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Papers in Honour of Ken Aplin edited by Julien Louys, Sue O'Connor and Kristofer M. Helgen
Helgen, Kristofer M., Julien Louys, and Sue O'Connor. 2020. The lives of creatures obscure, misunderstood, and wonderful: a volume in honour of Ken Aplin 1958-2019 ..........................
Armstrong, Kyle N., Ken Aplin, and Masaharu Motokawa. 2020. A new species of extinct False Vampire Bat (Megadermatidae: Macroderma) from the Kimberley Region of Western Australia 12s lere ebur gait enin asked i areae eei n
Cramb, Jonathan, Scott A. Hocknull, and Gilbert J. Price. 2020. Fossil Uromys (Rodentia: Murinae) from central Queensland, with a description of a new Middle PleistoGerigispecies wi. tio sem Roo ml ende desde e tee SUP Shall eph ad
Price, Gilbert J., Jonathan Cramb, Julien Louys, Kenny J. Travouillon, Eleanor M. A. Pease, Yue-xing Feng, Jian-xin Zhao, and Douglas Irvin. 2020. Late Quaternary fossil vertebrates of the Broken River karst area, northern Queensland, Australia ........................
Theden-Ringl, Fenja, Geoffrey S. Hope, Kathleen P. Hislop, and Benedict J. Keaney. 2020. Characterizing environmental change and species’ histories from stratified faunal records in southeastern Australia: a regional review and a case study for the early to middle Holocene ...................sssssssssssssssee ee ee ees
Brockwell, Sally, and Ken Aplin. 2020. Fauna on the floodplains: late Holocene culture and landscape on the sub-coastal plains of northern Australia ...................sssssssssss
Hawkins, Stuart, Fayeza Shasliz Arumdhati, Mirani Litster, Tse Siang Lim, Gina Basile, Mathieu Leclerc, Christian Reepmeyer, Tim Ryan Maloney, Clara Boulanger, Julien Louys, Mahirta, Geoff Clark, Gendro Keling, Richard C. Willan, Pratiwi Yuwono, and Sue O'Connor. 2020. Metal-Age maritime culture at Jareng Bori rockshelter, Pantar Island, eastern Indonesia .................sssssssssssenne ne
Frankham, Greta J., Linda E. Neaves, and Mark D. B. Eldridge. 2020. Genetic relationships of Long-nosed Potoroos Potorous tridactylus (Kerr, 1792) from the Bass Strait Islands, with notes on the subspecies Potorous tridactylus beporint Courtney- d OL hrana dn d cert dace iedyma pee nace ias pega inttr f aer ree p
Rowe, Kevin C., Helena A. Soini, Karen M. C. Rowe, Mark Adams, and Milos V. Novotny. 2020. Odorants differentiate Australian Rattus with increased Complexity in SYMpatry MRRNEEEEMEEMMMMMMMMMMM
Louys, Julien, Michael B. Herrera, Vicki A. Thomson, Andrew S. Wiewel, Stephen C. Donnellan, Sue O'Connor, and Ken Aplin. 2020. Expanding population edge craniometrics and genetics provide insights into dispersal of commensal rats through Nitsa Tengsarae Indonesia «Es xo d t s Rum ea es d eM eph nte uh se
Breed, William G., Chris M. Leigh, and Eleanor J. Peirce. 2020. Reproductive biology of the mice and rats (family Muridae) in New Guinea—diversity and evolution .........................
Suzuki, Hitoshi. 2020. Evolutionary history of the subgenus Mus in Eurasia with special emphasis on the House Mouse Mus musculus eenen
Richards, Stephen J., and Stephen C. Donnellan. 2020. Litoria aplini sp. nov., a new species of treefrog (Pelodryadidae) from Papua New Guinea ...............ssssssssssse
Records of the Australian Museum volume 72, issue no. 5 25 November 2020
AUSTRALIAN MUSEUM
Records of the Australian Museum (2020) vol. 72, issue no. 5, pp. 149—160 https://doi.org/10.3853/j.2201-4349.72.2020.1734
Records of the Australian Museum
a peer-reviewed open-access journal
published by the Australian Museum, Sydney communicating knowledge derived from our collections ISSN 0067-1975 (print), 2201-4349 (online)
The Lives of Creatures Obscure, Misunderstood, and Wonderful: A Volume in Honour of Ken Aplin 1958-2019
KRISTOFER M. HELGEN!?®, JULIEN Louys?®, Suge O’CONNOR?*®
! Australian Museum Research Institute, Australian Museum, 1 William Street, Sydney, Australia
? Australian Research Council Centre of Excellence for Biodiversity and Heritage, Canberra, Australia
? Australian Research Centre for Human Evolution, Environmental Futures Research Institute, Griffith University, Nathan Qld 4111, Australia
^ School of Culture, History, and Languages, College of Asia and the Pacific, The Australian National University, Canberra, Australia
He was always a modest man, but Ken was a genius and the toughest man we knew. He was also extraordinarily generous of spirit. The way he gave of himself, his time, and his hard-won stores of knowledge, was legendary amongst his friends and colleagues. We admired him and we loved him. Ken was a world-renowned comparative anatomist, vertebrate systematist, palaeontologist, and zooarchaeologist. He was a problem solver like few we've ever met, and a fieldworker and world traveller par excellence. Ken's personal and professional outlook embraced the whole world, in all its true facets and flavours, its complexities and eccentricities—he took the world, and all of us in it, as we came. His intellectual reputation extended well beyond Australia and was known to thousands of colleagues who may never have had the chance to meet him.
Ken was funny. It was a sense of humour that helped guide him in all situations, borne in part of never taking himself, or anything else whatsoever, too seriously. Ken belonged to that rare breed of truly grounded people. To say he didn't tend to stand on ceremony is to say the least. He preferred things practical and simple. He had little or no tolerance of honours. He took things as they were, not as they purported or professed or pretended to be. His refusal to kowtow to trend could come off as rather contrarian: he scoffed at anti-malarials and smart phones, even if most of the rest of us decided they were actually pretty useful! As a scientist, a common pattern for Ken would be to produce work of
Corresponding author: Kristofer M. Helgen Kris.Helgen(Australian. Museum
the most extraordinary calibre, and then publish it in the most obscure possible places. He took a personal pride and pleasure in such things. We admired him for it, though it had the effect that his work often wasn't recognized as widely for its brilliance as it should have been. But Ken sought no glory, period. In proper Aussie style, he was a true champion of the battlers and the underdogs, wherever he found them. Even when it came to his study animals, the more despised they might be in the public eye, the more he loved them. Snakes? Good. Rats? Even better. And the bigger the better.
Ken received many accolades across a varied and deeply respected academic and professional career, which included serving as Curator of Herpetology at the Western Australian Museum and as a Research Scientist at the CSIRO. Among his honours were his appointments as Research Associate at the American Museum of Natural History in New York and the Smithsonian in Washington DC, as well as receiving the Lifetime Achievement Award of the Australian Museum, an award very rarely bestowed. Especially important were Ken's lasting contributions to the conservation of wild landscapes and wild creatures, especially in Southeast Asia and New Guinea—he understood acutely that the world was a grander place than any of us can realize in our short time here. Despite his humility, we in the scientific community could see Ken's greatness, we recognized it, and hope to honour it in a small way with this volume. We have tried to capture the truly dizzying breadth and depth of Ken's knowledge and interests with the contributions presented
Received: 1 June 2020 Accepted: 26 October 2020 Published: 25 November 2020 (in print and online simultaneously) Publisher: The Australian Museum, Sydney, Australia (a statutory authority of, and principally funded by, the NSW State Government)
Citation: Helgen, Kristofer M., Julien Louys, and Sue O'Connor. 2020. The lives of creatures obscure, misunderstood, and wonderful: a volume in honour of Ken Aplin 1958—2019. In Papers in Honour of Ken Aplin, ed. Julien Louys, Sue O'Connor, and Kristofer M. Helgen. Records of the Australian Museum
72(5): 149—160. https://doi.org/10.3853/j.2201-4349.72.2020.1734
Copyright: O 2020 Helgen, Louys, O'Connor. This is an open access article licensed under a Creative Commons Attribution
4.0 International License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided
the original authors and source are credited.
CoL
150 Records of the Australian Museum (2020) Vol. 72
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Figure 1. Ken Aplin in his element in 2016, nra a late Holocene faunal depositi in southeastern NSW. Photo
courtesy of Brad Pillans.
in these pages. They describe new species of rodents, frogs, and bats, fossil and archaeological sites, and advances in rodent genetics and reproductive biology, amongst others. No topic was off bounds for Ken's piercing intellect, and even the smallest piece of scrappy bone or the tiniest crevice in a rock that harboured some animal remains was like finding treasure (Fig. 1).
Ken was a tough character. He chalked up most of the tropical diseases in the tropical disease textbook. Not the introductory textbook, but that massive reference book few physicians ever have to check out of the library. Malaria and typhus were long time traveling companions, and they came to know him as an opponent who gave as good as he got. A venomous snake bite and a broken back would not keep him down long, and were not to keep him from resuming the extraordinarily strenuous life of a New Guinea field biologist, a role he honed to perfection across four full decades of being ensconced in the natural and cultural worlds of that amazing island. So, literally backbreaking challenges could not tame him. When death stepped forward to introduce itself in his last phase of life, it did not at first realize the strength reflected straight back. Ken endured his final disease with the daunting Aplinian stoicism that all of us so admired. And that deep well of strength was to be seen not just in Ken, but in those around him who cared for him
and loved him, most of all of course Ken's wife Angela and his children. It was painful for all of us to watch his fight, and his pain, but we came to be grateful for the miracle of more time than we might have had any right to expect from the man once sickness set in. We'll remember these final years with sadness but will also remember the many years when Ken was such a whirlwind of strength and verve that to be anywhere near him was to be pulled in close to a world as exciting as any that could be dreamed.
Ken left a mark that resonates deeply on so many, and in so many places around the world. In tropical forests all across the great Indonesian island of Sulawesi, there flies a fruit bat called Nyctimene cephalotes aplini. A small fruit bat, with camouflage spotted green and khaki wings, this beautiful little bat has scattered forest seeds throughout its rainforest home for millions of years. We remember too the world's smallest bandicoot, Microperoyctes aplini, an exquisite gem of an animal with dark chocolate brown stripes ornamenting soft, fluffy brown fur. This little beast haunts the edges of lakes that dot the mountain vistas of north-western New Guinea: Ken's kind of place. Other creatures, too, were named in Ken's honour, one of the highest forms of recognition in the world of natural history. When we remember Ken, we also remember these beautiful and rare creatures, and be reminded of what a rare and beautiful soul the man was.
Helgen et al.: Papers in honour of Ken Aplin 151
Ken Aplin's Eponyms T extinct taxa Nyctimene cephalotes aplini Kitchener, Packer, and Suyanto, 1995 (Chiroptera: Pteropodidae) Microperoyctes aplini Helgen and Flannery, 2004 (Peramelemorphia: Peramelidae) tAlormys aplini Louys, O'Connor, Mahirta, Higgins, Hawkins, and Maloney, 2018 (Rodentia: Muridae) tUromys aplini Cramb, Hocknull, and Price, 2020 (Rodentia: Muridae) Litoria aplini Richards and Donnellan, 2020 (Anura: Pelodryadidae)
Taxa Described by Ken Aplin
T extinct taxa
Ordinal group names Suborder Agreodontia Beck, Travouillon, Aplin, Godthelp, and Archer, 2013
Infraorder Phascolarctomorphia Aplin and Archer, 1987 Infraorder Vombatomorphia Aplin and Archer, 1987
Family group names
TFamily Holoclemensiidae Aplin and Archer, 1987 Family Acrobatidae Aplin, 1987
Tribe Apodemini Lecompte, Aplin, Denys, Catzeflis, Chades, and Chevret, 2008 Tribe Arvicanthini Lecompte, Aplin, Denys, Catzeflis, Chades, and Chevret, 2008 Tribe Millardini Lecompte, Aplin, Denys, Catzeflis, Chades, and Chevret, 2008 Tribe Malacomyini Lecompte, Aplin, Denys, Catzeflis, Chades, and Chevret, 2008 Tribe Praomyini Lecompte, Aplin, Denys, Catzeflis, Chades, and Chevret, 2008
Genus group names
TWatutia Flannery, Hoch, and Aplin, 1989 Lemdubuoryctes Kear, Aplin, and Westerman, 2016 (junior synonym of Peroryctes)
Species group names
tDendrolagus noibano Flannery, Mountain, and Aplin, 1982 tProtemnodon tumbuna Flannery, Mountain, and Aplin, 1982 +Protemnodon nombe Flannery, Mountain, and Aplin, 1982 Litoria exophthalmia Tyler, Davies, and Aplin, 1986 Mallomys istapantap Flannery, Aplin, Groves, and Adams, 1989 Mallomys gunung Flannery, Aplin, Groves, and Adams, 1989 + Watutia novaeguineae Flannery, Hoch, and Aplin, 1989 Rattus timorensis Kitchener, Aplin, and Boeadi, 1991 Glaphyromorphus butlerorum Aplin, How, and Boeadi, 1993 Ramphotyphlops pilbarensis Aplin and Donnellan, 1993 Ramphotyphlops splendidus Aplin, 1998 Ramphotyphlops longissimus Aplin, 1998 Ramphotyphlops ganei Aplin, 1998 Diplodactylus klugei Aplin and Adams, 1998 Ctenotus maryani Aplin and Adams, 1998 Menetia surda cresswelli Aplin and Adams, 1998 {Petauroides ayamaruensis Aplin, 1999 Dactylopsila kambuayai Aplin, 1999 Pseudantechinus roryi Cooper, Aplin, and Adams, 2000 Varanus bushi Aplin, Fitch, and King, 2007 Delma tealei Maryan, Aplin, and Adams, 2007 Delma desmosa Maryan, Aplin, and Adams, 2007 TCoryphomys musseri Aplin and Helgen, 2010 Microhydromys argenteus Helgen, Leary, and Aplin, 2010 Phascogale tapoatafa wambenger Rhind and Aplin, 2015 Phascogale tapoatafa kimberleyensis Aplin and Rhind, 2015 Aprasia wicherina Maryan, Adams, and Aplin, 2015 Delma hebesa Maryan, Brennan, Adams, and Aplin, 2015 Rattus detentus Timm, Weijola, Aplin, Flannery, and Pine, 2016 +Peroryctes aruensis (Kear, Aplin, and Westerman, 2016) Halmaheramys wallacei Fabre, Reeve, Fitriana, Aplin, and Helgen, 2017 + Macroderma handae Aplin and Armstrong, 2020
152 Records of the Australian Museum (2020) Vol. 72 Ken Aplin's Bibliography
1977
Aplin, K. P. 1977. Preliminary faunal report. In Abercrombie Arch Shelter: An Excavation Near Bathurst, NSW, ed. I. Johnson. Australian Archaeology 6: 28—40.
https://doi.org/10.1080/03122417.1977.12093316
1981
Aplin, K. P. 1981. The Kamapuk Fauna: A Late Holocene Vertebrate Faunal Sequence from the Western Highlands District, Papua New Guinea. B.A. (Hons.) thesis. The Australian National University, Canberra.
