 2004 M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL Figure 4. Coronal cryostat sections (40 um thickness) through the caudal thalamus (a) and rostral thalamus (b) stained for cytochrome oxidase and Nissl substance, respectively. Please see Hassiotis and Ashwell (2003) for details of experimental ethics and animal acquisition. Note the large ventral posterior thalamic nucleus with lateral (VPL) and medial (VPM) compartments. In theria these two regions serve processing of somatosensory (touch) information from the body and head, respectively. Contrast the size of these two nuclei with the lateral geniculate nucleus (LG) processing visual information. The reticular thalamic nucleus, which is found in all therian mammals external to the ventral tier nuclei (i.e. embedded in the external medullary lamina to the left of 4a), appears to be absent from the echidna thalamus. The rostral thalamus (b) contains a large nucleus (anteromediodorsal -AMD) with no clear division into subnuclei. This may correspond to the mediodorsal nucleus of therians. 3v — third ventricle; eml — external medullary lamina; ml — medial lemniscus; ZI — zona incerta. horizontal limbs of the diagonal band of Broca, the magnocellular preoptic nucleus, the substantia innominata, nucleus of the ansa lenticularis, hypothalamic nuclei and the parabigeminal nucleus (Manger et al. 2002a). They proposed that the absence of cholinergic neurons from the hypothalamus might be related to the unusual features of monotreme sleep (Siegel et al. 1996, 1998). The catecholaminergic system of the monotreme brain appears to be very similar to that found in theria, but there were some minor differences in the form of the absence of A4, A3 and C3 groups from the locus coeruleus and caudal rhombencephalon. It should be noted however, that these are only small differences and this great similarity demonstrates the high degree of evolutionary conservatism in these neurons across amniote species (Manger et al. 2002b). Proc. Linn. Soc. N.S.W., 125, 2004 Serotonergic neurons in monotremes appear to fall into three groups: hypothalamic, rostral nuclear and caudal nuclear clusters. The rostral and caudal nuclear groups are found consistently across all mammals while the hypothalamic cluster, although not reported in other mammals, is found in most other species of vertebrates (Manger et al. 2002c). THE THALAMUS AND THALAMOCORTICAL PROJECTIONS Campbell and Hayhow (1971) identified several thalamic nuclei in echidna, which exhibited cyto- and myeloarchitectonic features resembling those found in other mammals (Figure 4). However, echidna thalamic nuclei are not as easily distinguished as those 293 ECHIDNA CENTRAL NERVOUS SYSTEM in opossums (Bodian 1939, 1942; Oswaldo-Cruz and Rocha-Miranda 1968; Benevento and Ebner 1971) or other commonly used laboratory mammals (Rose 1942; Rose and Woolsey 1949). Chemoarchitectural characteristics of the thalamus in echidnas and rats have been compared in sections stained for myelin, acetylcholinesterase (AChE), succinate dehydrogenase (SDH) and cytochrome oxidase (CO) by Regidor and Divac (1987). Numerous species differences were noted, but in general the thalamus is architecturally more homogenous in echidnas than in rats, especially within the anterior portion (Figure 4b). The large structure localized in the anteromediodorsal part of the thalamus of the echidna has been found to contain small amounts of acetylcholinesterase and oxidative enzymes; in this respect resembling the mediodorsal nucleus of rats. Regidor and Divac (1987) concluded that this brain structure of echidnas corresponds to the mediodorsal nucleus in placental species. Welker and Lende (1980) defined and described the thalamic nuclei that contribute major projections to the isocortex in echidna. Their purpose was to determine whether the echidna thalamus exhibited mammalian thalamocortical relations more similar to those found in metatheria, or to those in eutherian mammals. Welker and Lende also attempted to identify whether an enlarged thalamic nucleus was sending afferents to the enlarged frontal cortex. They performed a series of partial ablations of the somatic sensory, auditory, visual and motor areas, as well as in several different portions of the greatly enlarged frontal neocortex (see below) and demonstrated that the thalamocortical connections in the echidna are similar in most respects to those demonstrated in eutherian mammals. One unusual feature observed by Welker and Lende was a large nuclear mass in the dorso-fronto-medial thalamus (presumptive anteromediodorsal nucleus discussed above), which projects to the enlarged frontal cortex (Divac et al. 1987a, b). It has been hypothesised that this nuclear region is homologous to the eutherian mediodorsal nucleus. Their data also revealed that projections to separate motor and somatic sensory cortical areas from the thalamus were spatially distinct (Welker and Lende 1980). CORTICAL STRUCTURE AND ORGANIZATION Until the late 1800’s it was generally believed that all mammals possessed a corpus callosum (Turner 1890), a major fibre bundle connecting the neocortex of the two hemispheres of the brain. Elliott Smith 294 (1902, 1903) dispelled this notion in his early studies of comparative cortical organization. He showed that in monotremes and metatheria, the anterior commissure is the major cerebral commissure, being the sole connection between all parts of the neo- and paleocortex, with only a small archicortical commissural connection being present dorsally (the hippocampal commissure). Several striking aspects of gross cortical anatomy have been noted in Tachyglossus aculeatus. The most obvious of these is the high degree of gyrification (36% of isocortex buried in fissures), comparable to that in many eutherian mammals (e.g. cat 40%, squirrel monkey 39%). The second is the large proportion of the brain volume occupied by the cerebral cortex (43%), similar to values in eutheria (prosimians — 54%, Pirlot and Nelson 1978). Among the brains of eutheria, a highly gyrified cerebral cortex is usually considered as an attempt to maximise the number of cortical columns available for the processing of information. Therefore a highly gyrified cortex is considered the hallmark of more neurologically advanced mammals such as carnivores, primates and cetaceans. This raises the question as to why an animal like the echidna, which leads a solitary existence and has no known complex social life, has such a highly gyrified cortex. One principal difference between the — brains of the two living Australian monotremes is that the platypus’ cortex is quite smooth (lissencephalic), whereas the echidna cortex is complexly folded. Another most remarkable aspect of echidna neurobiology concerns cortical topography. Ziehen (1897, 1908), Brodmann (1909) and Schuster (1910), all published early observations on the cortex in the Monotremata. Brodmann examined the cortex in echidna and established that there is a typical six- layered distribution. The echidna has been noted to have a thinner cortex, perhaps due to its denser packing of neurons compared to the platypus (Abbie 1940). In both the echidna and the platypus, Ziehen (1897, 1908) showed that there was a change in the type of cortex between the anterior (olfactory) and posterior (sensorimotor) portions of the hemisphere, and Schuster (1910) confirmed his observations. Nevertheless, these early authors concluded that the plan upon which the monotreme brain is constructed conforms in every respect to the basic pattern prevailing among the vast majority of other mammals (Abbie 1940). To date, the most detailed anatomical study of the echidna cortex was performed in the 1940’s by Abbie. Since the 1940’s, no further in-depth anatomical studies have been done on the anatomy of the echidna cortex as a whole, although specific systems have been Proc. Linn. Soc. N.S.W., 125, 2004 M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL Welker and Lende (1980) Krubitzer et al. (1995) Rostral Figure 5. Electrophysiology and cytoarchitecture of the echidna cerebral cortex. The earliest study illustrated is by Lende (1964)(results shown in a summary diagram redrawn from Welker and Lende 1980). Greek letters denote major consistent sulci as delineated by Smith (1902). The Welker and Lende map shows only the externally visible gyral surface and indicates the position of motor (M), somatosensory (S1), visual (V) and auditory (A) cortices. The Ulinski map shows cytoarchitectural fields identified by that author (Ulinski 1984). Two coronal sections (i and ii) are illustrated with the positions of rostral (r) and caudal (c) fields of the somatosensory cortex (SM1) marked. The small lateral view shows the rostrocaudal positions from which these sections were taken. The figure below the two sections shows a flattened representation of cortex. Note that Ulinski’s “‘r’’ field lies rostral (and superior) to the deepest part of the o sulcus. The lower two illustrations summarize the findings of Krubitzer et al (1995). The smaller diagram shows a representation of the entire flattened cortical surface with the sulcal walls opened. Solid lines in the Krubitzer map indicate the deepest point of the sulci, while dotted lines indicate the sulcal rims at the external cortical surface. The larger illustration shows a drawing of the completely flattened cortical surface with the boundaries of functional areas indicated relative to B and sulci (thick grey lines). Note that the rostral somatosensory field in Krubitzer’s map lies caudal to the @ sulcus (cf Ulinski map). Ent — entorhinal cortex; Hi — hippocampus; M — manipulation cortex; Pir — piriform cortex; PV — parietal ventral somatosensory cortex; R -rostral somatosensory cortex; S1 — primary somatosensory cortex. Proc. Linn. Soc. N.S.W., 125, 2004 295 ECHIDNA CENTRAL NERVOUS SYSTEM studied. Abbie described the monotreme neocortex as comprising two broad components; one related to the hippocampus, labelled by him as parahippocampal regions and located in the anterior and medial parts; and the other related to the piriform cortex, labelled as the parapiriform regions, located posteriorly and laterally. He also defined sulcal boundaries to these regions. When labelling the cortex, Abbie adopted the system of Elliot Smith (1902), using Greek letters to name the major and deepest sulci (Figure 5). There are two pronounced sulci in the monotreme cortex, denoted as m and £. These divide the frontal cortex from the posterior motor and sensory cortices. More recent functional studies of the isocortex of Tachyglossus aculeatus have indicated that the primary motor, somatosensory, auditory and visual areas are located in the caudal half of the isocortex (Lende 1964)(Figure 5). Aside from the posterior location of these areas, the following relationships are unlike those described in any other therian mammals: the somatic sensory area is confined to the ventral portion of the lateral surface; the visual area is located dorsal to the somatic sensory area and borders the representation for the tail; the auditory area is located posterior to the visual and somatic sensory areas and borders the latter at the representation of the back. These relationships might be described as rotational dislocation of the areal relations relative to that found in eutherians in that the somatic sensory area has been displaced downward and backward, the auditory area upward, and the visual area upward and forward (Lende 1964). The somatosensory representation in the echidna is in some respects similar to that of other mammals. The area for the tail is found uppermost and the areas for hind limb, trunk, forelimb, and head are located laterally and ventrally, in that sequence. This is the same as the basic mammalian pattern of somatosensory area | (S1) as established by Woolsey (1952). A relatively large portion of somatosensory cortex in the echidna was found by Lende to be devoted to the head, and particularly the snout and tongue, as might be expected from the ant-eating habits of the echidna (Lende 1964). Physiological studies have indicated that more than 50% of the rostral cortex of the echidna has no attributable primary motor or sensory function and has been considered as an expanded prefrontal cortex (Welker and Lende 1980). If this interpretation is correct, then the proportion of isocortex in - Tachyglossus aculeatus occupied by the prefrontal area exceeds that in humans (29%) Divac et al. (1987a, b)(see section on thalamus and thalamocortical projection in previous pages). Ulinski’s study (1984) examined the 296 cytoarchitecture and thalamic afferents of the somatosensory area (SMI) in the echidna. His findings indicated that SMI contains two cytoarchitectonic fields. A caudal field with a well-developed layer IV present extends across the post @ gyrus and onto the floor of sulcus a The rostral field was reported to extend from the floor of sulcus a onto its rostral bank. It also was reported to have a well-developed layer IV but with a large number of pyramidal neurons in layer V. The remainder of the pre & gyrus was reported to contain a single cytoarchitectonic field with a thin layer IV and layer V heavily populated with larger pyramidal cells. This field corresponded to the physiologically defined motor area M1. Thalamic afferents to somatosensory area were examined by placing pressure injections of horseradish peroxidase into the two architectonic fields. The results indicated that the somatosensory area in Tachyglossus aculeatus contains two cytoarchitectonic fields that resemble areas 3a and 3b in some placental mammals, leading Ulinski to the conclusion that the collection of cytoarchitectonic fields corresponding