Aplin, K. P. 1981. Faunal remains from the archaeological sites in Mangrove Creek Catchment. In Mangrove Creek Dam Salvage Excavation Project, ed. V. Attenbrow, pp. 1—73. National Parks and Wildlife Service, New South Wales.
Aplin, K. P., R. Silcox, N. Stern, and E. Williams. 1981. An archaeological survey of the western shore of Lake Cawndilla, Kinchega National Park. In Darling Surveys I, ed. J. Hope, pp. 70-88. Canberra: The Australian National University, Occasional Papers in Prehistory, Research School of Pacific Studies.
1983
Aplin, K. P. 1983. Zygomaturus trilobus. A rhinoceran relative of the wombats and Euryzygoma dunense. Cheeky giant of the Pliocene. In Prehistoric Animals of Australia, ed. S. Quirk and M. Archer, pp. 58-61. Sydney: Australian Museum.
Flannery, T. F., M.-J. Mountain, and K. P. Aplin. 1983. Quaternary kangaroos (Macropodidae; Marsupialia) from Nombe Rockshelter, Papua New Guinea, with comments on megafaunal extinction in the New Guinea Highlands. Proceedings of the Linnean Society of New South Wales 107: 77—99.
1984
Archer, M., and K. P. Aplin. 1984. Humans among primates: stark naked in a crowd. In Vertebrate Zoogeography and Evolution in Australasia, ed. M. Archer and G. Clayton, pp. 949—993. Perth: Hesperion Press.
Coutts, P. J. F., K. Aplin, and N. Taylor. 1984. Archaeological investigations at Captain's Point, Mallacoota. In Coastal Archaeology in Southeastern Victoria, ed. P. J. F. Coutts, pp. 1—184. Records of the Victorian Archaeological Survey, No.14.
1986
Harding, H. R., K. P. Aplin, and C. D. Shorey. 1986. Cladistic analysis of marsupial sperm structure: methodological perspectives. Development, Growth and Differentiation Suppl. (1986): 70.
Tyler, M. J., M. Davies, and K. Aplin. 1986. A new stream- dwelling species of Litoria (Anura: Hylidae) from New Guinea. Transactions of the Royal Society of South Australia 110: 63—67.
1987
Aplin, K. P. 1987. Basicranial anatomy of the early Miocene diprotodontian Wynyardia bassiana (Marsupialia: Wynyardiidae) and its implications for wynyardiid phylogeny and classification. In Possums and Opossums: Studies in Evolution, ed. M. Archer, pp. 369-391. Sydney: Royal Zoological Society of NSW.
Aplin, K. P., and M. Archer. 1987. Recent advances in marsupial systematics, with a new syncretic classification of marsupials. In Possums and Opossums: Studies in Evolution, ed. M. Archer, pp. xv-Ixxii. Sydney: Royal Zoological Society of NSW.
Harding, H. R., K. P. Aplin, and C. D. Shorey. 1987. Parsimony analysis of marsupial sperm structure: a preliminary report. In New Horizons in Sperm Cell Research, ed. H. Mohri, pp. 375—385 Tokyo: Japan Science Society Press and New York: Garden and Breach Science Publishers.
Hughes, R. L., L. S. Hall, K. P. Aplin, and M. Archer. 1987. Organogenesis and fetal membranes in the New Guinea Pen-tailed Possum, Distoechurus pennatus (Acrobatidae: Marsupialia). In Possums and Opossums: Studies in Evolution, ed. M. Archer, pp. 715—724. Sydney: Royal Zoological Society of NSW.
1989
Donnellan, S. C., M. Adams, and K. P. Aplin. 1989. A biochemical and morphological study of Rana (Anura: Ranidae) from the Chimbu Province, Papua New Guinea. Herpetologica 45(3): 336-343.
Donnellan, S. C., and K. P. Aplin. 1989. Resolution of cryptic species in the New Guinean lizard, Sphenomorphus jobiensis (Scincidae) by electrophoresis. Copeia 1989(1): 81-88. https://doi.org/10.2307/1445608
Flannery, T. F., K. P. Aplin, C. P. Groves, and M. Adams. 1989. Revision ofthe New Guinean murid genus Mallomys (Rodentia), with description of two new species from subalpine habitats. Records of the Australian Museum 41(1): 83-105. https://doi.org/10.3853/).0067-1975.41.1989.137
Flannery, T. F., E. Hoch, and K. Aplin. 1989. Macropodines from the Pliocene Otibanda Formation, Papua New Guinea. A/cheringa
13: 145-152.
1990
Aplin, K. P. 1990. Basicranial Regions of Diprotodontian Marsupials: Anatomy, Ontogeny and Phylogeny. Unpublished Ph.D. thesis. University of New South Wales, Sydney.
Baverstock, P. R., T. F. Flannery, K. P. Aplin, J. Birrell, and M. Kreig. 1990. Albumin immunologic relationships of the bandicoots (Perameloidea: Marsupialia)—a preliminary report. In Bandicoots and Bilbies, ed. J. H. Seebeck, P. R. Brown, R. L. Wallis, and C. H. Kemper, pp. 13-18. Sydney: Surrey Beatty and Sons.
Harding, H. R., and K. P. Aplin. 1990. Phylogenetic affinities of the koala: a reassessment of the spermatozoal evidence. In Biology of the Koala, ed. A. Lee, K. Handasyde, and G. Sanson, pp. 1-31. Sydney: Surrey Beatty and Sons.
Hughes, R. L., L. S. Hall, M. Archer, and K. P. Aplin. 1990. Observations on placentation and development in Echymipera kalubu. In Bandicoots and Bilbies, ed. J. H. Seebeck, P. R. Brown, R. L. Wallis, and C. H. Kemper, pp. 259—270. Sydney: Surrey Beatty and Sons.
Kirsch, J. A. W., M. S. Springer, C. Krajewski, M. Archer, K. P. Aplin, and A. Dickerman. 1990. DNA/DNA hybridization studies of carnivorous marsupials. I. The intergeneric relationships of bandicoots (Marsupialia: Perameloidea). Journal of Molecular Evolution 30: 434—448. https://doi.org/10.1007/BF02101115
Springer, M. S., J. A. W. Kirsch, K. P. Aplin, and T. F. Flannery. 1990. DNA hybridization, cladistics and the phylogeny of phalangerid marsupials. Journal of Molecular Evolution 30: 298-311.
https://doi.org/10.1007/BF02100000
1991
Kitchener, D. J., K. P. Aplin, and Boeadi. 1991. A new species of Rattus from Gunung Mutis, south west Timor Island, Indonesia. Records of the Western Australian Museum 15: 445—461.
1992
Harding, H. R., and K. P. Aplin. 1992. Approaches to the study of structure-function relationships in spermatozoa. In Comparative Spermatology 20 Years Afier, ed. B. Baccetti, pp. 955—959. Italy: Serona Symposia Publications, Raven Press.
Mahony, M., S. C. Donnellan, and K. Aplin. 1992. Karyotypes of Australo-Papuan microhylid frogs (Anura Microhylidae). Herpetologica 48(2): 184—192.
Springer, M. S., G. McKay, K. P. Aplin, and J. A. W. Kirsch. 1992. Relationships among ringtail possums (Marsupialia: Pseudocheiridae) based on DNA hybridization. Australian Journal of Zoology 40: 423—435.
https://doi.org/10.1071/209920423
1993
Aplin, K. P., P. R. Baverstock, and S. C. Donnellan. 1993. Albumin immunological evidence for the time and mode of origin of the New Guinean terrestrial mammal fauna. Science in New Guinea 19: 131—145.
Aplin, K. P., and S. C. Donnellan. 1993. A new species of blindsnake, genus Ramphotyphlops, with a redescription of R. hamatus Storr 1982. Records of the Western Australian Museum 16: 243-256.
Aplin, K. P., and R. A. How. 1993. A window west: a perspective on Western Australian herpetology. In Herpetology in Australia. A Diverse Discipline, ed. D. Lunney and D. Ayers, pp. 337—345. Sydney: Royal Society of New South Wales. https://doi.org/10.7882/RZSNSW.1993.052
Aplin, K. P., R. A. How, and Boeadi. 1993. A new species of the Glaphyromorphus isolepis species group (Lacertilia: Scincidae) from Sumba Island, Indonesia. Records of the Western Australian Museum 16: 235—242.
Aplin, K. P., and P. A. Woolley. 1993. Notes on the distribution and reproduction of the Papuan Bandicoot, Microperoryctes papuensis (Peroryctidae, Peramelomorphia). Science in New Guinea 19: 109—112.
Colgan, D., T. F. Flannery, J. Trimble, and K. P. Aplin. 1993. Electrophoretic and morphological analysis of the systematics of the Phalanger orientalis (Marsupialia) species complex in New Guinea and the Solomon Islands. Australian Journal of Zoology 41: 355—378.
https://doi.org/10.1071/209930355
1995
Breed, W. G., and K. P. Aplin. 1995. Sperm morphology of murid rodents from New Guinea and the Solomon Islands— phylogenetic implications. Australian Journal of Zoology 43: 17-30.
Harding, H. R., K. P. Aplin, and M. Mazur. 1995. Ultrastructure of spermatozoa of Australian Blindsnakes, Ramphotyphlops spp. (Typhlopidae, Squamata): first observations on the mature spermatozoon of scolecophidian snakes. In Advances in Spermatozoal Phylogeny and Taxonomy, ed. B. G. M. Jamieson and J.-L. Justine, pp. 385—396. Paris: Memoires Muséum nationale d'Histoire naturelle, vol. 166.
Helgen et al.: Papers in honour of Ken Aplin 153
1996
Burbidge, A. A., and K. P. Aplin. 1996. Endangered: marsupial moles. Landscope 11: 34.
1998
Aplin, K. P. 1998. Three new blindsnakes (Squamata: Typhlopidae) from northwestern Australia. Records of the Western Australian Museum 19: 1—12.
Aplin, K. P. 1998. Vertebrate zoogeography of the Bird's Head of Irian Jaya, Indonesia. In Perspectives on the Bird s Head Peninsula of Irian Jaya, Indonesia: Proceedings of an Interdisciplinary Conference, ed. J. Miedema, C. Odé, and M. A. C. Dam, pp. 803—890. Leiden: Rodopi Publishers.
Aplin, K. P., and M. Adams. 1998. New species and subspecies of gekkonid and scincid lizards (Squamata) from the Carnarvon Basin region of Western Australia: morphological and genetic discrimination. Journal of the Royal Society of Western Australia 81: 201—223.
Pasveer, J. M. and K. P. Aplin. 1998. Late Pleistocene to Recent faunal succession and environmental change in lowland New Guinea: evidence from the Bird's Head of Irian Jaya, Indonesia. In Perspectives on the Bird s Head Peninsula of Irian Jaya, Indonesia: Proceedings of an Interdisciplinary Conference, ed. J. Miedema, C. Odé, and M. A. C. Dam, pp. 891—930. Leiden: Rodopi Publishers.
1999
Aplin, K. P. 1999. *Amateur" taxonomy in Australian herpet- ology—help or hindrance? Monitor, Journal of the Victorian Herpetological Society 10 (2/3).
Aplin, K. P., and S. C. Donnellan. 1999. An extended description of the Pilbara Death Adder, Acanthophis wellsi Hoser (Serpentes: Elapidae), with notes on the Desert Death Adder, A. pyrrhus, Boulenger, and identification ofa possible hybrid zone. Records of the Western Australian Museum 19: 277—298.
Aplin, K. P., J. M. Pasveer, and W. E. Boles. 1999. Late Quaternary vertebrates from the Bird's Head Peninsula, Irian Jaya, Indonesia, including descriptions of two previously unknown marsupial species. Records of the Western Australian Museum, Supplement no. 57: 351—387.
2000
Aplin, K. P., A. Paino, and L. Sleep. 2000. Building Frog Friendly Gardens. Perth: Western Australian Museum.
Cooper, N. K., K. P. Aplin, and M. Adams. 2000. A new species of false antechinus (Marsupialia: Dasyuromorphia: Dasyuridae) from the Pilbara region, Western Australia. Records of the Western Australian Museum 20: 115—136.
Donnellan, S. C., K. P. Aplin, and P. J. Dempsey. 2000. Genetic and morphological variation in Australian Christinus (Squamata: Gekkonidae): preliminary overview with recognition of a cryptic species on the Nullarbor Plain. Australian Journal of Zoology 48: 289—315.
https://doi.org/10.1071/2098015
McKenzie, N. L., J. K. Rolfe, K. P. Aplin, M. A. Cowan, and L. A. Smith. 2000. Herpetofauna of the southern Carnarvon Basin, Western Australia. Records of the Western Australian Museum Supplement no. 61: 335—360. https://doi.org/10.18195/issn.0313-122x.61.2000.335-360
Withers, P. C., K. P. Aplin, and Y. L. Werner. 2000. Metabolism and evaporative water loss of Western Australian geckos (Reptilia: Sauria: Gekkonomorpha). Australian Journal of Zoology 48: 111—126.
https://doi.org/10.1071/2099007
154 Records of the Australian Museum (2020) Vol. 72
2001
Aplin, K. P., N. K. Cooper, R. A. How, J. B. Hutchins, R. E. Johnstone, and L. A. Smith. 2001. Introduction to the checklists of vertebrates of Western Australia. Records of the Western Australian Museum Supplement no. 63: 1—7.
22x.63.2001.001-007
Aplin, K. P., and S. Johnson. 2001. Community involvement in the detection, surveillance and management of frog diseases: a case study from Alcoa frogWAtch, southwest Australia. In Developing Management Strategies to Control Amphibian Diseases: Decreasing the Risks Due to Communicable Diseases, ed. R. Speare, p. 47. Townsville: School of Public Health and Tropical Medicine, James Cook University.
Aplin, K. P., and P. Kirkpatrick. 2001. In pursuit ofthe frog fungus. Landscope 16(3): 10-16.