to areas 4, 3a, and 3b is a basic mammalian character which has been modified in metatherian and many eutherian mammals. In more recent times, Krubitzer et al. (1995) undertook a detailed study of monotreme cortical organization as part of a comparative approach to determining those features of the isocortex which characterise all the major lines of mammalian evolution, More specifically, their investigation was designed to determine the internal organization and number of somatosensory fields in the monotreme isocortex. The isocortices of both monotremes were found to contain four representations of the body surface. A large area that contained neurons predominantly responsive to cutaneous stimulation of the contralateral body surface was identified as the primary somatosensory area (S1). This was found caudal and ventral to the @ sulcus. Another somatosensory field (R) was identified rostral to S1. The topographic organization of R was similar to that found in S1, but neurons in R were responsive most often to light pressure and taps to peripheral body parts. Neurons in cortex located rostral to R were responsive to manipulation of joints and hard taps to the body. This field was termed the manipulation field (M) and occupies the position of the motor cortex identified by Lende. Note that Krubitzer’s M field occupies an area which Ulinski denoted as the rostral somatosensory field and Krubutzer’s R field occupies at least part of Ulinski’s caudal somatosensory field (Ulinski 1984)(Figure 5). Consequently the two studies are not easily reconciled. A parieto-ventral somatosensory field (PV) was also identified by Proc. Linn. Soc. N.S.W., 125, 2004 M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL rostral Figure 6. Nissl stained cryostat sections in the coronal plane through the cerebral cortex and olfactory bulb of the echidna. Please see Hassiotis and Ashwell (2003) for details of experimental ethics and animal acquisition. The inset drawings show: i) a lateral view of the echidna cerebral hemisphere indicating the planes of section shown in a, b, c and d, e, respectively; and ii) a line drawing of a coronal section showing the positions from which a, b, and c were taken. Figure 6a shows a lower power view of motor cortex (M) and the rostral field of somatosensory cortex (R). Figures 6b and c show motor cortex and S1 somatosensory cortex, respectively with layers indicated by Roman numerals. WM - subcortical white matter. Figures 6d and e show low and high power views of the olfactory bulb. Rectangle in d indicates the position of e. Note the lack of a clear tightly grouped monolayer of mitral cells (Mi) as is seen in therian mammals (Switzer and Johnson 1977). EK — ependyma of lateral ventricle; Gl — glomerular layer; [Gr — internal granular layer; IPL — internal plexiform layer; ON — olfactory nerve fibre layer; Pir — piriform cortex. Proc. Linn. Soc. N.S.W., 125, 2004 ; 297 ECHIDNA CENTRAL NERVOUS SYSTEM Krubitzer and was thought to be homologous to its therian counterparts (Krubitzer et al. 1995). The evidence for the existence of four separate somatosensory representations in somatosensory cortex was taken to indicate that cortical organization is more complex in the echidna than had been previously thought. Furthermore, although the two monotreme families have been quite separate for at least 55 million years (Richardson 1987), the similarity of cortical field organization in both monotremes studied suggested either that the original differentiation of sensory fields occurred very early in mammalian evolution, or that the potential for division of somatosensory cortex into numerous fields was highly constrained in evolution, so that both species arrived at the same result independently. Figure 6 shows the cytoarchitecture of the echidna motor and somatosensory cortices. Nomenclature for cortical areas is adopted from Krubitzer et al (1995). As in eutherian motor cortex (Figure 6b), the echidna M cortex is characterised by large pyramidal neurons in layer V. The S1 part of somatosensory cortex (Figure 6c) is characterised by a layer IV rich in densely packed small neurons. Several other aspects of cortical function in this species are also of note; particularly the apparent absence of an SII somatosensory representation and the relatively lateral position of the motor representation compared to that in eutheria (Krubitzer et al. 1995). Until recently, functional studies had failed to identify any parietal association cortex, but Krubitzer et al. (1995) was able to identify a topographically discrete, multimodal area between the primary sensory cortical areas, which may represent such an area. CONCLUDING REMARKS There are a number of unusual features of the anatomy of the brain and spinal cord of the echidna. 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(1996) The echidna Tachyglossus aculeatus combines REM and non-REM aspects in a single sleep state: Implications for the evolution of sleep. J. Neuroscience 16, 3500-3506. Siegel, J.M. Manger, P.R., Nienhuis, R., Fahringer, H.M., Pettigrew, J.D. (1998) Monotremes and the evolution of rapid eye movement sleep. Philosophical Transactions of the Royal Society of London Series B 353, 1147-1157. Simerly, R.B. (1995). Anatomical substrates of hypothalamic integration. In “The Rat Nervous System’. (Ed. G. Paxinos) pp. 353-376. (Academic Press, San Diego). Smith, G.E. (1902). “Descriptive and illustrative catalogue of the physiological series’, in The Museum of the Royal College of Surgeons of England, vol II, 2™ edn, Taylor and Francis, Red Lion Court, London, pp. 138-157 Smith, G.E. 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(1908) “Das centralnervensystem der monotremen und marsupialier’, Mikroskopische Anatomie, Semon’s Zoologische Forschungs, Bd 3, Teil 2, S 789-921. Proc. Linn. Soc. N.S.W., 125, 2004 Monotreme Tactile Mechanisms: From Sensory Nerves To Cerebral Cortex Mark J. Rows, D.A. MANS AND V. SAHAI School of Medical Sciences, The University of New South Wales, Sydney 2052, Australia Rowe, M.J., Mahns, D.A. and Sahai, V. (2004). Monotreme tactile mechanisms: from sensory nerves to cerebral cortex. Proceedings of the Linnean Society of New South Wales 125, 301-317. Electrophysiological recordings from single tactile sensory nerve fibres supplying the limb extremities in the echidna (Tachyglossus aculeatus) reveal a remarkable resemblance between monotreme peripheral tactile mechanisms and those of placental mammals. The similarities apply to a concatenation of attributes, including the classification of sensory fibre types and aspects of functional properties and tactile coding capacities. The analysis demonstrates that high-acuity tactile signalling from the distal forelimb in the monotreme is based upon a triad of major tactile fibre classes as is the case for placental mammals. Furthermore, the functional similarity between corresponding classes in monotreme and placental species suggests that peripheral tactile sensory mechanisms are highly conserved across evolutionarily-divergent mammalian orders. Evidence for a unique and striking dependence upon tactile sensory mechanisms in monotremes comes from both behavioural observations on the animals and from the exceptional prominence given to the representation of tactile inputs in the cerebral cortex of these species. In the platypus, for example, almost half of its lissencephalic cortex is allocated to the processing of inputs from the bill. Furthermore, within the specialized area of bill representation in the platypus cortex, the receptive fields of individual neurones are amongst the smallest ever recorded within the somatosensory areas of cortex (often 100) ol oO ale N Counts (Imp) (o) 100 Hz Counts (Imp) Figure 4. Precision of impulse patterning in the responses of echidna RA afferent fibers to vibrotactile stimuli. (A): Impulse traces show the tightly phase-locked pattern of response reflecting the periodicity of the 50 and 100 Hz vibrotactile stimuli. Quantitative measures of phase locking based on the vector strength or resultant, R (see Materials and Methods), were derived from the cycle histograms in (B), constructed to show the distribution of impulse occurrences throughout successive cycles of the vibration stimulus at the two frequencies. The analysis time in each CH corresponds to the cycle period of the vibration (modified from Mahns et al. 2003). class and a class of dynamically-sensitive tactile afferents with a distinctly broader bandwidth of sensitivity (Fig.5) reminiscent of placental PC fibres (Mahns et al. 2003). These putative PC fibres supplying the echidna forepaw had absolute response thresholds as low as ~5 um for the broad range of frequencies from ~50 to 300 Hz (Fig.5C) which may equip the animal to detect small vibratory perturbations set up by termites in either the soil or in timber material encountered in the animal’s use of the forepaw as an exploratory organ. Furthermore, these broad bandwidth vibrotactile sensors would be well-suited to serve as an early-warning system for the detection of ground-borne vibration signalling the movements of any predators or other animals in the vicinity (McIntyre 1980; Mahns et al. 2003). As the PC fibre responses to vibrotactile stimuli remained tightly phaselocked at frequencies up to and beyond 400 Hz, the metronome-like patterning in their discharge (Fig. 5A) would ensure that these fibres retained high acuity for signalling the temporal details of vibrotactile perturbations over a broad bandwidth of frequencies (Mahns et al. 2003). Although the bandwidth of vibrotactile responsiveness in the echidna PC fibres (Fig.5) did not extend to frequencies quite as high as those of placental PC fibres (Mahns et al. 2003) the explanation probably lies in the lower body Proc. Linn. Soc. N.S.W., 125, 2004 temperature of the echidna (~28-32°C; Grigg et al. 1992) rather than a fundamental difference in the receptors; in particular, as Sato (1961) has demonstrated that PC fibres in the cat display a displacement to lower frequencies in both bandwidth and peak sensitivity as temperature is lowered. The use of controlled sinusoidal vibration as a dynamic form of tactile stimulation in these studies permitted the precise quantification of both the frequency and intensive parameters of the stimuli. However, in addition, it provides a form of dynamic tactile stimulation that mimics in a controlled way the vibrational disturbances set up in the skin in association with relative movement between the skin and any textured surface encountered in the tactile exploratory movements of the forelimb. The properties revealed for echidna RA and PC fibres indicate that they are well suited to underpin the echidna’s capacity to signal and code information about textural changes in the ground surface or in the coarseness or fineness of objects encountered, such as sand, gravel or soil, in the tasks of locomotion, digging and burrowing. In summary, it appears that for the distal forepaw of the echidna, the tasks of tactile exploration and perception are based upon a triad of major tactile sensory fibre classes, comprising a broad SA class and both RA and PC classes with functional capacities 307 MONOTREME TACTILE MECHANISMS B 400 Hz Response (imp/sec) 0 20 40 60 80 100 Vibration Amplitude (microns) 40 oe ie fo segpee 1:1 Tuning —= Absolute Vibration Amplitude (microns) ©) 8 Vibration Frequency (HZ) Figure 5. (A): Temporal patterning in the vibrotactile responses of PC-like fibers in the echidna. Impulse traces and peristimulus time histograms (PSTHs) show the metronome-like impulse pattern at the 1:1 response level at vibrotactile frequencies of 50-400 Hz. Each PSTH was constructed from five consecutive responses to 20 cycles of vibration at each of the indicated frequencies. (B and C): Stimulus-response relations and vibrotactile frequency bandwidths for the putative PC class of echidna tactile afferent fiber. (B): Stimulus-response relations for a single PC-like fiber with RF on the lateral aspect of the forepaw glabrous skin, based on plots of the mean response (impulses/second) as a function of vibration amplitude at seven frequencies in the range 10-400 Hz. (C): Plots of absolute (solid lines) and 1:1 tuning thresholds (dashed lines) derived for five PC-like afferent fibers from stimulus-response data of the type shown in (B) (modified from Mahns et al. 2003). resembling those of the corresponding classes in placental mammals. The issue of whether the broad SA class might contain subsets will be resolved only with more detailed morpho-functional correlative analysis on both the receptor endings and the associated sensory nerve fibres. However, the breakdown of the echidna tactile sensory fibres into three broad classes resembling those in placental mammals suggests that peripheral mechanisms for tactile sensation in the distal glabrous skin are highly conserved across different mammalian orders (Mahns et al. 2003). CEREBRAL CORTICAL ORGANIZATION FOR TACTILE PROCESSING IN MONOTREMES The behavioural evidence for the pre- eminence of the tactile sense in the platypus, and 308 probably also in the echidna, has been re-inforced by electrophysiological studies on the organization of the cerebral cortex in these two species. In the platypus in particular, the allocation of neocortical space to tactile processing is quite spectacular as was demonstrated with both evoked potential and single-neuron microelectrode recording studies first undertaken in our laboratory in the 1970s (Bohringer and Rowe 1977; Rowe 1990), and more recently by Krubitzer et al. (1995a). With evoked potential mapping, a brief electrical stimulus delivered at a point on the skin surface activates sensory fibres that generate a _ synchronous input to the areas of cerebral cortex involved in processing information from that source of sensory input. From the cortical surface overlying these areas it is possible to record an evoked potential that is usually biphasic, consisting of an initial positive- going deflection followed by a larger negative-going Proc. Linn. Soc. N.S.W., 125, 2004 M.J. ROWE, D.A. MAHNS AND V. SAHAT ow ames SHS 2 RE sep et edt Re Sie SA +53 es 2s ft 4 Soo e tS S44 ae oes 4 Figure 6. Evoked potentials (positively downwards) recorded from the cortical surface of the platypus following bipolar stimulation of the anterolateral margin of the contralateral bill (1V, 100-microseconds pulse). Each recording was made from the position indicated by the dot at the left of the trace. Dotted lines indicate positions of large blood vessels in frontal region of hemisphere; view of hemisphere and whole brain (inset) from dorsolateral aspect. Stippled areas in inset represent focal projection sites and include sites at which positive-going responses exceed 100uV for bill (B), 30u,V for fore limb (FL) and 10uV for hind limb (HL). Zones between stippling and continuous lines include sites from which smaller responses could be recorded (from Bohringer and Rowe 1977). component (Fig.6, and Bohringer and Rowe 1977). The initial positive-going component is thought to arise from the direct excitatory action of thalamo-cortical afferent input on cortical neurons (Mountcastle and Poggio 1968), and therefore the cortical region from which this component can be recorded is thought to represent the projection focus for that source of input. The evoked potentials illustrated in Fig.6 were recorded in response to stimulation on the anterolateral margin of the bill and reveal that a vast area of the dorsal surface of the contralateral cerebral hemisphere is taken up with the processing of bill inputs. At each recording point indicated by the dots on the main figure, responses were also recorded to forelimb (FL) and hindlimb (HL) stimulation in the platypus allowing the isopotential contour maps for these and the bill Proc. Linn. Soc. N.S.W., 125, 2004 (B) inputs to be constructed on the inset figure, with the stippled areas indicating the focal projection sites for the three sources of input, and the zone between the stippling and continuous line a region from which smaller evoked potentials could be recorded. Single neuron mapping of the somatosensory cortex in monotremes More detailed single-neuron microelectrode recordings from as many as 250 individually- discriminated cortical neurons in up to 67 electrode penetrations in a given experiment on the platypus somatosensory cortex confirmed the highly ordered and complete cortical representation of tactile inputs from the contralateral body, extending from the mid- sagittal region of the hemisphere out to the region of the rhinal sulcus (Figs.7 and 8) on the ventrolateral 309 MONOTREME TACTILE MECHANISMS surface, the one sulcus present as an exception to the lissencephalic state of the platypus cerebral cortex. As the microelectrode mappings were based on inputs generated by light tactile stimulation of the skin surface they confirm the remarkable size of the cortical space devoted to tactile processing, in particular, from the bill. This is important as the electrical stimuli delivered to the skin in evoked potential mappings could have activated a combination of tactile and the putative electroreceptive afferent fibres postulated to be present in the bill of the platypus (see below). The continuity of tactile representation across the somatosensory cortex of the platypus was confirmed in several detailed mappings (Bohringer and Rowe 1977) and is apparent in Fig.8 which shows the tactile receptive fields outlined on the figurines for 60 individual neurons isolated in nine electrode penetrations made in a single anteroposterior plane in the posterior region of the hemisphere. While tactile receptive fields for individual neurons are up to ~15 cm” on regions such as the tail and trunk, those on the distal glabrous skin of the limbs were much smaller, while those on the bill, in particular, its anterior and lateral margins, were no more than Imm in diameter. These represent the smallest tactile receptive fields ever recorded in the cortex and are therefore capable of conferring great precision and fidelity upon the tasks of tactile localization and discrimination involving the bill. Where the electrode penetration was made normal to the cortical surface, the neurons encountered had very similar receptive field locations (Fig.8) indicative of the columnar organization, well described for the cerebral cortex of placental mammals (e.g. Mountcastle 1957), in which neurons of similar functional properties are grouped in columns oriented normal to the surface. In penetrations, such as number eight in Fig.8, that passed obliquely through successive cortical columns in the region of bill representation, there was a remarkably orderly and progressive shift in the representation of the bill surface that is apparent in the enlarged and expanded view of this electrode track in Fig.9, emphasizing once again the striking, fine-grain spatial resolution available within the area of bill representation in the somatosensory cortex of the platypus. The somatosensory cortex of the echidna, mapped by Lende (1964), also has a striking allocation of space to the tongue and snout representations but, Figure 7. Inset shows entry points (filled circles) for 54 microelectrode penetrations into the platypus cortex. The black areas on figurines show the combined receptive field areas for all neurons (up to 15) sampled in each of the penetrations. Dotted lines represent positions of large blood vessels in frontal region of hemisphere; view of hemisphere from dorsolateral aspect (from Bohringer and Rowe 1977). -————-—. —— — a _~ _—— 310 adh & & LOO att mS ~ a) Proc. Linn. Soc. N.S.W., 125, 2004 M.J. ROWE, D.A. MAHNS AND V. SAHAI aot RE 0 DEPTH BELOW CORTICAL SURFACE (mm) Figure 8. (A): Photograph of cortical surface of the platypus from dorsolateral aspect indicating a plane in which 9 penetrations were made. (B): Coronal section through hemisphere at the plane indicated in A, showing the course of the 9 penetrations. (C): Reconstruction of penetrations 1-9 indicating location of each neuron studied (filled circles) and its peripheral receptive field. No fields could be found for the first two neurons in penetration 4. Receptive fields for all neurons in each of the penetrations 7-9 were confined to the shaded areas on each of the associated figurines (from Bohringer and Rowe 1977). in addition, a prominent representation of the distal forearm, presumably reflecting its importance in digging and burrowing. Multiple representation of the body within the monotreme cerebral cortex For both the platypus and the echidna, the early cortical mapping studies in our laboratory (Bohringer and Rowe 1977; Rowe 1990) and that of Lende (1964), led to the conclusion that there was a single body representation in the contralateral cerebral hemisphere, conforming to the so-called primary somatosensory cortex (SJ) of other mammals. No evidence was found for a second representation that Proc. Linn. Soc. N.S.W., 125, 2004 might correspond to either the SII area that is well recognized in, for example, the cat and primate species (for review, Rowe 1990; Johnson 1990; Rowe et al. 1996; Zhang et al. 1996, 2001) or indeed, to any other areas of somatosensory representation that have been reported in some placental species (Kaas 1982, 1987; Krubitzer and Kaas 1990; Krubitzer et al. 1995b), and now more recently, for both the echidna and platypus (Krubitzer et al. 1995a). Krubitzer et al. (1995a) have proposed that there are four somatosensory representations within the contralateral cerebral cortex of both the platypus and the echidna which they designate the primary somatosensory cortex (S/), the Rostral deep field (R), the Manipulation field (M) and Sil MONOTREME TACTILE MECHANISMS Figure 9. Coronal section of platypus cerebral cortex showing the position of a microelectrode penetration (track 8 in Fig.8) made obliquely to the cortical surface. The filled circles indicate the locations of the 14 neurons studied. The receptive field for each neuron is indicated on the right which is an enlargement of the area on the ventral surface of the bill (modified from Bohringer and Rowe 1977). the Parietal Ventral field (PV). Furthermore, each is said to contain “a complete representation of the body surface’, a surprising claim to us, considering that the four areas are not all defined as being concerned with the processing of mechanosensory data from the body surface. The area given the S/ designation is concerned principally with cutaneous inputs, whereas area R immediately rostral to SJ is said to contain neurons that respond “most often to stimulation of deep receptors” and which required light taps or light pressure to the body surface to elicit a response. However, as the tactile stimuli employed in their study, whether brushing, tapping, or pressure forms of cutaneous stimulation, were neither quantified nor reproducible, it is difficult to see how reliable distinctions were made in terms of neural response thresholds between neurons in the different cortical representation areas. Topographic organization of the putative multiple areas of somatosensory representation in the monotreme cerebral cortex As responsiveness criteria may be insufficient to permit an unequivocal division of the monotreme somatosensory cortex into four distinct areas (Krubitzer et al. 1995a), such a differentation must therefore depend upon topographic considerations, in particular upon the extent to which the four putative areas constitute complete and discrete representational maps of the body. In the case of adjacent areas, whether, for example, SJ and R, or SI and PV, Krubitzer et al. (1995a) state that the boundaries coincide with a reversal in the representation of peripheral receptive fields as one progresses across a sequence of cortical recording sites. For example, in both platypus and echidna cortex, the receptive fields on the upper body may shift from proximal body locations, including the face, shoulder and upper limb, to the distal forelimb, and then move back to more proximal parts of the limb and shoulder. Similar reversals were seen across sequences of recording sites involving the hindlimb representations, leading to the interpretation by Krubitzer et al. (1995a) that the separate representations of proximal limb and associated trunk regions constitute parts of two distinct. body representations. However, an alternative interpretation must be considered, which emerges from the detailed studies carried out in the macaque monkey by Werner and Whitsel (1968, 1973) and Whitsel et al. (1969, 1971) for S7 in the postcentral gyrus, and for the more laterally-placed second somatosensory area, SII. 1 an ol sco Sb oe oma The representation of the body within the cerebral cortex: a reflection of the dermatomal trajectory The crucial observation emphasized by Werner and Whitsel was that the body maps in both SJ and SII of the cerebral cortex owe their essential topographic properties to the serial and overlapping projection of the dorsal roots. In the case of the macaque postcentral gyrus, the representation of spinal roots, from sacral through to cervical, forms a succession of antero-posteriorly oriented bands progressing from medial to lateral across the cortex (e.g., Figs. 6 and 8 in Werner and Whitsel 1973). Within the SI area of the macaque monkey, and indeed within the macaque S//J as well, the representations of the postaxial and preaxial arm and leg areas are separated by the representation of the more distal parts of the limbs, in particular, the digits and toes respectively. This effectively gives rise to a split representation of the upper parts of the limbs. In the case of the postcentral gyrus, one component lies Proc. Linn. Soc. N.S.W., 125, 2004 M.J. ROWE, D.A. MAHNS AND V. SAHAT medial to the distal limb representation, the other lateral to it, an arrangement represented schematically to illustrate this dermatomal trajectory in Fig.8 of Werner and Whitsel (1973). One may observe how the dermatomal trajectory generates the split representation of the upper regions of either forelimb or hindlimb within the cerebral cortex by examining, in a human anatomy text (e.g., Williams and Warwick 1980) the dermatomal boundaries of successive spinal roots associated with either the lower or upper regions of the body. In the case of the upper body, the ventral and dorsal axial lines of the upper limb mark a border between the innervation fields of C5 and T1. Therefore, in any representation of the body surface within the cerebral cortex, one might expect, with the central map being laid down according to the dermatomal trajectory (Werner and Whitsel 1968, 1973; Whitsel et al. 1969, 1971), that the central representation of the upper arm will be split along the axial lines with the input from the distal limb, carried over the C6, 7 and 8 roots, creating a clear separation in the cortical representation of lateral and medial surfaces of the upper arm. As the same fundamental plan and sequence for tactile dermatomes is also found in both the cat and monkey (Sherrington 1898; Kuhn 1953; Hekmatpanah 1961), one may assume that this organizational plan for spinal segmental innervation is a general one that would operate across mammalian orders, including the monotreme representatives. Is there multiple representation of the body within the platypus and echidna cerebral cortex? If one examines