Aplin, K. P., and P. Kirkpatrick. 2001. Chytridiomycosis in southwest Australia: historical sampling documents the date of introduction, rates of spread and seasonal epidemiology, and sheds new light on chytrid ecology. In Developing Management Strategies to Control Amphibian Diseases: Decreasing the Risks Due to Communicable Diseases, ed. R. Speare, p. 24. Townsville: School of Public Health and Tropical Medicine, James Cook University.
Aplin, K. P., and L. A. Smith. 2001. Checklist of the frogs and reptiles of Western Australia. Records of the Western Australian Museum Supplement no. 63: 51—74.
https://doi.org/10.18195/issn.0313-1
https://doi.org/10.18195/issn.0313-122x.63.2001.051-074
2002
Jacob, J., P. R. Brown, K. P. Aplin, and G. R. Singleton. 2002. Ecologically-based management of pest rodents in rice-based agro-ecosystems in southeast Asia. In Proceedings, Twentieth Vertebrate Pest Conference, Silver Legacy Resort-Casino, Reno, Nevada, March 4—7, 2002, ed. R. M. Timm and R. H. Schmidt, pp. 67—74. Davis: University of California, Davis.
O'Connor, S., K. P. Aplin, M. Spriggs, P. Veth, and L. K. Ayliffe. 2002. From savannah to rainforest: changing environments and human occupation at Liang Lemdubu, Aru Islands, Maluku (Indonesia). In Bridging Wallaces Line: The Environmental and Cultural History and Dynamics of the Southeast Asian- Australian Region, ed. P. Kershaw, B. David, N. Tapper, D. Penny, and J. Brown, pp. 279—306, Advances in Geoecology Series, no. 34. Reiskirchen: Catena Verlag.
2003
Aplin, K. P., P. R. Brown, J. Jacob, C. J. Krebs, and G. R. Singleton. 2003. Field methods for rodent studies in Asia and the Indo- Pacific. ACIAR Monograph no. 100, 223 pp. Canberra: Australian Centre for International Agricultural Research.
Aplin, K. P., T. Chesser, and J. ten Have. 2003. Evolutionary biology of the genus Rattus: profile of an archetypal rodent pest. In Rats, Mice and People: Rodent Biology and Management, ed. G. R. Singleton, L. A. Hinds, C. J. Krebs, and D. M. Spratt, pp. 487—498. ACIAR Monograph no. 96, 564 pp.
Aplin, K. P., A. Frost, N. P. Tuan, L. P. Lan, and N. M. Hung. 2003. Identification of rodents of the genus Bandicota in Vietnam and Cambodia. In Rats, Mice and People: Rodent Biology and Management, ed. G. R. Singleton, L. A. Hinds, C. J. Krebs, and D. M. Spratt, pp. 531—535. ACIAR Monograph no. 96, 564 pp.
Aplin, K. P., and G. R. Singleton. 2003. Balancing rodent management and small mammal conservation in agricultural landscapes: challenges for the present and the future. In Rats, Mice and People: Rodent Biology and Management, ed. G. R. Singleton, L. A. Hinds, C. J. Krebs, and D. M. Spratt, pp. 80—88. ACIAR Monograph no. 96, 564 pp.
Douangboupha, B., K. P. Aplin, and G. R. Singleton. 2003. Rodent outbreaks in the uplands of Laos: analysis of historical patterns and the identity of nuu khii. In Rats, Mice and People: Rodent Biology and Management, ed. G. R. Singleton, L. A. Hinds, C. J. Krebs, and D. M. Spratt, pp. 103-111. ACIAR Monograph no. 96, 564 pp.
Khamphoukeo, K., B. Douangboupha, K. P. Aplin, and G. R. Singleton. 2003. Pest and non-pest rodents in the upland agricultural landscape of Laos: a progress report. In Rats, Mice and People: Rodent Biology and Management, ed. G. R. Singleton, L. A. Hinds, C. J. Krebs, and D. M. Spratt, pp. 284—289. ACIAR Monograph no. 96, 564 pp.
Lan, L.P., K. P. Aplin, N. M. Hung, N. V. Quoc, H. V. Chien, N. D. Sang, and G. R. Singleton. 2003. Rodent communities and historical trends in rodent damage in the Mekong Delta of Vietnam: establishing an ecological basis for effective pest management. In Rats, Mice and People: Rodent Biology and Management, ed. G. R. Singleton, L. A. Hinds, C. J. Krebs, and D. M. Spratt, pp. 290—296. ACIAR Monograph no. 96, 564 pp.
Singleton, G. R, L. Smythe, G. Smith, D.M., Spratt, K. P. Aplin, and A. L. Smith. 2003. Rodent diseases in Southeast Asia and Australia: inventory of recent surveys. In Rats, Mice and People: Rodent Biology and Management, ed. G. R. Singleton, L. A. Hinds, C. J. Krebs, and D. M. Spratt, pp. 25-30. ACIAR Monograph no. 96, 564 pp.
2004
Joshi, R. C., E. B. Gergon, K. P. Aplin, G. R. Singleton, A. R. Martin, J. C. Cabigat, A. Cayong, N. V. Desamero, and L. S. Sebastian. 2004. Rodents and other small mammals in Banaue and Hungduan rice terraces, Philippines. /nternational Rice Research Notes 29(1): 44—46.
Mahirta, K. P. Aplin, D. Bulbeck, W. E. Boles, and P. Bellwood. 2004. Pia Hudale Rockshelter: a terminal Pleistocene occupation site on Roti Island, Nusa Tenggara Timur, Indonesia. In Quaternary Research in Indonesia, ed. S. Keates and J. M. Pasveer, pp. 361—380. Modern Quaternary Research in South East Asia No. 18. Rotterdam: A. A. Balkema.
Rabosky, D. L., K. P. Aplin, S. C. Donnellan, and S. B. Hedges. 2004. Molecular phylogeny of blindsnakes (Ramphotyphlops) from western Australia and resurrection of Ramphotyphlops bicolor (Peters, 1857). Australian Journal of Zoology 52: 531—548.
https://doi.org/10.1071/72004045 Suzuki, H., T. Shimada, M. Terashima, K. Tsuchiya, and K. P. Aplin. 2004. Temporal, spatial, and ecological modes of evolution of Eurasian Mus based on mitochondrial and nuclear gene
sequences. Molecular Phylogenetics and Evolution 33: 626—646.
https://doi.org/10.1016/j.ympev.2004.08.003
2005
Aplin, K., and J. Pasveer. 2005. Mammals and other vertebrates from late Quaternary archaeological sites on Pulau Kobroor, Aru Islands, eastern Indonesia. In Archaeology of the Aru Islands, ed. S. O'Connor, P. Veth, and M. Spriggs, pp. 41—62. Terra Australis 22. Canberra: Pandanus Press. https://doi.org/10.22459/TA22.2007.03
Chinen, A. M., H. Suzuki, K. P. Aplin, K. Tsuchiya, and S. Suzuki. 2005. Preliminary genetic characterization of two lineages of black rats (Rattus rattus sensu lato) in Japan, with evidence for introgression at multiple localities. Genes and Genetic Systems 80: 367—375.
https://doi.org/10.1266/ggs.80.367
Hope, G., and K. Aplin. 2005. Environmental change in the Aru Islands. In Archaeology of the Aru Islands, ed. S. O’ Connor, P. Veth, and M. Spriggs, pp. 25—40. Terra Australis 22. Canberra: Pandanus Press.
1) 22450
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O'Connor, S., K. Aplin, J. Pasveer, and G. Hope. 2005. Liang Nabulei Lisa: a Late Pleistocene and Holocene sequence from the Aru Islands. In Archaeology of the Aru Islands, ed. S. O'Connor, P. Veth, and M. Spriggs, pp. 125-161. Terra Australis 22. Canberra Pandanus Press. https://do TA22.2007.07
O’ Comi S Ks Aplin. K. Szabó, J. Pasveer, P. Veth, and M. Spriggs. 2005. Liang Lemdubu, a Pleistocene cave site in the Aru Islands. In Archaeology of the Aru Islands, ed. S. O' Connor, P. Veth, and M. Spriggs, pp. 171—204. Terra Australis 22. Canberra: Pandanus Press.
https://doi.org/10.22459/
i.org/10.2245
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2006
Aplin, K. P. 2006. Ten million years of rodent evolution in Australasia: phylogenetic evidence and a speculative historical biogeography. In Evolution and Biogeography of Australasian Vertebrates, ed. J. R. Merrick, M. Archer, G. M. Hickey, and M. S. Y. Lee, pp. 707—744. Oatlands, Sydney: Auscipub.
Aplin, K. P., P. R. Brown, G. R. Singleton, B. Douang Boupha, and K. Khamphoukeo. 2006. Rodents in the rice environments of Laos. In Rice in Laos, ed. J. M. Schiller, M. B. Chanphengxay, B. Linquist, and S. Appa Rao, pp. 291—308. Los Baños, Philippines: IRRI and Canberra: ACIAR.
Aplin, K. P., A. J. Fitch, and D. J. King. 2006. A new species of Varanus Merrem (Squamata: Varanidae) from the Pilbara region of Western Australia, with observations on sexual dimorphism in SIDSCLY ER zpecies: Zootaxa 1313: 1-38. https://doi.org/10.11646/zootaxa.1313.1.1
Dousneboupha, B., P R. Brow, K. Khamphoukeo, K. P. Aplin, and G. R. Singleton. 2006. Population dynamics of rodent pest species in upland farming systems of Lao PDR. The Lao Journal of Agriculture and Forestry Jan—June 2006: 109—121. Republished in Kasetsart Journal (Natural Science) 43: 125—131 (2009).
2007
David, B., A. Fairbairn, K. Aplin, L. Murepe, M. Green, J. Stanisic, M. Weisler, D. Simala, T. Kokents, J. Dop, and J. Muke. 2007. OJP, a terminal Pleistocene archaeological site from the Gulf Province lowlands , Papua New Guinea. Archaeology in Oceania 42: 31-33. https://doi.org/10.1002/j.1834-4453.2007.tb00013.x
Hope, G. s. and K. P. Aplin. 2007. Palaeontology of Papua. In The Ecology of Papua. Part One. The Ecology of Indonesia Series. Volume V1, ed. A. J. Marshall and B. M. Beehler, pp. 246—254. Singapore: Periplus Editions.
Maryan, B., K. P. Aplin, and M. Adams. 2007. Two new species of the Delma tincta group (Squamata: Pygopodidae) with remarks on patterns of species endemism in northwestern Australia. ROCORIS of the estern Australian Museum 23: 273—305. https://doi.org/10.18195/issn.0312-3162.23(3).2
O’ Comot s. and K. P. Aplin. 2007. A matter of balance: an overview of Pleistocene occupation history and the impact of the Last Glacial Phase in East Timor and the Aru Islands, eastern Indonesia: Milius A in Oceania 42: 82—90.
rg/10.22459/TA22.2007
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https://doi. Shimada, T. K. P. Apii P. Jenkins, and H. Suzuki. 2007. Rediscovery of Mus nitidulus Blyth (Rodentia, Muridae), an endemic murine rodent of the central basin of Myanmar. Zootaxa 1498(1): 45—68. https://doi.org/10.11646/zootaxa.1498.1.4 Shimada, T., K. P. Aplin, K. Jyogahara, K.-L. Lin, J.-P. Gonzalez, V. Herbreteau, and H. Suzuki. 2007. Complex phylogeographic structuring in a continental small mammal from East Asia, the rice field mouse, Mus caroli (Rodentia, Muridae). Mammal Study 32: 49—62.
https://doi.org/10.3106/1348-6160(2007)32[49:CPSIAC]2.0.CO;2
Helgen et al.: Papers in honour of Ken Aplin 155
Singleton, G. R., P. R. Brown, J. Jacob, K. P. Aplin, and Sudarmaji. 2007. Unwanted and unintended effects of culling: a case for ecologically-based rodent management. /ntegrative Zoology 2: 247—259. https://doi.org/10.1111/j.1749-4877.2007.00067.x
Veth, P. M., K. P. Aplin, L. Wallis, T. Manne, T. Pulsford, A. White, and A. Chappell. 2007. The archaeology of Montebello Islands, Northwest Australia: late Quaternary foragers on an arid coastline. BAR, International Series, 1668. Oxford: Archaeopress.
https://doi.org/10.30861/9781407301037
2008
Aplin, K., S. Donnellan, and J. Dell. 2008. The herpetofauna of Faure Island, Shark Bay, Western Australia. Records of the Western Australian Museum Supplement no. 75: 39-53. https://doi.org/10.18195/issn.0313-122x.75.2008.039-053
Lecompte, E., K. Aplin, C. Denys, F. Catzeflis, M. Chades, and P. Chevret. 2008. Phylogeny and biogeography of African Murinae based on mitochondrial and nuclear gene sequences, with a new tribal classification of the subfamily. BMC Evolutionary Biology 8: 199. https://doi.org/10.1186/1471-2148-8-199
O’Connor, S., K. Aplin, and S. Collins. 2008. A small salvage excavation in Windjana Gorge, Kimberley, Western Australia. Archaeology in OLE 15 IA 81.
https://doi.org/10.1002/j.1834-445 08.tb00032.x
2009
Angelakis, E., K. Khamphoukeo, D. Grice, P. N. Newton, V. Roux, K. Aplin, D. Raoult, and J. M. Rolain. 2009. Molecular detection of Bartonella species in rodents from the Lao PDR. Clinical VMACHOPIDRARY and prn 15 GAN dace https://doi.org/10.1111/j
Aplin, K. P. 2009. On the origin of rats. Australasian Science. April 2009: 28-31.
Shimada, T., J. J. Sato, K. P. Aplin, and H. Suzuki. 2009. Comparative analysis of evolutionary modes in Mc/r coat color gene in wild mice and mustelids. Genes and Genetic Systems 84: 225-231.
https://doi.or
1469-0691.2008.02177
rg/10.1266/99s.84.225
Sutton, A., M.-J. Mountain, K. Aplin, S. Bulmer, and T. Denham. 2009. Archaeozoological records for the highlands of New Guinea: a review of current evidence. Australian Archaeology 69: 41-58.
https://doi.org/10.1080/(
3122417.2009.11681900
2010
Aplin, K. P., and K. M. Helgen. 2010. Quaternary murid rodents of Timor part I: new material of Coryphomys buehleri Schaub, 1937, and description of a second species of the genus. Bulletin oft ble American Museum of Natural History 341: 1—80. https://doi.org/10.1206/692.1
Aplin, | K. P., and J. Lalsiamliana. 2010. Chronicle and impacts of the 2005-2009 mautam in Mizoram. In Rodent Outbreaks: Ecology and Impacts, ed. G. Singleton, S. Belmain, P. Brown, and B. Hardy. Los Baños, Philippines: International Rice Research Institute.
Aplin K., F. Ford, and P. Hiscock. 2010. Early Holocene human occupation and environment of the southeast Australian Alps: new evidence from the Yarrangobilly Plateau, New South Wales. In Altered Ecologies: Fire, Climate and Human Influence on Terrestrial Landscapes, ed. S. Haberle, J. Stevenson, and M. Prebble, pp. laa: OA 32. Canberra: ANU E Press.
https://doi.org/10 9/TA32.11.2010
156 Records of the Australian Museum (2020) Vol. 72
Aplin, K. P., K. M. Helgen, and D. P. Lunde. 2010. A review of Peroryctes broadbenti, the giant bandicoot of Papua New Guinea. American Museum Novitates 3696: 1—41. https://doi.org/10.1206/3696.2
David, B., J—M. Geneste, K. Aplin, J.-J. Delannoy, N. Araho, C. Clarkson, K. Connell, S. Haberle, B. Barker, L. Lamb, J. Stanisic, A. Fairbairn, R. Skelly, and C. Rowe. 2010. The Emo site (OAC), Gulf Province, Papua New Guinea: resolving long- standing questions of antiquity and implications for the history of the ancestral Hiri maritime trade. Australian Archaeology 70: 39-54. https://doi.org/10.1080/0312241 7.2010.11681910
Donnellan, S. C., K. P. Aplin, and T. Bertozzi. 2010. Species boundaries in the Rana arfaki group (Anura: Ranidae) and phylogenetic relationships to other New Guinean Rana. Zootaxa 2496: 49—62.
Helgen, K. M., T. Leary, and K. P. Aplin. 2010. A revision of Microhydromys (Rodentia: Murinae), with description of a new species from southern New Guinea. American Museum Novifates 3676: 1—22. https://doi.org/10.1206/632.1
Louys, I. K. P. Aplin, R. M. D. Beck, and M. Archer. 2010. Cranial anatomy of Oligo-Miocene koalas (Diprotodontia: Phascolarctidae): stages in the evolution of an extreme leaf-eating specialization: agen of Vertebrate Paleontology 29: 981—992. https://doi.org/10.1671/039.029.0412
McNiven, L, B. David, K. Aplin, M. Pivoru, W. Pivoru, A. Sexton, J. Brown, C. Clarkson, K. Connell, J. Stanisic, M. Weisler. S. Haberle, A. Fairbairn, and N. Kemp. 2010. Historicising the present: late Holocene emergence of a rainforest hunting camp, Gulf Province, Papua New Guinea. Australian Archaeology TI. 41—56.
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2010.116
Müunss: K., R. Retallick, K. R McDonald, D. Mendez, K. Aplin, P. Kirkpatrick, L. Berger, D. Hunter, H. B. Hines, R. Campbell, M. Pauza, M. Driessen, R. Speare, S. J. Richards, M. Mahony, A. Freeman, A. D. Phillott, J. Hero, K. Kriger, D. Driscoll, A. Felton, R. Puschendorf, and L. F. Skerratt. 2010. The distribution and host range of the pandemic disease chytridiomycosis in Australia, spanning surveys from 1956-2007. Ecology 91: ioo dans. https://doi.org/10.1890/09-1608.1
Nunome, M. C. Ishimori, K. P. Aplin, K. Tsuchiya, H. Yonekawa, K. Moriwaki, and H. Suzuki. 2010. Detection of recombinant haplotypes in wild mice (Mus musculus) provides new insights into the origin of Japanese mice. Molecular Ecology 19(12): 2404. P489. https ://doi.org/10.1111/j.1365-294X.2010.04651 .x
O’ keen S. K. pce E. [st Pierre, and Y.-x. Feng. 2010. Faces of the ancestors revealed: discovery and dating ofa Pleistocene-age Ed in Lene pas CaN East Timor. Antiquity 84: 649—665. https://doi.org/10.1017/S000: 3X00100146
O'Connor, S., A. Barham, ! M. Spriggs, P. Veth, K. Aplin, and E. St Pierre. 2010. Cave archaeology and sampling issues in the tropics: a case study from Lene Hara Cave, a 42,000 year old occupation site in East Timor, Island Southeast Asia. Australian Reis 71: 29-40. https://doi.org/10.1080/03122417.2010.11689382
Shimada, T., K. P Aplin, and H. Suzuki. 2010. Mus lepidoides (Muridae, Rodentia) of central Burma is a distinct species of potentially great evolutionary and biogeographic significance. AE Scene 27: 449-459. https://doi.org/10.2108/zsj.27.449
singletob. G. Rz, s. Bélnam. P. R. Brown, K. Aplin, and N. M. Htwe. 2010. Impacts of rodent outbreaks on food security in Asia. Wildlife Research 37. 355—359.
WR10084
https://doi.org/10.1071A
2011
Aplin, K. P., and E. Kale. 2011. Non-volant mammals of the Muller Range, Papua New Guinean. In Rapid Biological Assessments of the Nakanai Mountains and the Upper Strickland Basin: Surveying the Biodiversity of Papua New Guinea s Sublime Karst Environments, ed. S. J. Richards and B. G. Gamui, RAP Bulletin of Biological Assessment 60, pp. 211—221. Arlington: Conservation International.
Aplin, K. P., and M. Opiang. 2011. Mammals of the Nakanai Mountains, East New Britain Province, Papua New Guinea. In Rapid Biological Assessments of the Nakanai Mountains and the Upper Strickland Basin: Surveying the Biodiversity of Papua New Guinea’ Sublime Karst Environments, ed. S. J. Richards and B. G. Gamui, RAP Bulletin of Biological Assessment 60, pp. 85-103. Arlington: Conservation International.
Aplin K. P., H. Suzuki, A. A. Chinen, R. T. Chesser, J. ten Have, S. C. Donnellan, J. Austin, A. Frost, J.-P. Gonzalez, V. Herbreteau, F. Catzeflis, J. Soubrier, Y.-P. Fang, J. Robins, E. Matisoo-Smith, A. D. S. Bastos, I. Maryanto, M. H. Sinaga, C. Denys, R. A. Van Den Bussche, C. Conroy, K. Rowe, and A. Cooper. 2011. Multiple geographic origins of commensalism and complex dd history of Black Rats. PLoS ONE 6: e26357. https://doi.org/10.1371/journal.
agen’ K. N., and K. P. Ain. 2011. Bats of the Muller Range, Papua New Guinea. In Rapid Biological Assessments of the Nakanai Mountains and the Upper Strickland Basin: Surveying the Biodiversity of Papua New Guinea's Sublime Karst Environments, ed. S. J. Richards and B. G. Gamui, RAP Bulletin of Biological Assessment 60, pp. 222—234. Arlington: Conservation International.
Breed, W. G., S. Tan, C. M. Leigh, K. P. Aplin, K. Dvorakova- Hortova, and H. D. Moore. 2011. The morphology ofthe squirrel spermatozoon: a highly complex male gamete with a massive acrosome. Journal of Morphology 272: 883—889. https://doi.org/10.1002/jmor.10955
Bane M.I D. A. Steane, L. Joseph, D. K. Yeates, G. J. Jordan, D. Crayn, K. Aplin, D. J. Cantrill, L. G. Cook, M. D. Crisp, J. S. Keogh, J. Melville, C. Moritz, N. Porch, J. M. K. Sniderman, P. Sunnucks, and P. H. Weston. 2011. Decline of a biome: evolution, contraction, fragmentation, extinction and invasion of the Australian mesic zone biota. Journal of Biogeography 38): 1635-1656.
https://doi.org/10.1111/j.1365-269
pone.0026357
).2011.02535.x
David, B., J.-M. eneste. R. L. Whear, J.-J. Delannoy, M. Katherine, R. G. Gunn, C. Clarkson, H. Plisson, P. Lee, F. Petchey, C. Rowe, B. Barker, L. Lamb, W. Miller, S. Hoerlé, D. James, É. Boche, K. Aplin, I. J. McNiven, T. Richards, A. Fairbairn, and J. Matthews. 2011. Nawarla Gabarnmang, a 45,180 € 910 cal BP site in Jawoyn country, southwest Arnhem Land plateau. Australian Archaeology 73: 73—77. https://doi.org/10.1080/03122417.2011.11961928
Kambe, Y., T. Tanikawa, Y. Matsumoto, M. Tomozawa, K. P. Aplin, and H. Suzuki. 2011. Origin of agouti-melanistic polymorphism in wild Black Rats (Rattus rattus) inferred from Mcr gene SE d Spierice 28: 560—567. https://doi.org/10.2108/zsj.28.560
MCN I. J., B. David T Richards, K. Aplin, B. Asmussen, J. Mialanes, M. Leavesley, P. Faulkner, and S. Ulm. 2011. New direction in human colonisation ofthe Pacific: Lapita settlement of south coast New Guinea. Australian Archaeology 72: 1—6. https://doi.org/10.1080/03122417
O’Connor, S., A. Barham, K. Aplin, K. Dobney, A. Fairbairn, and M. Richards. 2011. The power of paradigms: examining the evidential basis for early to mid-Holocene pigs and pottery in Melanesia. Journal of Pacific Archaeology 2: 1—25.
2011.11690525
Rowe K. C., K. P. Aplin, P. R. Baverstock, and C. Moritz. 2011. Recent and rapid speciation with limited morphological disparity in ihe Eu Rattus. Systematic Biology 60: 188—203.
https oi.org/10.1093/sysbio/syq092
2012
McNiven, I. J., B. David, K. Aplin, J. Mialanes, B. Asmussen, S. Ulm, P. Faulkner, C. Rowe, and T. Richards. 2012. Terrestrial engagements by terminal Lapita maritime specialists on the southern Papuan coast. In Peopled Landscapes: Archaeological and Biogeographic Approaches to Landscapes, ed. S. Haberle and B. David. Terra Ausiraliss 34: 121-156. https://doi.org/10.22459/TA34.01.2012.05
McNiven, I. J., B. David. T. Richards, C. Rowe, M. Leavesley, J. Mialanes, S. P. Connaughton, B. Barker, K. Aplin, B. Asmussen, P. Faulkner, and S. Ulm. 2012. Response: Lapita on the south coast of Papua New Guinea: challenging new horizons in Pacific Archaeology. Australian Archaeology 75: 16—22.
Suzuki, H., and K. P. Aplin. 2012. Phylogeny and biogeography of the genus Mus in Eurasia. In Evolution of the House Mouse, ed. M. Macholán, S. J. E. Baird, P. Munclinger, and J. Piálek, pp. 35—64. Cambridge: Cambridge University Press.
81139044547.004
Westerman M., B. P. Kear, K. P. Aplin, R. W. Meredith, C. Emerling, and M. S. Springer. 2012. Phylogenetic relationships of living and recently extinct bandicoots based on nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 62: 97— rib
https://doi.org/10.1016/j.ymp
https://doi.org/10.1017/CBO97
2013
Aplin, K., and F. Ford. 2013. Murine rodents: late but highly successful invaders. In /nvasion Biology and Ecological Theory: Insights from a Continent in Transformation, ed. H. H. Prins and I. J. Gordon, pp. 196-240. Cambridge: Cambridge HBIVereit Pisos. https://doi.org/10.1017/CBO9781139565424.012
Conroy, C. JAB. Rowe, K. M. "Rows, P. L. Kamath, K. P. Aplin, L. Hui, D. K. James, C. Moritz, and J. L. Patton. 2013. Cryptic genetic diversity in Rattus of the San Francisco Bay region, California. Biological Invasions 15(4): 741—758. https://doi.org/10.1007/s10530-012-0323-9
Marin, J., S. C. Donnellan, S. B. Hedges, N. Puillandre, K. P. Aplin, P. Doughty, M. N. Hutchinson, A. Couloux, and N. Vidal. 2013. Hidden species diversity of Australian burrowing snakes (Ramphotyphlops). Biological Journal of the Linnean Society 110(2): 427-441. https://doi.org/10.1111/bij.12132
Petchey, F., S. Ulm, B. David, I. J. McNiven, B. Asmussen, H. Tomkins, N. Dolby, K. Aplin, T. Richards, C. Rowe, M. Leavesley, and H. Mandui. 2013. High-resolution radiocarbon dating of marine materials in archaeological contexts: radiocarbon marine reservoir variability between Anadara, Gafrarium, Batissa, Polymesoda spp. and Echinoidea at Caution Bay, Southern Coastal Papua New Guinea. Archaeological and dnehroporogtea! DeeHCeS sL i M 69—80. https://doi.org/10.1007/s12520-012 8-1
Shine, D. D. Wright, T Dehni K. Aplin, P. Hiscock, K. Parker, and R. Walton. 2013. Birriwilk rockshelter: a mid- to late Holocene site in Manilikarr Country, southwest Arnhem Land, Nocher Renton. Australian Archaeology 76: 69-78.
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Suzuki, H., M. Nunome, G. Kinoshita, K. P. Aplin, P. Vogel, A. P. Kryukov, M. L. Jin, S. H. Han, I. Maryanto, K. Tsuchiya, H. Ikeda, T. Shiroishi, H. Yonekawa, and K. Moriwaki. 2013. Evolutionary and dispersal history of Eurasian house mice Mus musculus clarified by more extensive geographic sampling of mitochondrial DNA. Heredity 111(5): 375—390. https://doi.org/10.103
Wright, D., P. Hiscock, and K. Aplin. 2013. Re-excavation of Dabangay, a mid-Holocene settlement site on Mabuyag in western Torres Strait. Queensland Archaeological Research 16: 15-32.
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Beck, R. M,, K. J. Travouillon, K. P. Aplin, H. Godthelp, and M. Archer. 2014. The osteology and systematics of the enigmatic Australian Oligo-Miocene metatherian Yalkaparidon (Yalkaparidontidae; Yalkaparidontia; ?Australidelphia; Marsupialia). Journal of Mammalian Evolution 21: 127—172. https://doi.org/10.1007/s10914-013-9236-3
Breed, W. G., C. M. Leigh, K. P. Aplin, A. A. Shahin, and N. L. Avenant. 2014, Morphological diversity and evolution of the spermatozoon in the mouse-related clade of rodents. Journal of Morphology 275: 540-547.