the receptive fields plotted for the platypus and echidna somatosensory cortex by Krubitzer et al. (1995a) in their Figs. 6 and 16, it might be argued that those fields on the proximal parts of the limb, on either side of the distal limb representation, are not clearly and systematically separated into representations of the medial and the lateral surfaces of the limb as might be expected in a perfect reflection of the dermatomal trajectory. However, this is hardly surprising on several grounds that are outlined in a recent review (Rowe, in press). The fundamental point to be emphasized is that the reversals in receptive field representation described by Krubitzer et al. are consistent with the sequence of representation that might be expected within a single body map whose plan is determined by the dermatomal trajectory. In view of these considerations we would re-emphasize our 1977 finding (Bohringer and Rowe 1977; Rowe 1990) of a single large representation of the contralateral body within the platypus cerebral cortex and the similar conclusion reached even earlier by Lende (1964) for Proc. Linn. Soc. N.S.W., 125, 2004 the echidna. Furthermore, we wish to emphasize the © fundamental importance and significance of Werner and Whitsel’s studies (1968, 1973; and Whitsel et al. 1969, 1971) on body representation within the cerebral cortex, and the need that arises from their studies, to take account of the dermatomal trajectory as the crucial determinant of representational topography within central neural systems. Sensory and perceptual specialization in monotremes: trigeminal electroreceptive mechanisms As the monotremes emerged in mammalian evolution on a separate line from therian mammals in the early Mesozoic (Dawson 1983; Rowe 1990; Augee and Gooden 1993) the possibility that some qualitatively different apparatus for neural sensing might have emerged was given some credence in the 1980s with both behavioural and electrophysiological studies reporting the presence of electroreception in monotremes (Scheich et al. 1986; Gregory et al. 1987, 1988, 1989a,b). These and subsequent reports suggested that electroreception might be associated with the bill of the platypus and the snout of the echidna, in each case in association with the trigeminal nerve rather than the lateral line system that is the principal basis of electroreception in certain fish (Cahn 1967; Bullock 1999). However, neither behavioural nor electrophysiological data have provided any suggestion of electroreception in association with other skin regions or somatosensory nerves of the monotremes, such as the median or ulnar nerves of the forelimb. The first behavioural evidence for electroreception in the platypus bill came in a short report from Scheich et al. (1986) that the platypus could detect weak electric fields with threshold strengths as low as 50-200 uV cm. Furthermore, they reported that an electroreceptive processing zone was present in the posterolateral region of the contralateral cerebral cortex on the caudal side, but next to, the map of bill mechanoreceptive input identified by Bohringer and Rowe (1977). In our view this is a puzzling finding on several grounds (for review, see Rowe 1990). First, in an earlier detailed electrophysiological mapping of the cerebral cortex we found that the cortical region concerned with tactile representation of the bill occupied almost the whole of the posterolateral region of cortex (Bohringer and Rowe 1977; Rowe 1990). Furthermore, we constructed separate cortical maps of bill representation based upon mechanical stimulation which was specific for tactile inputs, and electrical stimulation which would have activated both tactile and the putative electroreceptive afferents. 313 MONOTREME TACTILE MECHANISMS However, comparison of the two maps (Figs.6 and 7; and Bohringer and Rowe 1977) reveals no evidence of a separate area of bill representation in the map based upon the electrical stimulation, from that obtained with pure tactile stimulation. In contrast to the Scheich et al. (1986) report that electroreceptive-induced cortical evoked potentials were found next to the map of bill mechanoreceptor input, those by Iggo et al. (1992) and Krubitzer et al. (1995a) indicated that the electrosensory area of cortical representation lay entirely within the border of the tactile representation area defined for the bill in our earlier study (Bohringer and Rowe 1977; Rowe 1990). However, Krubitzer et al. (1995a) have described a “clear functional parcellation” within this SJ region whereby regions responsive to just tactile input were interdigitated with regions responsive to both tactile and electroreceptive inputs (e.g. Fig.10 in Krubitzer et al. 1995a). Furthermore, the regions of pure tactile sensitivity coincided with myelin and cytochrome oxidase-dense regions of SI while the regions of putative bimodal sensitivity coincided with myelin-light regions. What the significance of such differences in myelin density might be in this circumstance remains unclear. However, there may not be universal agreement with this assertion anyway, judging by the allocation of light and dark areas in their Fig.10B; for example, the myelin-dark region drawn to contain four recording sites of pure mechanosensitivity in part B of their figure appears rather paler in the adjacent tangentially- oriented photomicrograph than some areas represented as myelin-light in Fig.10B. It is likely to be difficult to make these distinctions reliably, in the tangentially- cut cortical sections, firstly without objective analysis of image density, and secondly, in the absence of systematic control for laminar depth across the extent of the cortical section under study. Electrosensory field-strength thresholds illustrated for cortical neurons in Fig.10 by Krubitzer et al. (1995a) ranged up to 900 uV cm'; however, it was not clear what field strength might have activated neurons in the purely tactile-sensitive regions. Behavioural and afferent fibre thresholds for electroreception A further concern in relation to claims for electroreception in monotremes arises over the thresholds reported for the phenomenon. At the behavioural level, Scheich et al. (1986) reported field strength values as low ~50-200 uV cm’! for the platypus, and, more recently, values of 20uVcm''! have been reported (Manger and Pettigrew 1996; Pettigrew 1999). However, individual trigeminal afferents, believed to be of the electrosensitive class, had 314 threshold field strengths of ~4mV cm’! (Gregory et al. 1988, 1989b). As these values were based on a substantial fibre sample, and as these values are vastly higher than reported behavioural thresholds for the platypus, one might infer that any putative electroreceptive sense must depend upon some other source of afferent input. Proposals that spatial summation, based upon convergence of a number of trigeminal electroreceptive afferents onto central neurons, may confer the observed behavioural thresholds upon the animal are difficult to accept when none of the sampled afferent fibres has thresholds low enough to account for this. To the extent that there are contentious issues associated with the claims for electroception within the monotreme order of mammals, it is important that further rigorously controlled investigations be pursued to resolve such issues, in particular, more detailed behavioural studies based upon objective analysis rather than anecdotal accounts of the movement patterns of the platypus and echidna in relation to presumed electrosensory stimuli. REFERENCES Augee, M. and Gooden, B. (1993). “Echidnas of Australia and New Guinea’. Sydney, University of New South Wales Press. Belov, K. and Hellman, L. (2003). Platypus immunoglobulin M and the divergence of the two extant monotreme lineages. 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N.S.W., 125, 2004 317 t ee om, ay fe - + | i ; Ly 1 ed eT 8g me) Sai ‘ Bor i udtict i vie ot ie yb ecules i ash mye bey, ¥ ay ~ a ; ae doe ot he i : ihn eu Hon, a Bic it yy ary; bak a. nis Nadie. 1a a Veet SY. a juan) toni ' ‘< 7 6 ety A deka oa ALR. 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CB v4 ae (LA... Ctiemat GT. Adh me f at Raves ee rei A A ; WHT. Cel ReA 5 Mae a af wy bueegte ut ah Ae sag ; the. ie okre: rt vet ain a Sonne! tf 4 leper Negnelpay The Role of Push Rods in Platypus and Echidna — Some Speculations U. PROSKE AND J.E. GREGORY Department of Physiology, Monash University, Clayton, VIC 3800 (uwe.proske @med.monash.edu.au) Proske, U. and Gregory, J.E. (2004). The role of push rods in platypus and echidna — some speculations. Proceedings of the Linean Society of New South Wales 125, 319-326. This is a review of the structure and innervation of the mechanosensory organ, the push-rod, in skin of the platypus bill and echidna snout. Four receptor types can be identified in association with push rods in platypus and echidna: (i) central vesicle chain receptors, (ii) peripheral vesicle chain receptors, (iii) Merkel endings and (iv) paciniform corpuscles. Function of the vesicle chain receptors remains unknown. Merkel endings are known to be slowly adapting with irregular discharge (SAI) while paciniform corpuscles are rapidly adapting vibration-sensitive (RA). Recordings made from echidna nose skin have identified both SAIs and RAs. In addition, responses typical of SAII endings (regular discharge) and rapidly adapting, but vibration insensitive, responses were observed. It was concluded that the push rod in monotremes is not associated with mechanoreceptors unique to the group. Skin of the platypus bill and the echidna nose contains erectile tissue. It is conjectured that blood engorgement inflates the skin to facilitate contact between push rods and the external environment. In addition platypus push-rods have a ring of contractile tissue around their tips which, on contracting, restricts mobility of the rod, perhaps when the platypus leaves the water. Possible cooperative roles between electroreceptors and mechanoreceptors are discussed. Manuscript received 3 September 2003, accepted for publication 22 October 2003. KEYWORDS: Cutaneous receptors; electroreceptors; mechanoreceptors; Merkel receptors; push rod; sensation; slowly adapting; vibration. INTRODUCTION In a recent publication we presented some speculations about the role of electroreceptors in the detection of prey items by platypus and echidna (Proske and Gregory 2003). Here we want to extend these speculations to the mechanoreceptors. This, therefore, is a review account of our own work and that of others and speculations based on those observations. THE PLATYPUS PUSH ROD The bill of the platypus has two prominent sense organ structures, the electroreceptors associated with sensory-innervated mucous glands, and mechanoreceptors associated with the push rods. They are distributed differently, the electroreceptors being lined up in rostro-caudally directed rows, the push rods distributed more or less uniformly across the bill, with particularly high concentrations on the edge of the bill (Fig. 1). In the platypus, although it has not been stated explicitly, it appears that push rods are absent from skin, including glabrous skin, of other parts of the body. In other words, push-rods seem to be specifically associated with the bill, as are the electroreceptors. There have been estimated to be 46,500 push rods in the platypus (Manger and Pettigrew 1996), distributed in the skin covering the outside of the upper and lower bill and lining the mouth (Fig. 1a). They are especially numerous around the margins of the upper bill. The platypus push rods (Fig. 2a) are about 70 um in diameter and 400 um long, spanning nearly the full thickness of the epidermis. They consist of a column of flattened spinous cells attached rather loosely to the surrounding epidermis, which allows them relatively free movement in all directions. The rounded tip of the push rod protrudes slightly above the surrounding skin surface. Each push rod is innervated by 25 to 40 myelinated nerve fibres. These terminate in 4 types of sensory ending. The first type is the central vesicle chain receptor, supplied by 5 to 8 medium sized myelinated axons. The receptor consists of axon terminals running ROLE OF PUSH RODS IN MONOTREMES Figure 1. Dorsal and ventral view of the platypus bill (A) and lateral view of the echidna snout (B). Dots indicate the distribution pattern of push rods, which are more numerous near the edge of the bill and tip of the snout. (A, rearranged from Andres and von During, 1984; B, from Proske, 1997.) vertically up the push rod near the centre, with a regular array of vesicle-like protrusions along their length. The second type, the peripheral vesicle chain receptor, is similar, but the myelinated axons (as many as 18) supplying it are thinner and their terminals are located towards the periphery of the push rod. The function of the vesicle chain receptors is still unknown. Their structure is somewhat reminiscent of Meissner corpuscles found in the glabrous skin of many mammals. Meissner corpuscles consist of a coiled arrangement of endings from a number of myelinated axons that terminate between layers of flattened Schwann cells, and they are known to be rapidly adapting mechanoreceptors. If, as is likely, vesicle chain receptors are mechanosensitive, their disposition in the centre and around the periphery of 320 the push rod appears favourable for detecting compression or bending of the rod, and perhaps in this way the direction in which a stimulus is acting can be signalled. The receptors extend to within just a few cells of the skin surface, giving rise to the suggestion that they would also be well placed to function as thermoreceptors (Catania 1995). The third type of receptor in the push rod is the Merkel cell. Up to 12 Merkel cells are located at the base of each push rod, and each myelinated afferent nerve fibre supplying them branches to supply 6 to 8 Merkel cells. The functional properties of Merkel cell receptor complexes have been well documented in other mammalian and avian species, where they are one of the common receptor types found. They are slowly adapting mechanoreceptors, giving rise to what in other mammals are termed Type I responses characterised by a high degree of variability in the train of impulses discharged when stimulated. Figure 3 shows an example, recorded from the echidna. The fourth type of ending is a group of 3 to 6 paciniform corpuscles, lying in the dermis immediately underneath the column of epidermal cells comprising the push rod. These are one of the most intensively studied types of cutaneous receptor and like the Merkel cell, they are found ubiquitously amongst birds and mammals. They are a rapidly adapting mechanoreceptor giving highly phasic responses to stimulation and able to signal faithfully vibratory stimuli at frequencies up to about 1,000 Hz. Figure 3 shows an example, again recorded from the echidna. The paciniform corpuscles at the bottom of the platypus push rods have been reported to be arranged with their long axes oriented either strictly parallel or perpendicular to the skin surface, and at right angles to each other, thus building up a three - dimensional system for transducing any direction of movement of the push rod (Andres & von During 1984). This further implies that the push rods are free to move and not constrained to displacement in any particular direction. Proc. Linn. Soc. N.S.W., 125, 2004 U. PROSKE AND J.E. GREGORY Figure 2. Schematic representations of push rods in the platypus bill (A) and echidna snout (B), showing the 4 types of sensory terminals: lamellated, paciniform corpuscles (p, Imr), Merkel cells (m, Mc), central vesicle chain receptors (cvc) and peripheral vesicle chain receptors (pvc), the latter 2 having a single label (ver) in B. The platypus push rod is attached rather loosely to the surrounding epidermis, allowing relatively independent movement in all directions, while the echidna push rod appears more firmly anchored in the epidermis. (A, modified from Andres and von During, 1984; B, modified from Andres et al, 1991.) THE ECHIDNA PUSH ROD In the echidna push rods are scattered across the surface of the snout, becoming fewer at its base, in the region where the hairs begin (Fig. 1b). They are especially dense near the tip of the snout, which is also the region where the electroreceptors are found. As for the platypus, there is no evidence of push rods in skin of other parts of the body. The structure of the echidna push rod follows the same plan as that in the platypus with two differences. Its nerve supply is less dense and the column of cells comprising the rod is less mobile than in the platypus — there is a less clearly differentiated Proc. Linn. Soc. N.S.W., 125, 2004 boundary between the column of spinous cells with tonofibrils and the surrounding epidermis. The echidna push rods (Fig. 2b) are smaller than those in the platypus and are innervated by only about half as many myelinated nerve fibres. However, they contain larger numbers of Merkel cells and paciniform corpuscles (Andres et al. 1991). RESPONSE PROPERTIES OF THE RECEPTORS What emerges is that the push rod is a specialised mechanoreceptor complex, having separate receptor systems for signalling steady stimuli and phasic or vibratory stimuli, and if the speculations 321 ROLE OF PUSH RODS IN MONOTREMES Vibration receptor 53 Hz 433 Hz ttt AL WWWWWW Slowly-adapting Type I Figure 3. Responses of single receptors in the echidna snout, shown as action potentials recorded in dissected nerve filaments. The vibration sensitive mechanoreceptor is shown responding in a 1:1 entrained fashion to stimulation at just above threshold amplitude and at the frequencies indicated. The stimulus is represented in the traces below the action potentials. This type of response is characteristic of paciniform corpuscles. The slowly adapting Type I response in the lower part of the figure is typical of Merkel cell receptors. It is characterised by an absence of resting or unstimulated discharge, a sensitivity to stimulus velocity and an adapting, irregular discharge during constant skin indentation. The stimulus is shown in the trace below the train of action potentials, and consisted of a linearly increasing skin indentation to an amplitude that was then held constant. (From Iggo et al, 1996.) about vesicle chain receptors are correct, a third system for signalling the direction in which a mechanical stimulus is acting. In an electrophysiological study of tactile receptors in the echidna, Iggo et al. (1996) recorded from 44 mechanoreceptor afferents with receptive fields in the skin of the snout. The receptors were not identified morphologically but some attempt was made to associate a response with the site of a push rod. 322 While it could not be ascertained with certainty that particular responses arose from within a push rod, two types of responses observed were typical of endings found at the base of push rods and identified as such in other mammals. In our study we classified responses into 4 types, based on similarities to the 4 main types of response seen in other mammals and for which the morphological identity of the receptors has been established. Proc. Linn. Soc. N.S.W., 125, 2004 U. PROSKE AND J.E. GREGORY The first type of receptor was characterised by a rapidly adapting response to mechanical stimulation of the bill skin and a high sensitivity to vibration at frequencies up to about 800 Hz (Fig. 3). This type of response is characteristic of Pacinian and paciniform corpuscles in other mammals and can confidently be ascribed to the paciniform corpuscles found in the snout skin and especially prominent at the base of pushrods. The second type of response was an irregular, slowly adapting discharge (Fig. 3). These SAI responses have been known for some time to be generated in other mammals by Merkel cell receptors, the second ending type found prominently at the base of push rods. The examples encountered in the echidna typically had very small receptive fields, with a diameter well below 100 um and a threshold for a response to skin displacement of 4 um. Such small, low threshold fields are consistent with their belonging to Merkel receptors at the base of push rods that are somewhat shielded from stimuli not directly applied to the push rod tip. The most numerous type of response to mechanical stimulation was a slowly adapting, regular discharge, termed a Type II response in other mammals. These have been identified with Ruffini endings in the dermis, signalling preferentially stretch of the skin. A similar morphological type is present in echidna snout skin, but not in direct association with push rods. The fourth type of response was a rapidly adapting discharge to skin stimulation. These receptors were distinguished from the other rapidly adapting group by not showing a sensitivity to vibration at high frequencies, and they responded well only to frequencies below 300 Hz. The study shed no light on the function of vesicle chain receptors. Perhaps they are the rapidly adapting receptors unresponsive to high frequencies of vibration, but the number found seemed too few, at only about 15% of the sample studied. It may be that one or both types of vesicle chain receptor are responsible for at least some of the Type II slowly adapting responses, which accounted for nearly half the sample studied. What can be said is that no completely new mechanoreceptive responses, unknown in other species, were seen in the echidna, either in this study or in an earlier one by the same group (Iggo et al. 1985). There is some evidence that the sensitivity of the push rod complex to external stimuli is not fixed, but can be varied, and in two ways. First, in both echidna and platypus, there is a venous cavernous system in the bill or snout, beneath the skin. It is postulated that engorgement of the venous sinus may Proc. Linn. Soc. N.S.W., 125, 2004 change the mechanical properties of the skin, affecting ° transmission of mechanical stimuli to the receptors at the base of the push rod and effectively changing its sensitivity. It may also lead to protrusion of rod tips beyond the skin surface, to allow them to present more effectively to environmental stimuli. Secondly, Manger et al. (1998) have described a ring of contractile material around the tip of the platypus push rod. Perhaps this is used to restrict the mobility of the push rod and in this way also change its effective sensitivity. Why might it be desirable to change the sensitivity of the push rod by altering the mechanical coupling between the stimulus and the receptors? Perhaps the answer for the platypus lies in the different environments, air and water, the animal inhabits. An appropriate sensitivity for one medium may not be optimal for the other and the platypus may have evolved a means of adjusting between the two. A similar argument could apply to the echidna, which uses the tactile receptors in the snout both in the ground and above it. However there are no reports of the presence of contractile cells in association with echidna push rods, which, anyway, seem to be much less mechanically independent of the surrounding skin. COMPARISONS WITH OTHER ANIMALS The push rod theme reaches its most exuberant expression in the star-nosed mole. Protruding from the snout of this animal is an extraordinary array of 22 radiating fleshy pink fingers entirely covered with thousands of small domes. These domes are the tips of Eimer’s organs, which have a remarkable similarity to the push rods in monotremes, except for containing fewer sensory endings and being innervated by fewer sensory nerve fibres than push rods (Fig. 4). There is a single Merkel receptor at the base and a single lamellated receptor immediately underneath in the dermis. A single vesiculated nerve terminal runs up the centre of the Eimer’s organ and 5 - 10 at the periphery (Catania 1995). Eimer’s organs are believed to be tactile sense organs, responsive to the onset and offset of depression of the papilla and to sustained compression (Catania & Kaas 1995). The star-nosed mole is a burrowing insectivore living almost entirely underground in swamps and bogs. As in the monotremes, the function of the vesiculated receptor terminals has not been conclusively established. It is assumed that the star is a sensitive tactile organ used by the mole to explore its environment for prey. Some other members of the family Talpidae, moles, shrew moles and desmans, have Eimer’s organs (Catania 2000), but only the star- 323 ROLE OF PUSH RODS IN MONOTREMES IN (pe % erie es) .g ue Ap SANA teed ~ Vesicle chain-like receptor Bx . va 4 17 Nt Ss EATEN EIT EN EA) MeN EN DEN DEN DEN SON MS Noe SBN Merkel cell a ANS, Ys Veployk Lamellated corpuscle eae SAIS mA IRAIN tite Figure 4. Schematic of an Eimer’s organ in the star-nosed mole, showing the single Merkel cell and lamellated corpuscle, as well as the central and peripheral neural processes similar to the central and peripheral vesicle chain receptors of the monotreme push rod. (Modified from Catania, 1995.) nosed mole has developed such an elaborate structure to deploy them. FUNCTIONAL CONSIDERATIONS If it is accepted that push rods subserve a mechanosensory function and are designed to raise fidelity of transmission of static and dynamic stimuli, it poses the question of why such an arrangement is needed. Speculation about the mechanoreceptors in monotremes has focussed on two aspects. WwW No - In the platypus, there is a close association between the electrosensory and tactile senses, and cells in the cerebral cortex have been found that receive inputs from both modalities (Iggo et al. 1992; Manger et al. 1996). Pettigrew et al. (1998) have postulated a system for determining the distance and perhaps direction of live objects underwater, based on a cooperation between the two senses. An animal like a shrimp would generate both an electrical pulse and a mechanical pressure wave when it flicked its tail. The electrical pulse would arrive at the platypus first and Proc. Linn. Soc. N.S.W., 125, 2004 U. PROSKE AND J.E. GREGORY be detected by the electroreceptors. After a delay depending on the shrimp’s distance, the mechanical pressure wave would arrive and the receptors in the pushrods would be stimulated. Suitably tuned cells in the cortex would detect the delay between the two inputs, and thus distance could be computed. Directional information could also be derived if the push rods have a directional sensitivity as speculated, and this would be combined with the hypothesised directional information derived from _ the electroreceptors (Manger et al. 1996). A second possible role for push rods concerns our speculation about close-range electrolocation (Proske and Gregory, 2003). As the platypus fossicks about on the bottom of a stream or pond, pushing its bill into the detritus looking for live prey, its electroreceptors will allow it to distinguish animate from inanimate objects. It is conceivable that the alerting signal from the electroreceptors receives confirmation of the presence of a prey item from the detailed mechanosensory signals provided by push rods. Similarly for the echidna, there may be a functional cooperativity between inputs from electroreceptors and mechanoreceptors as the animal pushes its nose into the soil in its search for prey. Speculating more broadly about push rods as vehicles for mechanoreceptor