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O’ le S., G. Robertsi, and K. P. Aplin. 2014. Are osseous artefacts a window to perishable material culture? Implications of an unusually complex bone tool from the Late Pleistocene of East Timor. Journal of Human Evolution 67: 108—119. https://doi.org/10.1016/j.jhevol.2013.12.002
Robins, J. H., V. Tintinger, K. P. Aplin, M. Hingston, E. Matisoo- Smith, D. Penny, and S. D. Lavery. 2014. Phylogenetic species identification in Rattus highlights rapid radiation and morphological similarity of New Guinean species. PLoS ONE 9(5): e98002. https://doi.org/10.1371/journal.pone.0098002
Thomson, V., K. P. Aplin, A. Cooper, S. Hisheh, H. Suzuki, I. Maryanto, G. Yap, and S. C. Donnellan. 2014. Molecular genetic evidence for the place of origin ofthe Pacific Rat, Rattus exulans. PLoS ONE AE AS
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Aplin, K. P. 2015. Family Notoryctidae (marsupial moles). In Handbook of the Mammals of the World. Vol 5. Marsupials and Monotremes, ed. D. E. Wilson and R. A. Mittermeier, pp. 210-219. Barcelona: Lynx Edicions.
Aplin, K. P. 2015. Family Acrobatidae (feather-tailed gliders and feather-tailed possum). In Handbook of the Mammals of the World. Vol 5. Marsupials and Monotremes, ed. D. E. Wilson and R. A. Mittermeier, pp. 574—591. Barcelona: Lynx Edicions.
Aplin, K., K. N. Armstrong, and J. Novera. 2015. Mammals of Manus and Mussau Islands. In A Rapid Biodiversity Survey of Papua New Guineas Manus and Mussau Islands, ed. N. Whitmore, pp. 50-68. Goroka, Papua New Guinea: Wildlife Conservation Society Papua New Guinea Program.
Aplin, K. P., and J. S. Lamaris. 2015. Non-flying mammals. In A Rapid Biodiversity Assessment of Papua New Guinea s Hindenburg Wall Region, ed. S. J. Richards and N. Whitmore, pp. 131-165. Goroka, Papua New Guinea: Wildlife Conservation Society Papua New Guinea Program.
Aplin, K. P., S. G. Rhind, J. ten Have, and R. T. Chesser. 2015. Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a Topi Pag donis Zootaxa 4055(1): 1-73.
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Armstrong, K. N., K. P. Aplin, and J. S. Lamaris. 2015. Bats. In A Rapid Biodiversity Assessment of Papua New Guinea s Hindenburg Wall Region, ed. S. J. Richards and N. Whitmore, pp. 166-181. Goroka, Papua New Guinea: Wildlife Conservation Society Papua New Guinea Program.
Armstrong, K. N., J. Novera, and K. Aplin. 2015. Acoustic survey of the echolocating bats of Manus and Mussau Islands. In A Rapid Biodiversity Survey of Papua New Guinea s Manus and Mussau Islands, ed. N. Whitmore, pp. 69-85. Goroka, Papua New Guinea: Wildlife Conservation Society Papua New Guinea Program.
David, B., K. Aplin, F. Petchey, R. Skelly, J. Mialanes, H. Jones- Amin, J. Stanisic, B. Barker, and L. Lamb. 2015. Kumukumu 1, a hilltop site in the Aird Hills: implications for occupational trends and dynamics in the Kikori River delta, south coast of aoe INE Guinea. Quaternary International 385: 7—26. https://doi.org/10.1016/j.quaint.2014.06.058
aed, B.. J . Mialanes, F. Petchey, K. Aplin, J. M. Geneste, R. Skelly, and C. Rowe. 2015. Archaeological investigations at Waredaru and the origins of the Keipte Kuyumen clan estate, upper Kikori River, Papua New Guinea. PALEO. Revue d Archéologie Préhistorique 26: 33—57. https://doi.org/10.4000/paleo.2890
Gagan, M. K., L. K. Ayliffe, G. K. Smith, J. C. Hellstrom, H. Scott-Gagan, R. N. Drysdale, N. Anderson, B. W. Suwargadi, K. P., Aplin, J.-x. Zhao, C. W. Groves, W. S. Hantoro, and T. Djubiantono. 2015. Geoarchaeological finds below Liang Bua (Flores, Indonesia): a split-level cave system for Homo floresiensis?. Palaeogeography, Palaeoclimatology, i E 440: 533—550. https://doi.org/10.1016/j.palaeo.2015.09.021
Kagl, J. P, N. Whitmore, and K. P. Aplin. 2015. Traditional and local ecological knowledge. In A Rapid Biodiversity Assessment of Papua New Guinea 5 Hindenburg Wall Region, ed. S. J. Richards and N. Whitmore, pp. 3-13. Goroka, Papua New Guinea: Wildlife Conservation Society Papua New Guinea Program.
Maryan, B., M. Adams, and K. P. Aplin. 2015. Taxonomic resolution of the Aprasia repens species-group (Squamata: Pygopodidae) from the Geraldton Sandplains: a description of a new species and additional mainland records of A. clairae. Records of the Coser Australian Museum 30(1): 12-32. https://doi.org/10.18195/issn.0312
Maryan: B. I. Bosnien, M. Rcs. and K. P. Aplin. 2015. Molecular and morphological assessment of De/ma australis Kluge (Squamata: Pygopodidae), with a description of a new species from the biodiversity ‘hotspot’ of southwestern Western Australia. Zootaxa 3946: 301—330. https://doi.org/10.11646/zootaxa.3946.3.1
McDowell, M. C., D. Haouchar, K. P. Aplin, M. Bunce, A. Baynes, and G. Prideaux. 2015. Morphological and molecular evidence supports specific recognition of the recently extinct Bettongia anhydra (Marsupialia: Macropodidae). Journal of Mammalogy 96: 287—296. https://doi.org/10.1C
Motokawa, M., S. Shimoinaba, S. Kawada, and K. Aplin. 2015. Rediscovery of the holotype of Mus bowersii var. okinavensis Namiye, 1909 (Mammalia: Rodentia: Muridae). Bulletin of the National Museum of Nature and Science Series A (Zoology) 41: 131-136.
Richards, S. J., and K. Aplin. 2015. Herpetofauna of Manus and Mussau Islands. In A Rapid Biodiversity Survey of Papua New Guineas Manus and Mussau Islands, ed. N. Whitmore, pp. 31-37. Goroka, Papua New Guinea: Wildlife Conservation Society Papua New Guinea Program.
Woxvold, I., B. Ken, and K. P. Aplin. 2015. Birds. In A Rapid Biodiversity Assessment of Papua New Guinea s Hindenburg Wall Region, ed. S. J. Richards and N. Whitmore, pp. 103-130. Goroka, Papua New Guinea: Wildlife Conservation Society Papua New Guinea Program.
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2016
Alfano, N., J. Michaux, S. Morand, K. Aplin, K. Tsangaras, U. Lóber, P. H. Fabre, Y. Fitriana, G. Semiadi, Y. Ishida, K. M. Helgen, A. L. Roca, M. V. Eiden, and A. D. Greenwood. 2016. Endogenous gibbon ape leukemia virus identified in a rodent (Melomys burtoni subsp.) from Wallacea (Indonesia). Journal of Virology pM. 3107 BLEU, https://doi.org/10.1128/JVI 23-16
Jit K. T. Màn tud V. Attenbrow. 2016. Using a 3-stage burning categorization to assess post-depositional degradation of archaeofaunal assemblages: some observations based on multiple prehistoric sites in Australasia. Journal of Archaeological Science: Reports 7: 700—714.
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Aplin, K., C. Rowe, H. Peck, B. Asmussen, S. Ulm, P. Faulkner, and T. Richards. 2016. The natural setting of Caution Bay: climate, landforms, biota, and environmental zones. In Archaeological Research at Caution Bay, Papua New Guinea: Cultural, Linguistic and Environmental Setting, ed. T. Richards, B. David, K. Aplin, and I. J. McNiven, pp. 75-111. Caution Bay Studies in Archaeology, 1. Oxford: Archaeopress.
Beck, R. M. D., N. M. Warburton, M. Archer, S. J. Hand, and K. P. Aplin. 2016. Going underground: postcranial morphology ofthe early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians. Memoirs of Museum lacloria 74: 151-171. https://doi.org/10.24199/j.mmv.2016.74.14
David, B., H. J senini T. Richards, J. Mialanes, B. Asmussen, F. Petchey, K. Aplin, M. Leavesley, I. J. McNiven, C. Zetzmann, and C. Rowe. 2016. Ruisasi 1 and the earliest evidence of mass- produced ceramics in Caution Bay (Port Moresby Region), Papua New Guinea. Journal of Pacific Archaeology 7: 41—60.
Kear, B. P., K. P. Aplin, and M. Westerman. 2016. Bandicoot fossils and DNA elucidate lineage antiquity amongst xeric-adapted Australsstan DD CHINE Reports 6: 37537. https://doi.org/10.1038/srep3753
engin M. C. S. O’ Cohr: and K. Aplin. 2016. A >46,000- year-old kangaroo bone implement from Carpenter's Gap 1 (Kimberley, northwest Australia). Quaternary Science Reviews 154: 199-213.
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Richards TSB: Dav K. dos and I. J. McNiven. 2016. Archaeological Research at Caution Bay, Papua New Guinea: Cultural, Linguistic and Environmental Setting. Caution Bay Studies in Archaeology 1. Oxford: Archaeopress.
Richards, T. H., B. David, K. P. Aplin, and I. J. McNiven. 2016. The Caution Bay Project field and laboratory methods. In Archaeological Research at Caution Bay, Papua New Guinea: Cultural, Linguistic and Environmental Setting ed. T. Richards, B. David, K. Aplin, and I. J. McNiven, pp. 145-175. Caution Bay Studies in Archaeology, 1. Oxford: Archaeopress.
Richards, T., B. David, K. Aplin, I. J. McNiven, and M. Leavesley. 2016. Introduction to the Caution Bay Archaeology Project. In Archaeological Research at Caution Bay, Papua New Guinea: Cultural, Linguistic and Environmental Setting ed. T. Richards, B. David, K. Aplin, and I. J. McNiven, pp. 1—7. Caution Bay Studies in Archaeology, 1. Oxford: Archaeopress.
Timm, R. M., V. Weijola, K. P. Aplin, S. C. Donnellan, T. F. Flannery, V. Thomson, and R. H. Pine. 2016. A new species of Rattus (Rodentia: Muridae) from Manus Island, Papua New Guinea. Journal of Mammalogy 97. 1—18.
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Aplin, K., S. O'Connor, D. Bulbeck, P. J. Piper, B. Marwick, E. St Pierre, and F. Aziz. 2017. The Walandawe Tradition from Southeast Sulawesi and osseous artifact traditions in Island Southeast Asia. In Osseous Projectile Weaponry: Towards an Understanding of Pleistocene Cultural Variability, ed. M. C. Langley, pp. 137208, Dordrecht: Springer. https://doi.org/10.1007/978-94-024-0899-7_13
Aplin, K. P., and M. Opiang. 2017. Non-volant mammals (rodents and marsupials). In Biodiversity Assessment of the PNG LNG Upstream Project Area, Southern Highlands and Hela Provinces, Papua New Guinea, ed. S. J. Richards, pp. 141—208. Port Moresby: ExxonMobil PNG Limited.
Armstrong, K., and K. P. Aplin. 2017. Enhancing biological monitoring with genetic information. In Biodiversity Assessment of the PNG LNG Upstream Project Area, Southern Highlands and Hela Provinces, Papua New Guinea, ed. S. J. Richards, pp. 255-269. Port Moresby: ExxonMobil PNG Limited.
Barker, B., L. Lamb, J. J. Delannoy, B. David, R. Gunn, E. Chalmin, G. Castets, K. Aplin, B. Sadier, I. Moffat, and J. Mialanes. 2017. Archaeology of JSARN-124 site 3, central-western Arnhem Land: determining the age of the so-called ‘Genyornis’ painting. In The Archaeology of Rock Art in Western Arnhem Land, Australia, ed. B. David, P. S. C. Tagon, J.-J. Delannoy, and J.-M. Geneste, pp. 423—496. Canberra: The Australian National University Press. https://doi.org/10.22459/TA47.11.2017.15
David, B., J. J. Delannoy, R. Gunn, L. M. Brady, F. Petchey, J. Mialanes, E. Chalmin, J. M. Geneste, I. Moffat, K. Aplin, and M. Katherine. 2017. Determining the age of paintings at JSARN-113/23, Jawoyn Country, central-western Arnhem Land plateau. In The Archaeology of Rock Art in Western Arnhem Land, Australia, ed. B. David, P. S. C. Taçon, J.-J. Delannoy, and J.-M. Geneste, pp. 371—422. Canberra: The Australian National University Press. https://doi.org/10.22459/TA47.11.2017.14
David, B., J. J. Delannoy, R. Gunn, E. Chalmin, G. Castets, F Petchey, K. Aplin, M. O’Farrell, I. Moffat, J. Mialanes, and J. M. Geneste. 2017. Dating painted Panel El at Nawarla Gabarnmang, central-western Arnhem Land plateau. In The Archaeology of Rock Art in Western Arnhem Land, Australia, ed. B. David, P. S. C. Taçon, J.-J. Delannoy, and J.-M. Geneste, pp. 245—302. Canberra: The Australian National University Press. https://doi.org/10.22459/TA47.11.2017.11
Denys, C., P. J. Taylor, and K. P. Aplin. 2017. Family Muridae (true mice and rats, gerbils and relatives). In Handbook of the Mammals of the World. Vol. 7 Rodents II, ed. D. E. Wilson, T. E. Lacher Jr, and R. A. Mittermeier, pp. 536—886. Barcelona: Lynx Edicions.
Fabre, P.-H., A. H. Reeve, Y. Fitriana, K. P. Aplin, and K. M. Helgen. 2017. A new species of Halmaheramys (Rodentia: Muridae) from Bisa and Obi Islands (North Maluku Province, Indonesia). META of GP 99: 187—208. https://doi.org/10.10
Hawkins, S. S. o Connor. T. Maloney, M. Litster, S. Kealy, J. Fenner, K. Aplin, C. Boulanger, S. Brockwell, R. Willan, E. Piotto, and J. Louys. 2017. Oldest human occupation of Wallacea at Laili Cave, Timor-Leste, shows broad-spectrum foraging responses to late Pleistocene environments. Quaternary Science Reviews 171: 58-72. https://doi.org/10.1016/j.quascirev.2017.07.008
Jackson, S. M., C. P. Groves, P. J. S. Fleming, K. P. Aplin, M. D. B. Eldridge, A. Gonzalez, and K. M. Helgen. 2017. The wayward dog: is the Australian native dog or Dingo a distinct species? Zootaxa: Kr 201—224.
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Louys, J., S. Kealy, S. O'Connor, G. J. Price, S. Hawkins, K. Aplin, Y. Rizal, J. Zaim, Mahirta, D. A. Tanudirjo, W. D. Santoso, A. R. Hidayah, A. Trihascaryo, R. Wood, J. Bevitt, and T. Clark. 2017. Differential preservation of vertebrates in Southeast Asian caves. International Journal of Speleology 46: 379—408. https://doi.o 806X.46.3.2131
O’ COEAOR S., A. Barhánr K. Aplin, and T. Maloney. 2017. Cave stratigraphies and cave breccias: implications for sediment accumulation and removal models and interpreting the record of human occupation. Journal of Archaeological Science 7T: 143- 159.
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Woxvold, L, and K. P. Aplin. 2017. Camera trap monitoring of terrestrial birds and mammals: a pilot study. In Biodiversity Assessment of the PNG LNG Upstream Project Area, Southern Highlands and Hela Provinces, Papua New Guinea, ed. S. J. Richards, pp. 121-140. Port Moresby: ExxonMobil PNG Limited.
2018
Aplin, K., K. N. Armstrong, and I. Woxvold. 2018. Non-volant mammals. In /dentification Guide to Flora and Fauna of the Hides Range and the Agogo Range (Moro), Papua New Guinea, ed. S. J. Richards, pp. 117-144. Port Moresby: ExxonMobil PNG Limited.
Beaumont, P., S. O'Connor, M. Leclerc, and K. Aplin. 2018. Diversity in early New Guinea pottery traditions; north coast ceramics from Lachitu, Taora, Watinglo and Paleflatu. Journal of Pacific Archaeology 10(1): 15—32.
Chakma, N., N. J. Sarker, S. Belmain, S. U. Sarker, K. Aplin, and S. K. Sarker. 2018. New records of rodent species in Bangladesh: taxonomic studies from rodent outbreak areas in the Chittagong Hill Tracts. SH Journal of Zoology 46(2): 217—230. https://doi.org/10 9/bjz 0)2.39055
Eldridge, M. D. Ss. Potter, K. M. Helgen, M. H. Sinaga, K. P. Aplin, T. F. Flannery, and R. N. Johnson. 2018. Phylogenetic analysis of the tree-kangaroos (Dendrolagus) reveals multiple divergent lineages within New Guinea. Molecular Phylogenetics and Evolution 127: 589-599.
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Fabre, P Hi Y. S. Fitriana, G. Axeiniádi M. Pages, K. Aplin, N. Supriatna, and K. M. Helgen. 2018. New record of Melomys burtoni (Mammalia, Rodentia, Murinae) from Halmahera (North Moluccas, Indonesia): a review of Moluccan Melomys. Mammalia 82(3): 218—247. https://doi.org/10.1515/mammalia-2016-01 37
Hawkins, S., S. C. Samper Carro, J. Louys, K. Aplin, S. O'Connor, and Mahirta. 2018. Human palaeoecological interactions and owl roosting at Tron Bon Lei, Alor Island, eastern Indonesia. Journal of Coastal and Island Archaeology 13: 371—387 https://doi.org/10.1080/15564894.2017.1285834
Louys, J., M. Herrera, S. Hawkins, K. Aplin, C. Reepmeyer, F. Hopf, S. C. Donnellan, S. O'Connor, and D. A. Tanudirjo. 2018. Neolithic dispersal implications of murids from late Holocene archaeological and modern natural deposits in the Talaud Islands, northern Sulawesi. In The Archaeology of Sulawesi: Current Research on the Pleistocene to the Historic Period, ed. S. O'Connor, D. Bulbeck, and J. Meyer, pp. 223-242. Terra Australis 48. Canberra: ANU E Press. https://doi.org/10.22459/TA48.11.2018.14
Maloney. T. s. O'Connor, R. Wood, K. Aplin, and J. Balme. 2018. Carpenters Gap 1: a 47,000 year old record of indigenous adaption and innovation. Quaternary Science Reviews 191: 204—228.
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O'Connor, S., D. Bulbeck, P. J. Piper, F. Aziz, B. Marwick, F. Campos, J. Fenner, K. Aplin, F. Suryatman, T. Maloney, B Hakim, and R. Wood. 2018. The human occupation record of Gua Mo’o hono shelter, Towuti-Routa region of Southeastern Sulawesi. In The Archaeology of Sulawesi: Current Research on the Pleistocene to the Historic Period, ed. S. O'Connor, D. Bulbeck, and J. Meyer, pp. 117-152. Terra Australis 48. Canberra: ANU E Press. https://doi.org/10.22459/TA48.11.2018.09
Pahl, T., H. J. Nisletiat Y. Wang, A. S. Achmadi, K. C. Rowe, K. Aplin, and W. G. Breed. 2018. Sperm morphology of the Rattini—are the interspecific differences due to variation in intensity of intermale sperm competition? Reproduction, F ae and Devetonment 30(11): 1434—1442. https://doi.orc
Theden- Rinel. F,K.P. Hislon: K. Aplin, R. Grün, and M. R. Schurr. 2018. The chronology and environmental context of a cave deposit and associated faunal assemblage including megafauna teeth near Wee Jasper, southeastern Australia. The Holocene 28(9): 1467-1482. https://doi.org/10.1177/095968361 8777073
Thomson, V.,A. Wiewel, A. Chinen, I. Maryanto, M. H. Sinaga, R. How, K. Aplin, and H. Suzuki. 2018. A perspective for resolving the systematics of Rattus, the vertebrates with the most influence on himar welfare. Ad RE Mc 431—452.
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Balme, J., S. O'Connor, T. Maloney, D. Vannieuwenhuyse, K. Aplin, and I. E. Dilkes-Hall. 2019. Long-term occupation on the edge of the desert: Riwi Cave in the southern Kimberley, Western Austral, Archaeology in Oceania 54(1): 35—52. https://doi.org/10.1002/arco.5166
David, B., K. "fne H. Peck, R. Skelly, M. Leavesley, J. Mialanes, K. Szabó, B. Koppel, F. Petchey, T. Richards, and S. Ulm. 2019. Moiapu 3: settlement on Moiapu Hill at the very end of Lapita, Caution Bay hinterland. In Debating Lapita: Distribution, Chronology, Society and Subsistence, ed. S. Bedford and M. Spriggs, pp. 61-88. Terre Fur 52. Canberra: ANU Press.
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2020
Armstrong, K. N., K. P. Aplin, and M. Motokawa. 2020. A new species of extinct False Vampire Bat (Megadermatidae: Macroderma) from the Kimberley Region of Western Australia.
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Brockwell, S., and K. P. Aplin. 2020. Fauna on the floodplains: late Holocene culture and landscape on the sub-coastal plains of northern Australia. In Papers in Honour of Ken Aplin, ed. Julien Louys, Sue O'Connor, and Kristofer M. Helgen. Records of the Australian Museum 72(5): 225—236. https://doi.org/10.3853/j.2201-4349.72.2020.1728
Kealy, S., S. C. Donnellan, K. J. Mitchell, M. Herrera, K. Aplin, S. O'Connor, and J. Louys. 2020. Phylogenetic relationships of the cuscuses (Diprotodontia: Phalangeridae) of Island Southeast Asia and Melanesia based on the mitochondrial ND2 gene. Australian Mammalogy 42(3): 266—276. https://doi.org/10.1071/AM18050
Louys, J., M. B. Herrera, V. A. Thomson, A. S. Wiewel, S. C. Donnellan, S. O'Connor, and K. P. Aplin. 2020. Expanding population edge craniometrics and genetics provide insights into dispersal of commensal rats through Nusa Tenggara, Indonesia. In Papers in Honour of Ken Aplin, ed. Julien Louys, Sue O’Connor, and Kristofer M. Helgen. Records of the Australian Museum E PSI SUE, https://doi.org/10.3853/j.2201-4349.72.2020.1730
Miokiswicz J. I aJ. Louys, R. M. D. Beck, P. Mahoney, K. Aplin, and S. O'Connor. 2020. Island rule and bone metabolism in fossil murines from Timor. Biological Journal of the Linnean Society 129(3): T 386, https://doi.org/10.1093/biolinnean/blz197
Roberts, P., J. Coire J. Zech, C. Shipton, S. Kealy, S. Samper Carro, S. Hawkins, C. Boulanger, S. Marzo, B. Fiedler, N. Boivin, Mahirta, K. Aplin, and S. O'Connor. 2020. Isotopic evidence for initial coastal colonization and subsequent diversification in the human occupation of Wallacea. Nature Communications 11(1): 1-11.
https://doi.org/10.1038/s41467-020-15969-4
ACKNOWLEDGEMENTS. This introduction and remembrance of Ken Aplin is based on the eulogy given by Kris Helgen on 21 January 2019, at Ken’s funeral at Pappinbarra, New South Wales.
We thank Ken’s family, including his father, Kenneth Sr, his sisters Sue, Jenny, and Val, and his older children Nick, Lucy, Felix, and Charlotte, for their kindness and fellowship. We especially thank Ken’s wife and partner, Dr Angela Frost, and their daughter Sophie, for sharing Ken with us, and for so many other things.
As editors, we thank the authors who have contributed these papers to this volume in celebration of Ken, and the many reviewers who assessed and improved our contributions. We also thank Rebecca Johnson, Kim McKay, Cameron Slatyer, Sandy Ingleby, Anja Divljan, Harry Parnaby, and Lauren Helgen for support.
We are deeply grateful to Dr Shane McEvey, editor of the Records of the Australian Museum, for the opportunity to edit this volume, and for his remarkable support in producing it on our behalf.
Records of the Australian Museum (2020) vol. 72, issue no. 5, pp. 161—174 https://doi.org/10.3853/j.2201-4349.72.2020.1732
Records of the Australian Museum
a peer-reviewed open-access journal
published by the Australian Museum, Sydney communicating knowledge derived from our collections ISSN 0067-1975 (print), 2201-4349 (online)
A New Species of Extinct False Vampire Bat (Megadermatidae: Macroderma) from the Kimberley Region of Western Australia
KYLE N. ARMsTRONG??($, KEN APLIN!*7@, AND MASAHARU MoTOKAwa!
' The Kyoto University Museum, Yoshida Honmachi, Sakyo-ku, Kyoto, 606-8501, Japan ? School of Biological Sciences, The University of Adelaide SA 5005, Australia ? South Australian Museum, North Terrace, Adelaide SA 5000, Australia
^ Australian Museum Research Associate, 1 William Street, Sydney NSW 2010, Australia
ABSTRACT. A new species of False Vampire Bat (Megadermatidae), Macroderma handae sp. nov., is described from dental, dentary and maxillary fragments recovered from limestone deposits at Dingo Gap, Oscar Range, in the Kimberley region of Western Australia. This material is likely to be of Pliocene age, or early Pleistocene, based on biocorrelation within the same sample. The absence of the P? indicates that it is more derived than Miocene taxa including M. malugara and M. godthelpi, but its phylogenetic position relative to M. koppa could not be determined. It appears to be slightly smaller than M. gigas and M. koppa based on the size of M! and M.. It can be distinguished from M. gigas by the lesser degree of fenestration in the maxilla; and from all other species of Macroderma by the shape ofthe protofossa of the M!, plus the M, protoconid relatively high and of proportionally greater area within the trigonid. Other material collected, but not identified completely or described, includes several lower canines from a species of emballonurid, and a dentary with M, , representing a vespertilionid bat. Given the wear striations observed on the M; of the newly-described Macroderma species, we suggest that it was a predator of small vertebrates, including possibly the chiropteran co-inhabitants of the cave. This new species of Macroderma is the sixth species recognized in the genus so far, and the second from the Pliocene.
Introduction Hand & Sigé, 2018). Early megadermatid lineages are
represented by modern extant taxa in the genera Lavia
The family Megadermatidae (False Vampire Bats) has a long history that began in the mid-Eocene with its divergence from the Craseonycteridae c. 44—43 Ma, based on molecular dating methods (9596 credibility interval 47—39 Ma; Teeling et al., 2005; Foley et al., 2015). Until recently, the oldest known megadermatid fossil was considered to be Necromantis adichaster Weithofer, 1887, represented in the Quercy Phosphorites Formation, France, but this genus 1s now accepted to be part of a distinct family (Necromantidae; Sigé, 2011; Ravel et al., 2016;
and Cardioderma, based on their inferred phylogenetic position (Hand, 1985; but see Kaňuch et al., 2015). The oldest megadermatid fossils, however, are: Saharaderma pseudovampyrus Gunnell et al., 2008 from early Oligocene deposits in Egypt (33.9—28.4 Ma), which shows similarities to Cardioderma and Lavia, and with which it may form a distinct African clade (Gunnell et al., 2008); and Megaderma lopezae Sevilla, 1990 from early Oligocene deposits in Spain. The remaining eight described Afro-European species of extinct Megaderma are represented in deposits that range
Keywords: Macroderma; Megaderma; Ghost Bat; False Vampire Bat; new species; Pliocene Taxonomic registration: urn:lsid:zoobank.org:pub:D252F DFE-7C93-4E13-8DC7-FFB585ACC16F
Corresponding author: Kyle N. Armstrong kyle.armstrong@adelaide.edu.au
Received: 3 February 2020 Accepted: 20 August 2020 Published: 25 November 2020 (in print and online simultaneously) Publisher: The Australian Museum, Sydney, Australia (a statutory authority of, and principally funded by, the NSW State Government)
Citation: Armstrong, Kyle N., Ken Aplin, and Masaharu Motokawa. 2020. A new species of extinct False Vampire Bat (Megadermatidae: Macroderma) from the Kimberley Region of Western Australia. In Papers in Honour of Ken Aplin, ed. Julien Louys, Sue O’Connor, and Kristofer M. Helgen. Records of the Australian Museum 72(5): 161—174. https://doi.org/10.3853/j.2201-4349.72.2020.1732
Copyright: © 2020 Armstrong, Aplin, Motokawa. This is an open access article licensed under a Creative Commons Attribution 4.0 International License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any
medium, provided the original authors and source are credited.
COL
T Ken Aplin 1958-2019, deceased
162 Records of the Australian Museum (2020) Vol. 72
Figure 1. Location of Dingo Gap (star) in the Kimberley region of Western Australia,
plus Riversleigh World Heritage Area (cross), Wellington Caves in New South Wales 0 a
(circle; type locality of Macroderma koppa), and various sites where subfossil and guano of M. gigas have been found (triangles) (information from Cook, 1960; Bridge, 1975; Molnar et al., 1984; Hand, 1996; Mahoney ef al., 2008; and Ken Aplin unpublished
data from islands of northwestern Western Australia).
in age from the early Miocene (e.g., Megaderma brailloni Sigé, 1968 from the Aquitanian) to the Pleistocene (e.g., Megaderma watwat Bate, 1937) (reviewed in Sigé, 1976; Sevilla, 1990; Ziegler, 1993).