stimulation, it is of interest that they are located exclusively in bill skin of the platypus and in nose skin of the echidna. Being at the front end of the animal, perhaps they subserve some kind of teletactile function, that is, detection of disturbances in the water or in the soil created by prey at some distance ahead of the animal. Vertebrates have evolved a number of ways of stimulating, at a point, a population of receptors with both static and dynamic properties. An example that comes to mind is the vibrissae of mammals. Here it is known that the various afferents supplying each vibrissa project to the same area of cerebral cortex to form identifiable ‘barrels’ (Miller et al. 2001). It suggests that central processing of tactile information coming from these structures requires the presence of both static and dynamic components of the stimulus and the processing is done by neurones lying in close proximity to one another. For push rods, the presence of a mobile column of epidermal cells that is able to excite paciniform and Merkel cell receptor types at the same time, provides the opportunity for similar central processing of static and dynamic stimuli. It remains to determine why it is necessary to process the information in this way. Presumably the nature of the sensory experience evoked by stimulation of a push rod is dependent on the simultaneous presentation of static and dynamic features of the stimulus. Proc. Linn. Soc. N.S.W., 125, 2004 REFERENCES Andres, K. and Von During, M. (1984). The platypus bill. A structural and functional model of a pattern- like arrangement of different cutaneous sensory receptors. In Sensory Receptor Mechanisms (ed. W. Hamann and A. Iggo). World Scientific, Singapore. Andres, K.H., Von During, M., Iggo, A. and Proske, U. (1991). The anatomy and fine structure of the echidna Tachyglossus aculeatus snout with respect to its different trigeminal sensory receptors including the electroreceptors. Anatomy and Embryology 184, 371-393. Catania, K.C. (1995). Structure and innervation of the sensory organs on the snout of the star-nosed mole. Journal of Comparative Neurology 351, 536-548. Catania, K.C. (2000). Epidermal sensory organs of moles, shrew moles, and desmans: a study of the family talpidae with comments on the function and evolution of Eimer’s organ. Brain, Behavior & Evolution 56, 146-174. Catania, K.C. and Kaas, J.H. (1995). Organization of the somatosensory cortex of the star-nosed mole. Journal of Comparative Neurology 351, 549- 567. Iggo, A., Gregory, J.E. and Proske, U. (1992). The central projection of electrosensory information in the platypus. Journal of Physiology 447, 449-465. Iggo, A., Gregory, J.-E. and Proske, U. (1996). Studies of mechanoreceptors in skin of the snout of the echidna Tachyglossus aculeatus. Somatosensensory and Motor Research 13, 129- 138. Iggo, A., McIntyre, A.K. and Proske, U. (1985). Responses of mechanoreceptors and thermoreceptors in skin of the snout of the echidna Tachyglossus aculeatus. Proceedings of the Royal Society of London - Series B: Biological Sciences 223, 261-277. Manger, P.R., Calford, M.B. and Pettigrew, J.D. (1996). Properties of electrosensory neurones in the cortex of the platypus (Omnithorhynchus anatinus): implications for processing of electrosensory stimuli. Proceedings of the Royal Society of London - Series B: Biological Sciences 263, 611-617. Manger, P.R., Keast, J.R., Pettigrew, J.D. and Troutt, L. (1998). Distribution and putative function of autonomic nerve fibres in the bill skin of the platypus (Ormithorhynchus anatinus). Philosophical Transactions of the Royal Society of London - Series B: Biological Sciences 353, 1159-1170. Manger, P.R. and Pettigrew, J.D. (1996). Ultrastructure, number, distribution and innervation of electroreceptors and mechanoreceptors in the bill skin of the platypus, Ornithorhynchus anatinus. Brain, Behavior & Evolution 48, 27- 54. 825 ROLE OF PUSH RODS IN MONOTREMES Miller, B., Blake, N.M.J. and Woolsey, T.A. (2001). Barrel cortex: structural organisation, development and early plasticity. In Plasticity of Adult Barrel Cortex (Ed. M. Kossut), pp. 3-45. (F.P. Graham, Johnson City TN, USA). Pettigrew, J.D., Manger, P.R. and Fine, S.L. (1998). The sensory world of the platypus. Philosophical Transactions of the Royal Society of London - Series B: Biological Sciences 353, 1199-1210. Proske, U. (1997). Echidna on the nose. Nature Australia 25, 58-63. Proske, U. and Gregory, J.E. (2003). Electrolocation in the platypus — some speculations. Comparative Biochemistry and Physiology 136, 821-825. 326 Proc. Linn. Soc. N.S.W., 125, 2004 Obituary MERVYN EDWARD GRIFFITHS 1914-2003 Mervyn Griffiths (who always preferred to be called “Merv’’) was born in Sydney on 8" July 1914, was educated at North Sydney Boys’ High School and obtained his Bachelor Degree in Zoology with first Class Honours in 1937, followed by his Master of Science in 1938 at Sydney University Merv first began publishing in the scientific literature in 1936 with a paper on The colour changes in batoid fishes in the Society’s Proceedings and contributed six further papers to this journal between 1936 and 1942. After completing his Master studies, Merv was awarded the Travelling Research Scholarship for the 1851 Exhibition, which took him to McGill University in Montreal, Harvard University and the National Institute for Medical Research in London. From this work he produced 10 papers, including research on diabetes mellitus and the secretory functions of the pituitary. In 1941 Merv returned to the University of Sydney where he continued his work on diabetes in the Department of Medicine, where he was a Linnean Macleay Fellow in Physiology in 1941. The fellowship was renewed in 1942, but only a few months into his second fellowship year he joined the Royal Australian Air Force. Merv was in the Empire Air Training Scheme during 1943 in Edmonton, Canada and became a Pilot Officer in 1944. He held the post of Commanding Officer of the 3rd Malaria Control Unit in Darwin from 1944-45 and left the RAAF in February 1946 at the age of 31, continuing his involvement in diabetes research as a Zoologist in the Institute of Anatomy in Canberra. He became the Senior Biochemist at the Institute in 1949, publishing his work on the biochemistry of diabetes through to 1957, when he returned to zoology, joining the C.S.I.R.O. Wildlife Survey Section [which later became the Division of Wildlife Research] as a Senior Research Officer in June 1957. In 1959 he was awarded his Doctor of Science Degree by Sydney University for his thesis entitled The Relationship of the Pituitary Gland to Experimental Diabetes and the Action of Insulin. Although his initial published works at the C.S.I.R.O. were concerned with rabbits, Merv became interested in the biology of marsupials, particularly macropods. In 1965 he published his first paper on the biology of the group for which his research is best known, and which became his consuming passion - the Monotremes. His monograph on the Echidnas was published in 1968. During his time at the Division of Wildlife Research, Merv was the scientific director of two films, The Echidna and the Comparative Biology of Lactation, both of which won awards. Merv retired in October 1975 from the C.S.ILR.O. Division of Wildlife Research as Senior Principal Research Officer, but not from the field of wildlife research. His interests broadened, although the monotremes remained his prime focus. His classic work The Biology of the Monotremes in 1978 pulled together all of the disparate research carried out on the group to that time. Merv was variously honoured by scientific societies, including being awarded the Peter Aitken Medal by the South Australian Museum in 1988 and becoming a Fellow of the Royal Zoological Society of N.S.W. in 1991. In his “retirement” Merv researched and published widely in a number of fields, where his own work and collaboration with colleagues and friends resulted in a further 33 publications to add to his pre-retirement total of forty-three. Merv Griffiths died on 6" May 2003. The above summary of his academic life cannot adequately describe his contribution to biological science in Australia. Throughout his academic career Merv remained a great “generalist” in a world of increasing numbers of scientific “specialists”. His encouragement, generous advice, support and sometimes cajoling, are deeply appreciated by many of these specialists. 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Aid} eeeaes or 3 vane: b ainearee aie es | ef ioe pag ieee aia Sir J ta Se aS Lae. eta obipioig tn 16r00 dad bsinaven ioe CCE n=! % | phn 2 = ak “pettiins: dizonyiaiddor: wieriny: qontiye bal ee ay SO eit, aa As anor Prag inis = ae oe 2 aftvedi Yo alton aft en eons OR. aloteoieberaliden lepine 5 ES a VOR tees iw banc ao hy . & Ww ae ‘<4 BOOK REVIEW Admiral Doenitz’s Legacy Paul Adam School of Biological, Earth and Environmental Sciences, University of New South Wales, NSW 2052 Murray, D.R. 2003 Seeds of concern. The genetic manipulation of plants. UNSW Press.156pp. ISBN 0 86840 460 8. $34.95. Over the last decade there has been increasing public concern about the development and use of genetically modified organisms. This has been manifest in media coverage involving ‘shock-horror’ headlines such as “Frankenfoods’, litigation, product boycotts, increased sales of ‘organic’ products, illegal destruction of genetically modified crops and, recently, the rejection by Zambia of food aid which may have contained genetically modified grains. Opposition has been global, but at its most intense in Europe. What is the basis for the concern and is it justified? Some of the concerns are clearly unnecessary, reflecting an unfortunate lack of basic scientific knowledge amongst the media and general public. Alarmist stories about how eating genetically modified organisms involves consuming DNA, as if non- modified organisms lack DNA, do not add to the credibility of journalists or their editors. For products which are extracted, highly refined and purified, whether their origin is from modified or unmodified organisms is irrelevant in terms of the end qualities and properties. Other issues have more substance, but at least in Europe, the GM debate is only a symptom of a much broader concern about the nature of modern agriculture. Public confidence has been disturbed by the outbreaks of both BSE and Foot and Mouth disease, which are seen as components of a broader malaise. Neither disease of course has anything to do with genetic modification, although particularly in the case of BSE, significant portions of the media and the public believe it does. To understand the origins of the perceived malaise it is necessary to go back to the First World War. For the first time submarines proved to be an important weapon, and convoys of food supplies were disrupted. Between the wars submarine technologies were substantially developed. Agriculture was little changed, indeed for much of the period European agriculture was in decline, as cheap imports from north America and the southern hemisphere satisfied the market. During the Second World War U-boats maintained a blockade which almost brought Britain to defeat. The post war response was — ‘never again’, and the still prevailing policy of self sufficiency was developed. Governments plan to win the last war, so the fact that the weapons of mass destruction which brought the Second World War to an end changed the nature of any future global war, did not influence policy development. From some perspectives the self sufficiency policy could be judged a success. Who in 1945 would have anticipated the vast European Union surpluses, or that in 2003 British livestock-feed grains would be exported to Australia? Nevertheless it is the cost of this success which is now being questioned. The drive for increased production and efficiency was powered by subsidies, both direct and, indirect, and the developing agribusiness companies. Synthetic pesticides and herbicides (such as DDT and MCPA) were first used on a large scale towards the end of the war, and in the immediate post war years usage burgeoned, being proclaimed as an example of the new scientific approach to farming. There was little external scrutiny of government programs and the administrative bureaucracies were captive to their clients — farmers and agribusiness. The first expressions of concern surfaced with the publication of Rachel Carson’s Silent Spring (1962). Both the publication and its author were subject to sustained attack by both government and agribusiness, but the basic thesis was increasingly supported by independent evidence. The publication of Silent Spring was one of the key events leading to the modern environmental movement, resulted in the banning in the west, if not globally, of some pesticides and the institution of greater scrutiny of new chemicals. Nevertheless these were only minor hiccups on the way to industrialization of agriculture. Other changes included, in northern Europe, the decline of mixed farming in favour of specialization, the loss of woodlands, hedgerows and BOOK REVIEW - SEEDS OF CONCERN wetlands, the loss of genetic diversity amongst crops as many local varieties were replaced by a few new cultivars, and increased use of nitrogen fertilizers resulting in greener, but floristically simpler, basically Lolium monocultures, pastures. New crops came to prominence, most notably oilseed rape (known in more sensitive nations such as Australia as canola) converting the landscapes of Constable to ones more akin to those of van Gogh. Lifestock production was increased through adoption of so-called factory farming techniques, changing, for example, chicken from a luxury to a convenience food but raising widespread community concerns about animal welfare and creating substantial environmental problems associated with effluent management. The changing face of the countryside, increasing awareness of the impacts on biodiversity, and concerns about potential impacts on human health has led to an upsurge of public disquiet (Shoard 1980, Harvey 1997, 2001, Humphrys 2001, Green 2002), but reform of the European agricultural system, although debated for several decades has been slow to eventuate. The idealized countryside of an urban population is often a bucolic dream, the product of a Romantic imagination, and ignores the earlier extensive changes wrought by the Agricultural Revolution and enclosures (Fox and Butlin 1979), but the issues raised by commentators such as Harvey (1997) are nevertheless well documented and cannot lightly be dismissed. However, the public concern over the consequences of agricultural policy runs contrary to the public expectation, also developed since 1945, of a never ending supply of cheap food. This expectation is well summarized by Gummer (2001) (John Gummer is a former UK Minister for Agriculture and Secretary of State for the Environment). “Many in the rich world, who do not blanch at forking out £30,000 for a more fashionable motor car, will refuse to expend threepence more on a fresher lettuce or a tastier loaf of bread. Food, which ought to rank highest among our spending priorities, has been relegated to the rank of necessity, and in this advanced civilization of ours, only luxuries deserve to be prized. We take necessaries as our right and expect them to be delivered at a discount. So it is that food prices demand a smaller and smaller proportion of a prosperous household’s income and take less time than ever for the average worker to earn. What is more, now that packaging, distribution and preparation are necessary on-costs, the basic food content of what we buy represents an even smaller proportion of what we pay. With one in three meals eaten out of the home and most of the rest to a growing 330 extent pre-prepared, that proportion will continue to fall”. In demanding cheap food the public neglects to take into consideration the considerable subsidies paid out of the taxpayers’ pocket to farmers by governments. The trend to cheap food has also been assisted by the growth of supermarket chains. For many foodstuffs, prices to farmers are determined by a global oligopoly of retailers. With the availability of cheap airfreight a further consequence of the growth of the supermarket chains is the abolition of seasons, with the same range of produce being available globally year round. This means that the shelves of European supermarkets may be replete with sugar peas from Zambia, green beans from Kenya, salad greens from Tanzania and southern Africa, even brussel sprouts from Australia. When the costs of freight and packaging are considered the return to farmers must be very small, and does not take into account environmental degradation and disruption of traditional agricultural economies. Additionally agriculture in developing countries is adversely affected by dumping of surpluses from elsewhere — tomato producers in West Africa for example cannot compete even in their local market against heavily subsidized Italian canned tomatoes (Bradshaw 2003). There is little doubt that if global agriculture had remained as it was in 1945 it would have been impossible to support the current human population of about six billion. Currently global food production is capable of providing adequate nutrition for the world population (Waterlow et al. 1998), famine and malnutrition are the consequences of failures of political systems, not of agriculture. What is less certain is whether, without extensive adoption of new molecular techniques, it will be possible to feed the predicted human population in 2050 of eight billion (Waterlow et al. 1998), particularly given the added impacts of global warming. The increased production is due in part to an increase in the land area devoted to agriculture but is largely the consequence of new technologies. The development of agricultural chemicals, mechanization and application of conventional plant and animal breeding can rightly be regarded as scientific and engineering triumphs. Nevertheless broader questions of ecological sustainability were not part of the agenda until recently, and the true costs of food production have rarely been calculated. How is this discussion relevant to any debate about use of genetically modified organisms in Australia? Firstly, despite talk about free trade and level Proc. Linn. Soc. N.S.W., 125, 2004 P. ADAM playing fields, world agricultural markets are still heavily influenced by government subsidies and policy intervention. Whether the original motivation for these government programs can still be justified (and in the case of the much maligned European Common Agricultural Policy, maintenance of the social structure of rural communities was as important as ensuring self- sufficiency) it is unlikely that there is political will for change. Australia’s agricultural future, both in terms of access to overseas markets and ability to control imports, will be very much determined by what happens in Washington and Brussels. The US is currently seeking action by the World Trade Organisation to require the European Union to lift its restrictions on genetically modified food (Sanger 2003). The outcome of these proceedings will have global implications, determining for example whether Australia could similarly impose controls on imports of genetically modified organisms. President Bush has argued that Europe’s approach has discouraged use of genetically modified foods in the third World and this contributed to continuing famine in Africa (Sanger 2003). Most would argue that African famines have a number of causes, and that absence of genetically modified crops is not one of them. There will be continuing pressure for increased efficiency (as measured in production of cheap food) and this will have social and economic consequences. Secondly agribusiness is global, and increasingly vertically integrated, so that much plant breeding and development of genetically modified plants will be in the hands of a few multinationals, with the results imported into Australia. The growing hegemony of a few agribusiness company has considerable implications beyond the issue of genetic modification. Any choices that farmers might have in terms of crop varieties or chemicals are becoming increasingly illusory. President Eisenhower famously warned of the influence of the military industrial complex — his words would equally apply to agribusiness. To date, the introduction of genetically modified crops in Australia has been, as it also has been in Europe, a public relations disaster for agribusiness. However, information about the nature of the genetically modified crops has not been readily accessible to the public. David Murray’s ‘Seeds of Concern’ attempts to fill the gap, with the objective of promoting informed debate. The author is a distinguished plant scientist so his critique cannot be dismissed by the more enthusiastic proponents of the technology as well intentional but misinformed. However, while the book is aimed, in part, at the intelligent lay person, the assumed level of chemical/biochemical knowledge is Proc. Linn. Soc. N.S.W., 125, 2004 high and may well deter the intended audience. The technology to achieve genetic manipulation exists, and the genie cannot be put back in the bottle. Murray recognizes this and discusses possibilities for using the technologies which could potentially be of considerable benefit to humankind. Unfortunately the benefits of the majority of genetically modified plants released to date accrue to agribusiness and farmers rather than the consumer. Genetic manipulation is possible in the whole range of organisms. Modification of microorganisms for industrial processes has not attracted much public attention, modification of domestic animals (including farmed fish) is still largely at the trial stage, but genetically modified plants are in the landscape and market place and very much in the public eye. Murray provides a very succinct introduction to plant cell biology and the techniques of genetic manipulation. In commercially released genetically modified plants the most frequent changes are the introduction of herbicide resistance or of genes which express insecticidal compounds. Murray explains the underlying basis for both changes, but also discusses actual and potential drawbacks with the use of plants modified in these ways in the field. It is far from clear that in the long term these approaches will have benefit. The number of potential applications of genetic modification which have been touted in the media is very large. Murray explains how in some cases, such as lowering caffeine levels in coffee or preventing expression of polyphenol oxidases, the proponents are ignorant of the biological function of these compounds or of consumer requirements. The yield costs of genetic modification are often, as Murray points out, given little consideration. What Murray regards as failings of current patenting regimes and of Australia’s Plant Breeders Rights legislation are discussed at some length. I would support the critique although recognizing that commercial interests would be expected to take a different view. These difficulties also arise with conventional plant breeding, but, with the heavy investment in genetic modification are likely to be more apparent as companies seek to protect what they regard as their intellectual property. The regulatory framework for release of genetically modified organisms currently applying in Australia is discussed in some detail. While this regime is important, Murray does not raise the issue of general lack of regulation of many agricultural activities. Farmers world wide argue that they are over regulated, and arguments in favour of less regulation abound (see Ridley 2001, Pennington 2001, and the continuing opposition by farmers in Eastern Australia to control of land clearing). While it is the case that some aspects 331 BOOK REVIEW - SEEDS OF CONCERN of agriculture are heavily regulated, key decisions are left to farmers. The cover of “Seeds of Concern’ shows a field of canola (rape); the decision of European farmers to adopt broad acre rape cultivation had a profound visual and ecological impact, but was not one in which the broader community participated. The change in the northern hemisphere from spring to winter cereal cultivation has had major effects on many bird species, but again was a decision of farmers and agribusiness. In the Australian context changes from pastoralism to cropping, or the spread of cotton growing (both changes which may be facilitated by the development of genetically modified plants) will have social, ecological and environmental consequences, but the decisions will be made by landholders with few avenues for external scrutiny, let alone any requirement for approval. Even if the specific concerns of the Gene Technology Regulator are met, the broader questions raised by agricultural change will remain unaddressed. These are clearly issues beyond those which Dr. Murray set out to address, but they do need to be placed on the policy agenda. Dr. Murray also manages within the compass of his admirably concise book to mount a defence of Mendel against claims that his results might have been ‘polished’, express skepticism about proposals to clone thylacines, and have a few swipes against the pontifications of the great and the good. Despite in a few places being ill served in matters of layout by the publisher, this is an important contribution to the debate about genetically manipulated plants and of the future of agriculture. There are indeed ‘seeds of concern’, perhaps leading to seeds of doubt. REFERENCES Bradshaw, S. (2003) Unfair meal gives taste of global trade pitfalls. Broadcast on ABC ‘Landline’ 13 April 2003. Transcript at http://www.abc.net.au/ landline/stories/s828402.htm Carson, R. (1962) Silent spring. (Houghton Miflin, Boston). Fox, H.S.A. & Butlin, R.A. (Eds) (1979) Change in the countryside. (Institute Of British Geographers, London). Green, B. (2002) The farmed landscape: the ecology and conservation of diversity. In Remaking the landscape. The changing face of Britain. Ed. J. Jenkins pp.183-210. (Profile Books, London). Gummer, J. (2001) Farming and the CAP. In A countryside for all. The future of rural Britain. (Ed. M. Sissons). pp.57-68. (Vintage, London). Harvey, G. (1997) The killing of the countryside. (Jonathan Cape, London). Harvey, G. (2001) Reinventing agriculture. In A countryside for all. The future of rural Britain. (Ed. M. Sissons). pp./7-88. (Vintage, London). Humphrys, J. (2001) The great food gamble. (Hodder and Stoughton, London). Pennington, M. (2001) Deregulating the land: an alternative route to urban and rural regeneration. In A countryside for all. The future of rural Britain. (Ed. M. Sissons). pp.35-47. (Vintage, London). Ridley, M. (2001) Denationalising the land. In A countryside for all. The future of rural Britain. (Ed. M. Sissons). pp.25-33. (Vintage, London). Sanger, D. (2003) Bush blames European rules on GM food for hunger in Africa. The Sydney Morning Herald, May 23. p.12. Shoard, M. (1980) The theft of the countryside. (Temple Smith, London). Waterlow, J.C., Armstrong, D.G., Fowden, L. & Riley, R. (Eds) (1998) Feeding a world population of more than eight billion people. A Challenge to science. (Oxford University Press, Oxford). Proc. Linn. Soc. N.S.W., 125, 2004 BOOK REVIEW M.L. Augee Wellington Caves Fossil Studies Center, 89 Caves Road, Wellington NSW 2820 Duyker, Edward. 