Australia has excellent representation of megadermatid fossil taxa, beginning from the mid-Cenozoic and extending to subfossil recent material (Molnar et al., 1984; Hand, 1996). Most have been discovered in the freshwater limestone deposits of Riversleigh World Heritage Area, northwestern Queensland, which has a rich diversity of bat species from the families Mystacinidae (Hand et al., 1998), Emballonuridae (Archer et al., 2006; King, 2013), Rhinonycteridae (Sigé et al., 1982; Hand, 1997a; Hand & Archer, 2005), Hipposideridae (Hand, 1997b; Hand, 1998a, 1998b), Molossidae (Hand, 1990; Hand et al., 1997), and Vespertilionidae (Menu et al., 2002).
The genus Megaderma is thought to have entered Australia after the middle Miocene, and the small-sized Megaderma richardsi from the early Pleistocene Rackham's Roost Site at Riversleigh is its only known representative in Australia (Hand, 1995; Woodhead et al., 2016). Four extinct Australian megadermatid taxa have been referred to the endemic genus Macroderma—M. godthelpi Hand, 1985 from the early Miocene Microsite and middle Miocene Gag Site, Riversleigh; M. malugara Hand, 1996 from the middle Miocene Gotham City Site, Riversleigh; an unnamed species of Macroderma from the middle Miocene Henk's Hollow Site, Riversleigh (Hand, 1996); and M. koppa Hand, Dawson & Augee, 1988 from the Pliocene deposits of Big Sink, Wellington Caves, New South Wales (Hand et al., 1988). The
remaining two extinct megadermatid taxa from Australia have not been given a formal binomial name—Dwornamor Variant from the middle Miocene Gag Site, Riversleigh (Hand, 1985); and Megadermatidae indet. from the middle Miocene Henk's Hollow Site, Riversleigh (Hand, 1996).
The extant Macroderma gigas (Dobson, 1880) is currently distributed across northern Australia, from the Pilbara and Kimberley regions of Western Australia, through the Top End of the Northern Territory and part of the Gulf Coastal and Mt Isa Inlier bioregions of the Northern Territory and northwestern Queensland, to Cape York, Queensland (Worthington Wilmer et al., 1999; Churchill, 2008). It contracted from areas further south in the Holocene (Molnar et al., 1984), and has declined further since the arrival of Europeans (Churchill & Helman, 1990; Churchill, 2008; Woinarski et al., 2014; Augusteyn et al., 2018; Armstrong et al., 2019). This taxon is also represented in the early Pleistocene deposit of Rackham's Roost, Riversleigh (Hand, 1996; Woodhead et al., 2016), as well as many sites of Pleistocene and Holocene age around Australia (Molnar et al., 1984). In Western Australia, fossil and subfossil bat material has been discovered in very few localities, though M. gigas is a conspicuous presence in numerous limestone caves in the south-west corner (reviews in Cook, 1960; Bridge, 1975; Baynes etal., 1975; Molnar et al., 1984; Armstrong & Anstee, 2000), and few of these caves are now used by bats of any species (Armstrong et al., 2005). Megadermatid fossils have also been discovered further north on Barrow Island and the Monte Bello Islands off the Pilbara coast (Ken Aplin, unpublished observations).
Armstrong et al.: New extinct species of Kimberley Macroderma 163
More recently, a limestone deposit from Dingo Gap in the Kimberley region, north-west of Fitzroy Crossing (Fig. 1), has produced material from a range of fossil mammals, which includes at least three species of bat. One of these is clearly a megadermatid, which is described here as a new species. The other bat species are not sufficiently well represented for identification or formal description, but they do provide context for the occurrence of the megadermatid bones and teeth.
Methods
Scanning electron micrographs were taken with a Jeol JSM6060B microscope. Holotype and paratype material was examined and illustrated in comparison with a specimen of M. gigas from the CSIRO Australian National Wildlife Collection (ANWC), Canberra (CM568, male, collected from Mt Etna, Queensland), as well as material in the Western Australian Museum (WAM: three dentaries from M. gigas specimens M3415, M18284 and M18575; all from the Pilbara region of Western Australia). Descriptions are made in comparison with information in Hand (1985, 1995, 1996) and Hand et al. (1988). Measurements were made from SEM images using the software ImageJ (Rasband, 1997—2005; Abramoff et al., 2004). Measurements of the newly described species made for direct comparison with M. gigas correspond to a subset of those in Hand (1985) and are numbered accordingly (Fig. 2). Additional measurements made for descriptive purposes are indicated by letters (Table 1). Higher level systematics follow Simmons & Cirranello (2020). Anatomical terminology follows Hand (1985), Hand et al. (1988), and Hand (1996).
Systematics
Chiroptera Blumenbach, 1779
Yinpterochiroptera Springer, Teeling, Madsen, Stanhope & de Jong, 2001
Rhinolophoidea Gray, 1825 Megadermatidae H. Allen, 1864
Macroderma Miller, 1906
Macroderma handae sp. nov. Aplin and Armstrong
1:lsid:zoobank.ora:act:018A 744D-3AE6-44C0-988E-018C963EI
Figs 3-8
Holotype. Fragment of left dentary containing a mostly intact M,, broken P,, M, and M,, and alveoli of single-rooted P, and C, (WAM 2020.4.1; Figs 3A,B and 4A,C,E,G). Paratypes. A second fragment of left dentary with alveoli of incisors, C,, P, and P,, and first two molars (WAM 2020.4.2; Fig. 3D,E); athird fragment of left dentary containing a worn M, and one alveolus of M, (WAM 2020.4.3; Fig. 3C); palatal fragment of left maxilla with lingual alveoli of P^ and M' (WAM 2020.4.4; Fig. 5B,C); fragment of right maxilla with alveoli of C! and P^ (WAM 2020.4.5; Fig. 5A); right M! (WAM
2020.4.6; Fig. 6A,C); right V? fragment (WAM 2020 4.10; Fig. 6E); anterior portion of right C! (WAM 2020.4.7; Fig. 7A); right C! with broken paracone (principal cusp, sensu Hand, 1985; WAM 2020.4.9; Fig. 7C,D); left P (WAM 2020.4.8; Fig. 8A,B,D); left M, in poor condition (WAM 2020.4.11; Fig. 4D); left P^ with damaged paracone (principal cusp; WAM 2020.4.12; Fig. SE-H). All type material is lodged in the Western Australian Museum.
Type locality, lithology, and age. Material was collected from a cemented accumulation of bone material that formed on the floor of a cave in a carbonate-rich stratigraphic sequence at Dingo Gap, Oscar Range, Kimberley region, Western Australia (17?40'S 125?13'E, Fig. 1). The location is part of the marginal reef slope and basinal facies of the northern face of the Oscar Range (Stephens & Sumner, 2003). This range forms the northern edge of the Canning Basin, and is the remnant of an Upper Devonian marine reef complex.
The bone accumulation was in a hard limestone matrix and consisted of teeth and small bone fragments of mammals, particularly rodents (Muridae: Hydromyini (sensu Smissen & Rowe, 2018); Rattus was absent). Further details of the fauna in this collection are not yet available. It is more likely to be an accumulation from a cave floor beneath a megadermatid bat roost site rather than a pellet accumulation from an owl given that larger jaw fragments were absent. Dental material from other bats was also present, including an unknown species of bat (Fig. 9A—D), canines from an emballonurid (probably Taphozous sp.; Fig. 9E-L), and a lower row of molars from an unidentified vespertilionid (Fig. 9M,N). Given the absence of Rattus, which is thought to have reached Australia by at least the mid-Pleistocene (Rowe et al., 2019), the material is aged tentatively as Pliocene or early Pleistocene.
Diagnosis. Referred to the genus Macroderma Miller, 1906 on the basis of the large size of the M'? (within the lower part of the size range of M. gigas and M. koppa; Table 1; cf. Hand, 1995: 52), the M! with elongated heel, and markedly lingually displaced mesostyle (cf. Megaderma richardsi; Hand, 1995: 66); M, , paracristid (sensu Hand, 1995, 1996; = protocristid sensu Hand, 1985, who used both terms) longer than metacristid; M, ; reduced metaconid contribution to the cristid obliqua; M,_; robust and continuous anterior, labial (= buccal) and posterior cingula (see Hand, 1996: 373).
Compared with Macroderma gigas—Maxilla fenestrated (Fig. 5B,C), but not to the degree seen in M. gigas (cf. Hand, 1985: 31); anterior part of dentary thickened, though relatively gracile compared with that of M. gigas (dentary depth below M, protoconid less in M. handae; Table 1; Fig. 3A,F); most molar measurements smaller than the average for M. gigas, or within the lower part of the size range (Table 1); the shape of the M! protofossa (whose edges are defined by the preprotocrista and postprotocrista) is rounded rather than triangular (Fig. 6A—D); M, paraconid lower, and protruding less anteriorly past the protoconid (trigonid less expanded anteriorly than in M. gigas); M; protoconid relatively high and of proportionally greater area within the trigonid (more than half in occlusal view (Fig. 4A,B); and M, talonid proportionally larger with respect to the trigonid (Fig. 4A,B). No protostyle cusp on P^, which is obvious in M. gigas (Fig. 8E,F).
Compared with M. koppa (see Hand et al., 1988: 344—346)—Anterior upper tooth row relatively shorter in M. handae, alveoli of C! and P^ indicating overlap of crowns
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Table 1. Measurements (mm; Fig. 2) of the holotype dentary and M, (WAM 2020.4.1), and the paratypes M' (WAM 2020.4.6) and C! (WAM 2020.4.7) of Macroderma handae sp. nov., in comparison with M. gigas and M. koppa (values and character numbers are from Hand, 1985: 23,25; Hand et al., 1988: 349; mean and range in parentheses; RR indicates measurements from M. gigas in Rackham's Roost, see Hand, 1996: 370; letters in the first column represent measurements made in the present study only; * measurement from paratype WAM 2020.4.2).
holotype dentary and M, M. handae M. gigas M. koppa 3 Dentary depth below M, protoconid 3.5, 3.42* 3.92 (3.40—4.90) RR: 3.45 4.2 (4.445) 10 M, length (sum measurements 14 + 15) 3.21 3.78 (3.41—4.17) RR: 3.27 4.2 (3.9-4.1) 14 M, trigonid length 1.73 2.41 (1.91-2.79) RR: 2.10 2.5 (2.3-2.5) 15 M; talonid length 1.48 1.41 (1.00-1.88) RR: 1.19 1.6 (1.3-1.6) 2] M, trigonid width 2.36 2.38 (2.05-2.68) 2.8 (2.4-2.6) 22 M, talonid width 2.16 2.31 (1.86—2.85) 2.6 (2.2-2.5) 27 M, paracristid length 1.44 1.72 (1.38-1.92) — 28 M, metacristid length 1.04 1.25 (0.98-1.65) — A M, protoconid height (not illustrated) 3.19 — — B Mental foramen width (not illustrated) 0.53, 0.55 — — paratypes M! and C! M. handae M. gigas M. koppa 14 M! labial (buccal) length 3.53 3.93 (3.36-4.40) RR: 3.36, 3.52. 4.1(4.0-4.2) 18 M! lingual length 3.13 4.24 (3.60—4.76) RR: 3.59, 3.85 4.0 21 M! width 3.95 4.15 (3.65—4.63) RR: 3.433, 3.94 444 (4.1—4.3) 25 M! metacone apex to metastyle 2.15 2.73 (2.36-2.88) — 28 M! paracone to heel 2.43 3.20 (2.29-3.66) — 30 M! heel inflexions 1.49 2.34 (1.84-3.54) — 32 M! length through protocone 1.70 2.44 (2.08-2.90) — C M! protofossa width 1.20 — — D M! heel width 1.37 — — E C! height (not illustrated) 4.29 — — Left M2
Figure 2. Dental measurements taken from the left M, and the right M', based on Hand (1985).
(Fig. 5A; cf. Hand et al., 1988: 345, fig. 2b,c); the shape of the M! protofossa (with edges defined by the preprotocrista and postprotocrista) 1s rounded rather than triangular; molar measurements smaller than the values for M. koppa (Table 1; cf. Hand et al., 1988: 349); anterior part of dentary relatively gracile compared with that of M. koppa (dentary depth below M, protoconid less in M. handae; Table 1); M, paraconid relatively low, and protruding less anteriorly past the protoconid due to anterior compression of the trigonid (Fig. 4C,E; cf. Hand et al., 1988: 345, fig. 2a); M, protoconid relatively high and of proportionally greater area within the trigonid (more than half in occlusal view; Fig. 4A); entoconid
smaller than hypoconulid (Fig. 4E,G; cf. Hand et al., 1988: 345, fig. 2a); the P, is of a similar shape in both species (Fig. 8A,B,D; cf. Hand et al., 1988: 345, fig. 2a).
Compared with M. malugara Hand, 1996—P? absent in M. handae; slightly smaller size of M! and M, (Table 1; cf. Hand, 1996: 368); the shape of the M! protofossa (whose edges are defined by the preprotocrista and postprotocrista) is rounded rather than triangular; M, paraconid relatively low, and protruding less anteriorly past the protoconid due to anterior compression of the trigonid (Fig. 4A,C,E; cf. Hand, 1996: 366—367, pl. 48k—m); M, protoconid relatively high and of proportionally greater area of the trigonid (more than
Armstrong ef al.: New extinct species of Kimberley Macroderma 165
Figure 3. Scanning electron micrographs of holotype and paratype material of Macroderma handae sp. nov. (A) lateral view of the left dentary of holotype WAM 2020.4.1 with mostly intact M», broken P,, M, and M,, and alveoli of single-rooted P, and C,; (B) occlusal view of the holotype WAM 2020.4.1 anterior to the M,; (C) occlusal view of a fragment of left dentary, paratype WAM 2020.4.3; (D, E) lateral and occlusal view of a fragment of left dentary, paratype WAM 2020.4.2; (F) digital photograph of the left dentary of M. gigas WAM M18284. Scale bars 1 mm.