2003. Citizen Labillardiére: a naturalist’s life in revolution and exploration (1755-1834) Edward Duyker. The Miegunyah Press, Melbourne 2003. $59.95 This book is a splendid example of the need for academic presses in Australia; Miegunyah Press is an imprint of Melbourne University Publishing. It is a scholarly, extremely well documented work covering a poorly known but important aspect of Australia’s early history. As such it cannot be considered an “easy read” nor a likely undertaking for a commercial publisher. It is however a goldmine of information and will be a valuable reference source for anyone seeking information on the roots of natural history study in Australia or the contributions of French scientists to such study. The use of this book as a reference is greatly facilitated by three complete indexes (botanical, zoological and general) and a fully detailed bibliography. The amount of research into resources in several languages required to produce this book is amazing. Such detail insures its use for many years to come. “Citizen Labillardiére” can be read simply as a biography of a very interesting man who lived through very interesting times. He stands astride the “troubles” in France as the country shifted from the Ancien Régime to the National Assembly and constitutional monarchy; to a republic overtaken by the “Reign of Terror’; to an Empire (Napoleon); to the restoration of the monarchy; and then to the final overthrow of the Bourbons. The “citizen” in the book title indicates Labillardiére’s general political leaning, but it is also the story of a man of science trying for the middle road in turbulent politics. It is therefore a story of survival and adventure. Much of the adventure comes from travel, often arduous, over much of the globe. After graduating in medicine (1772), Labillardiére immediately begins his life of travel with ajourney to England. There he establishes connections with Joseph Banks and other British scientists that will serve him well in the future. He travels through the Alps and Lebanon, always collecting botanical specimens and possessing a remarkable enthusiasm for climbing mountains. Then comes his big break; he is appointed as a naturalist on the expedition to be lead by Bruny d’ Entrecasteaux in search of the missing La Perouse (last seen by the British at Botany Bay on 10 March 1788). This small fleet, consisting of the Recherche and the Espérance, left France on 28 September 1791. After stops at Tenerife and the Cape of Good Hope, where Labillardi¢re made many collections, they landed in Van Diemen’s Land, at Recherche Bay, on 20 April 1792. There Labillardiére collected 5,000 specimens in five weeks. Sailing north in search of La Perouse, the expedition went to New Caledonia, New Guinea and the Solomons before returning to Recherche Bay. At that time Labillardiére records positive encounters with the native people; indeed he seems to have been a sympathetic and astute observer of the Tasmanian natives. Leaving Van Diemen’s Land, the expedition went to Tonga, passing close by the tip of New Zealand, and to New Caledonia. A return to Van Diemen’s Land, with the great likelihood of discovering that it was not part of the mainland and discovering the strait later to be discovered by Bass, was not to be, and d’Entrecasteaux died at sea on 19 July 1793. Already falling apart with rifts between royalists and republicans, the expedition died when, upon reaching Java, the Dutch were found to be at war with France. The Dutch imprisoned Labillardiére and others, although not the royalists who collaborated with them. Labillardiére was finally released on 29 March 1795 and, after spending six months at Ile-de- France enroute, was back in Paris in March 1796. Meanwhile most of his collection of botanical specimens had been taken to England and were claimed by the royalist French court in exile there. Banks however used his strong influence and the collection was returned to Labillardiére. There was an odd interlude in the life of Jacques-Julien Labillardiére when, in the summer of 1796, he went to Italy as part of a commission to oversee seizure of Italian art and other treasures taken as tribute after General Napoleon Bonaparte conquered that country. In 1800 Labillardiére was made a full member of the Académie des sciences. Besides the political turmoils of his time, Labillardiére also straddled a revolution in biological science, during the change from strict application of Linnean principles (which saw taxonomy as a rigid BOOK REVIEW - CITIZEN LABILLARDIERE set of categories, determined in the case of plants by the structure of the sexual parts of flowers) to a more natural approach (systematics including as much evidence as possible, especially physiological and morphological). Labillardiére’s “Sertum austro- caledonicum” published in 1824 abandoned the strict Linnean approach. It should be noted, however, that well before that time zoologists (such as the towering figure of Baron Cuvier) had widely accepted the natural system. Not unlike today when about the only resistance to modern systematics, such as cladistics, comes from botanical taxonomists, many of whom one suspects strongly regret that upstart Antoine-Laurent de Jussieu who started tinkering with the Linnean system in 1789. Labillardiére died 8 January 1834 aged 79. He had a truly adventurous life and made a significant contribution to the advancement of science, although the claim by Duyker that he was “one of the founders of botany, zoology and ethnography in Australia” is overly generous. Labillardiére was basically a collector with a sometimes off-hand approach to record keeping. Nonetheless, he did record many “firsts” and many names he gave to plants are valid today. 334 Proc. Linn. Soc. N.S.W., 125, 2004 INSTRUCTIONS FOR AUTHORS (this is an abbreviated form — the full instructions can be obtained from our web site or from the Secretary) 1. The Proceedings of the Linnean Society of New South Wales publishes original research papers dealing with any topic of natural science, particularly biological and earth sciences. 2. Manuscripts should be submitted to the Editor (M.L. Augee, PO Box 82, Kingsford NSW 2032). All manuscripts are sent to at least two referees and in the first instance three hard copies, including all figures and tables, must be supplied. Text must be set at one and a half or double spacing. 3. The final version, incorporating referees’ and editor’s comments, must be supplied on floppy disc or CD in WORD for PC format (Mac discs will not be accepted). Photographs must be supplied as black and white prints or as .TIF files (Jpeg is not acceptable). Figures other than photographs must be in hardcopy, EXCEL or WORD files. The text file must contain absolutely no auto-formatting or track changes. Tables and/or figures must be separate from the text file. 4. References are cited in the text by the authors’ last name and year of publication (Smith 1987). For three of more authors the citation is (Smith et al. 1988; Smith and Jones 2000). Notice that commas are not used between the authors’ names and the year. The format for the reference list is: Journal articles: Smith, B.S. (1987). A tale of extinction. Journal of Paleontological Fiction 23, 35-78. Smith, B.S., Wesson, R.I. and Luger, W.K. (1988). Levels of oxygen in the blood of dead Ringtail Possums. Australian Journal of Sleep 230, 23-53. Chapters or papers within an edited work: Ralp, P.H. (2001). The use of ethanol in field studies. In ‘Field techniques’ (Eds. K. Thurstle and P.J. Green) pp. 34-41. (Northwood Press, Sydney). Books: Young, V.H. (1998). “The story of the wombat’. (Wallaby Press, Brisbane). 5. An abstract of no more than 200 words is required. Sections in the body of the paper usually include: INTRODUCTION, MATERIALS AND METHODS, RESULTS, DISCUSSION, ACKNOWLEDGEMENTS and REFERENCES. Some topics, especially taxonomic, may require variation. 6. Subheadings within the above sections should be in the form: Bold heading set against left margin This is the form for the first level headings and the first line of text underneath is indented Underlined heading set against left margin This is the next level, and again the first line of text underneath is indented. Further subheadings should be avoided. Italics are not to be used for headings but are reserved for genus and species names. 7. Up to 10 KEYWORDS are required. These are often used in computer search engines, so the more specific the terms the better. ‘Australian’ for example is useless. Please put in alphabetical order. 8. Paragraphs are to be set off by a tab indentation without skipping a line. Do not auto-format the first line (i.e. by using the “first line” command in WORD). All auto-formatting can be fatal when transferring a manuscript into the publisher platform. 9. Details of setting up the manuscript: Use 12 point Times New Roman font. Do not justify Margins should be: 3 cm top, 2.5 cm bottom, 3 cm left and 2.5 cm right. This is the area available for text; headers and footers are outside these margins. 10. Figures can be line drawings, photographs or computer-generated graphics. No figures will be accepted INSTRUCTIONS FOR AUTHORS larger than 15.5 X 23 cm. Width of lines and sizes of letters in figures must be large enough to allow reduction to half page size. If a scale is required it must be presented as a bar within the figure. It is the editor’s prerogative to reduce or enlarge figures as necessary and statements such as “natural size” or “4X” in the legend are unacceptable. While there is no objection to full page size figures, it is journal policy to have the legend on the same page whenever possible and figures should not be so large as to exclude the legend. Figure legends should be placed together on a separate page at the end of the manuscript. 11. Tables must be provided on separate pages at the end of the manuscript and NOT placed within the text. It is essential that table legends are not set within the table but are supplied separately as with figure legends. It may be necessary to reduce or enlarge tables but the legends must remain in the same font as the text (10 point Times New Roman in the final form). While the draft manuscript text is expected to be in 12 point type, it may be necessary to use smaller font size for large tables. Do not use vertical lines in tables unless absolutely necessary to demark data columns. Keep horizontal lines to a minimum. Do not place the table in a box border. WORD or EXCEL tables are acceptable. 12. Details of punctuation, scientific nomenclature, etc. are to be found in the complete instructions. 336 Proc. Linn. Soc. N.S.W., 125, 2004 Part 2 — monotreme papers 217 227 235 243 259 273 PATE 279 287 301 319 ‘Grant, T.R. Captures, capture mortality, age and sex ratios of platypuses, Ornithorhynchus anatinus, during studies over 30 years in the upper Shoalhaven River in New South Wales. Grant, T.R., Griffiths, M. and Temple-Smith, P.D. Breeding in a free-ranging population of platypuses, Ornithorhynchus anatinus, in the upper Shoalhaven River, New South Wales - a 27 year study. Grant, T.R. Depth and substrate selection by platypuses, Ornithorhynchus anatinus, in the lower Hastings River, New South Wales. Lunney, D., Grant, T. and Matthews, A. Distribution of the platypus in the Bellinger catchment from community knowledge and field survey and its relationship to river disturbance. ; Grant, T.R., Lowry, M.B., Pease, B., Walford, T.R. and Graham, K. Reducing the by-catch of platypuses (Ornithorhynchus anatinus) in commercial and recreational fishing gear in New South Wales. Bethge, P., Munks, S., Otley, H. and Nicol, S. Platypus burrow temperatures at a subalpine Tasmanian lake. Higgins, D.P. Ultrasonography of the reproductive tract of the short-beaked echidna (Tachyglossus aculeatus). Higgins, D.P., Tobias, G. and Stone, G.M. Excretion profiles of some reproductive steroids in the faeces of captive female short-beaked echidna (Tachyglossus aculeatus) and long-beaked echidna (Zaglossus sp.). Hassiotis, M., Paxinos, G. and Ashwell, K.W.S. Anatomy of the central nervous system of the Australian echidna. Rowe, M.J., Mahns, D.A. and Sahai, V. Monotreme tactile mechanisms: from sensory nerves to cerebral cortex. - Proske, U. and Gregory, J.E. The role of push rods in platypus and echidna — some speculations. Part 3 — obituary, book reviews, instructions to authors 327 Obituary: Mervyn Edward Griffiths 1914-2003. 329 Adam, P. Admiral Doenitz’s legacy. A review of the book “Seeds of concern. The genetic manipulation of plants” by D.R. Murray (2003). 333 Augee, M.L. A review of the book “Citizen Labillardiére: a naturalist’s life in revolution and exploration (1755- 335 1834)” by Edward Duyker. Instructions for authors. PROCEEDINGS OF THE LINNEAN SOCIETY OF N.S.W. VOLUME 125 Issued 20 February 2004 CONTENTS Part 1 — general contributions { 43 Dif, 67 73 oF 110 115 141 aete) Moulds, T. Review of Australian cave guano ecosystems with a checklist of guano invertebrates. Young, GC. Nd BY=\Voyalt-laM o)e-Volanyatarele= lore R-Vatalcoelic= =) <0] 1M (olf-(erele -1ana Mica) Micelaamlai-Msi(e)\>0) mm lal= Mi al=1al (ere) 0)(e Mex=1alt| ol=\e| >a B)(e/¢[=)(6)0)/0r-m ( @1a}l fe) elefel- bm Miiiave)e)(e)anre)se)ar-) Micelaamcveleltal=r-t)(-108) Australia and Lord Howe Island. Weaver, H.J. and Aberton, J.G. A survey of ectoparasite species on small mammals during autumn and winter at Anglesea, Victoria. Allen, S., Marsh, H. and Hodgson, A. Occurrence and conservation of the dugong (Sirenia: Dugongidae) in New South Wales. 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