166 Records of the Australian Museum (2020) Vol. 72
Figure 4. Scanning electron micrographs of holotype and paratype material of Macroderma handae sp. nov. (A, C, E, G) occlusal, lingual, labial, and labial-oblique views of the left M, from the holotype WAM 2020.4.1; (B, D, F H) corresponding views of the left M, of M. gigas ANWC CM568; (7) occlusal view of left M3, paratype WAM 2020.4.11; (J) left M, of M. gigas ANWC CM568. Scale bars 1 mm.
Armstrong ef al.: New extinct species of Kimberley Macroderma 167
Figure 5. Scanning electron micrographs of paratype material of Macroderma handae sp. nov. (A) fragment of the right maxilla with alveoli of the C! and P^, paratype WAM 2020.4.5; (B) palatal fragment of left maxilla with lingual alveoli of P^ and M!, paratype WAM 2020.4.4; (C) detail of the blood vessel fenestrations in paratype WAM 2020.4.4; (D) probable wear striations on the M;, paratype WAM 2020.4.11; (E) wear striations from M. gigas ANWC CM568. Scale bars 1 mm, except where indicated otherwise.
168 Records of the Australian Museum (2020) Vol. 72
Figure 6. Scanning electron micrographs of paratype material of Macroderma handae sp. nov. (A, C) occlusal-oblique views of a right Mi, paratype WAM 2020.6; (B, D) corresponding views of the right M! of M. gigas ANWC CM568; (E) occlusal view of a fragment of a right MP, paratype WAM 2020.44.10; (F) corresponding view of the right M? of M. gigas ANWC CM568. Scale bars 1 mm.
Armstrong ef al.: New extinct species of Kimberley Macroderma 169
Figure 7. Scanning electron micrographs of paratype material of Macroderma handae sp. nov. (A) anterior half of a right C!, paratype WAM 2020.4.7; (B) lingual view of a right C! of M. gigas ANWC CM568; (C, D) labial and lingual views of a right C! with a damaged paracone, paratype WAM 2020.4.9. Scale bars 1 mm.
half in occlusal view; Fig. 4A; cf. Hand, 1996: 366—367, pl. 48m); greater development of M5 hypoconulid (Fig. 4A; cf. Hand, 1996: 366—367, pl. 48m).
Compared with M. godthelpi Hand, 1985—C' and M! and M; slightly larger in size in M. handae, and M, with greater protoconid height (Table 1; cf. measurements in Hand, 1985: 8—9; see also Sigé et al., 1982 for measurement key); taller and more robust C! (Table 1E; Fig. 7A,C,D; cf. Hand, 1985: 9.12, fig. 5a,b); loss of P?; proportionally greater contribution of the cingulum to the height of the P, (cf. Hand, 1985: 13, fig. 6c); M, paraconid relatively low, and protruding less anteriorly past the protoconid due to anterior compression of the trigonid (Fig. 4A,C,E; cf. Hand, 1985: 11, fig. 4a,b,c); M, protoconid relatively high and of proportionally greater area of the trigonid (more than half in occlusal view; Fig. 4A; cf. Hand, 1985: 11, fig. 4c).
Description. The anterior part of the dentary is thickened, though relatively gracile and shallower in depth compared to M. koppa and M. gigas, with likely two lower incisors
per side (paratype WAM 2020.4.2; anterior detail not shown in Fig. 3A,B,D,E). Two premolars are present—P, and P,, in addition to the M, (Fig. 3A,B), and the M, (Fig. 4I).
There is marked extension posterolingually of the C,, similar to M. gigas (Fig. 7A—D). The P, has a proportionally large cingulum, as can be seen in occlusal view, which gives the tooth the appearance of a “witches hat" when viewed from either the labial or lingual side (Fig. 8A,B,D).
The M, is shorter than, or equal in length to, the tall- crowned M, (Fig. 3A). The paracristid of the M, is longer than the metacristid (Fig. 4A). There is relatively little contribution of the M, metaconid to the cristid obliqua (Fig. 4A). The M, hypoconulid is situated posteriorly (Fig. 4A). The anterior, labial, and posterior cingula are robust and continuous (Fig. 4A,E,G). There is no development of the entostylid (Fig. 4A).
The maxilla is rugose and fenestrated, with grooves of blood vessels along the surface (Fig. 5B,C). The condition of the infraorbital foramen (a key feature separating M.
170 Records of the Australian Museum (2020) Vol. 72
Figure 8. Scanning electron micrographs of paratype material of Macroderma handae sp. nov. (A, B, D) lingual, labial and occlusal views of a left P, paratype WAM 2020.4.8; (C) labial view of the left P, of M. gigas ANWC CM568; (E, G, H) occlusal, lingual-oblique, and posterior views of a damaged left P^, paratype WAM 2020.4.12; (F) occlusal view ofa left P^ of M. gigas ANWC CM568. Scale bars 1 mm.
Armstrong ef al.: New extinct species of Kimberley Macroderma 171
F
Figure 9. Scanning electron micrographs of other unidentified and undescribed bat material recovered from the same deposit at Dingo Gap. (4—D) WAM 2020.4.13; (E-G) right C, of an emballonurid, WAM 2020.4.14; (H) right C, of an emballonurid, WAM 2020.4.15; (4, J) left C; of an emballonurid, WAM 2020.4.16; (K, L) left C, of an emballonurid, WAM 2020.4.17; (M, N) lingual and occlusal views of a fragment of dentary of a vespertilionid containing M,—M; (M, is on the right in both views), WAM 2020.4.18. Scale bars 1 mm.
172 Records of the Australian Museum (2020) Vol. 72
Megadermatidae indet. Megaderma spp. Dwornamor Variant Macroderma godthelpi Macroderma sp.
Macroderma malugara
n
.. Macroderma handae sp. nov.
Macroderma koppa
* * *
5 W l Macroderma gigas
Figure 10. Inferred relative phylogenetic position of Macroderma handae sp. nov. based on observable synapomorphic features (modified
after Hand, 1996; numbers indicate the development of potential apomorphic character states, as detailed in that reference).
koppa |two foramina] and M. gigas [one foramen]; Fig. 5A) cannot be observed.
The P? is absent, as indicated by the absence of an alveolus between those of the canine and P^ (paratype WAM 2020.4.5; Fig. 5A). The alveolus of C! and anterobuccal/ anterolabial alveolus of P^ indicate that the crowns of these teeth overlapped in the tooth row (Fig. 5A). The heel of the P" is broad, and the posterior edge is at right angles to the paracone (it is angled close to 45? lingually in M. gigas; Fig. 8E-H). There is no protostyle cusp, which is obvious in M. gigas (Fig. SE,F).
The M! has a broad labial (buccal, sensu Hand, 1996) shelf, though narrower than that of M. gigas (Fig. 6A,B), and a markedly lingually displaced mesostyle (cf. Megaderma richardsi, Hand, 1995). The preprotocrista and postprotocrista are curved, giving the protofossa a rounded shape, which contrasts with the more triangular form of other Macroderma species (Hand et al., 1988: 345, fig. 2c; Hand, 1985: 10, fig. 3c, 1996: 366—367, pl. 48d), and also Megaderma richardsi (Hand, 1995: pl. 1b,c). Both the M! and M? have tall crowns, and appear to be slightly compressed anteroposteriorly relative to Macroderma gigas (Fig. 6A—F).
Unidirectional wear striations are observable on the left M5, which resemble those found on the teeth of the predatory M. gigas that crush the bones of prey (Fig. 5D,E).
Etymology. Named in honour of Professor Suzanne (“Sue”) J. Hand ofthe University of New South Wales, in recognition of her previous extensive work on fossils of this family, and her extraordinary, sustained, and ongoing work on fossils that has helped piece together the rich history of the Australasian mammal fauna.
Discussion
Phylogenetic relationships
The phylogenetic position of Macroderma handae relative to most megadermatids can be estimated based on the presence of various synapomorphies that characterize subclades within the family (character sets 1—5 listed in Hand, 1996: 373) (Fig. 10). It displays the following apomorphic conditions: (a) Characterizing it as part of the Megaderma- Macroderma clade: M, shorter than or equal in length to M.. (b) Distinguishing it from the Megaderma clade: M! with elongated heel, and markedly lingually displaced mesostyle (cf. Megaderma richardsi, Hand, 1995); in the M5: the paracristid longer than metacristid, reduced metaconid contribution to the cristid obliqua; robust, continuous anterior, labial and posterior cingula. (c) Distinguishing it from Macroderma godthelpi: large-sized, tall-crowned teeth; M, with robust and broad anterior cingulum. (d) Distinguishing it from M. malugara: P? absent; C, markedly posterolingually-extended; M, » larger and more posteriorly- situated hypoconulid; and preentocristid further reduced. The phylogenetic position of M. handae relative to M. koppa and M. gigas could not, however, be determined unambiguously based on the material from Dingo Gap because the condition of the infraorbital foramen (one or two foramina) and some other diagnostic features could not be observed.
Australian Pliocene megadermatid diversity
The new species M. handae represents the second Pliocene species of Macroderma discovered to date, together with
Armstrong et al.: New extinct species of Kimberley Macroderma 173
M. koppa. The age of the Big Sink Site of Wellington Caves in New South Wales has also been estimated as Pliocene, though it has not been dated other than on the basis of biocorrelation with better-dated faunas (reviews in Hand, 1996; Dawson et al., 1999), and the inferred plesiomorphic condition of M. koppa (Dawson et al., 1999: 284). Both sites lack Rattus material, though they have representatives of the Old Endemic murid radiation (Hydromyini, sensu Smissen & Rowe, 2018), so their likely age is at least somewhere between the first Australian murid radiations and the invasion of Rattus (Aplin, 2006; Rowe et al., 2019). The species M. handae and M. koppa might have been contemporaneous, or alternatively they could have arisen at slightly different ages sometime from the late Miocene to early Pliocene. While M. handae appears slightly smaller on the basis of a few molar measurements, it is not markedly so. Thus, it might have been an earlier or allopatric taxon. A proposed common name for M. handae is the Kimberley False Vampire Bat.
Chiropteran assemblage
Several other bat species were recovered from the same assemblage that contained M. handae (Fig. 9). The lack of molars, or those in an unbroken condition, precluded identification to species, or species description. But on the basis of canine morphology (the position of cingular cusps), an emballonurid species, most likely representing the genus Taphozous, is present. A small vespertilionid species was also present. Based on the wear striations on the M, of M. handae (Fig. 5D), probably derived from crushing the bones of vertebrates, these smaller bat species might well have been prey, as well as co-inhabitants of the roost. Body parts of the species Taphozous georgianus, Rhinonicteris aurantia and Vespadelus finlaysoni have all been observed in the prey accumulations of modern M. gigas in the Pilbara region of Western Australia and Northern Territory (Churchill, 2008; K. N. Armstrong personal observations).
ACKNOWLEDGEMENTS. The authors are grateful to the Kyoto University Museum for support and use of facilities; to Hidetoshi Nagamasu for help with the SEM; the Western Australian Museum and the Australian National Wildlife Collection for the loan of material; Alexander Baynes of the Western Australian Museum for information on the collection and careful editing of the manuscript; and to Robin Beck, Gilbert Price, and Sue Hand, for helpful review comments that resulted in a much-improved text. KPA acknowledges the support of the Kyoto University Museum, which hosted him as a visiting professor; and KNA gratefully acknowledges the support of a JSPS Postdoctoral Fellowship for Foreign Researchers while at the Kyoto University Museum. Ken Aplin is a posthumous author on this publication. He collected and discovered the material described herein, designed the study, took the images together with KNA, and made numerous contributions and comments as the earlier versions of the manuscript were drafted. Some details about Ken's Dingo Gap collection are unfortunately not available.
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Records of the Australian Museum (2020) vol. 72, issue no. 5, pp. 175-191 https://doi.org/10.3853/].2201-4349.72.2020.1731
Records of the Australian Museum
a peer-reviewed open-access journal
published by the Australian Museum, Sydney communicating knowledge derived from our collections ISSN 0067-1975 (print), 2201-4349 (online)
Fossil Uromys (Rodentia: Murinae) from Central Queensland,
with a Description of a New Middle Pleistocene Species
JONATHAN CRAMB'^(9, Scott A. HOCKNULL?(9, AND GILBERT J. PRICE?®
! Geosciences, Queensland Museum, 122 Gerler Road, Hendra QLD 4011, Australia ? School of Earth and Environmental Sciences, The University of Queensland, Brisbane QLD 4072, Australia
? School of BioSciences, Faculty of Science, University of Melbourne, Melbourne VIC 3010, Australia
ABSTRACT. The first fossil species of Uromys (Giant Naked-tailed Rats) is described, as well as the southern- most records of the genus based on palaeontological data. Uromys aplini sp. nov. lived during the Middle Pleistocene in the area around Mount Etna, eastern central Queensland, but was probably driven extinct by climate-mediated habitat loss sometime after 205 ka but before c. 90 ka. A second species, the extant U. caudimaculatus, occurred in the area during the Late Pleistocene, but became locally extinct prior to the Last Glacial Maximum. These fossils indicate an unexpectedly high diversity of species of Uromys in Australia, suggesting a long occupation of the continent. Phylogenetic analysis places U. aplini together with other species of Uromys endemic to Australia, at the base of the radiation of the genus. This may indicate that the initial diversification of Uromys occurred in Australia rather than New Guinea, as has previously been thought. These new Quaternary records of Uromys occur approximately 550 km south of the southern-most modern record for the genus, indicating that Uromys was able to cross the southern
St Lawrence biogeographic barrier, possibly twice during the Pleistocene.
Introduction
Uromys (commonly called “Giant Rats" or “Giant Naked- tailed Rats") is a genus of generally very large murine rodents whose species are found on mainland and continental islands of northern Sahul (Australia and New Guinea), and the Melanesian island archipelago (Fig. 1). They belong to the tribe Hydromyini, in a subclade called the Uromys division (colloquially known as the “Mosaic-tailed Rats"), that also includes four related genera: Melomys, Paramelomys, Protochromys, and Solomys (Musser & Carleton, 2005; Lecompte et al., 2008; Aplin & Helgen, 2010). The ecology and conservation status of extant species of Uromys was summarized by Flannery (1995a, 1995b), Breed & Ford (2007), Moore (2008), and Moore & Winter (2008). These authors noted that many species are presently endangered, critically endangered or presumed extinct.
Keywords: Hydromyini; Muridae; rainforest; extinction
Currently, 11 species of Uromys are recognized. Two widely distributed and morphologically variable species occur on mainland New Guinea (U. anak and U. caudimactulatus, the latter also occurring on several nearby islands), with a further four near threatened to critically endangered species that are endemic to the nearby islands of Biak (U. boeadii), Awai (U. emmae), New Britain (U. neobrittanicus) and Kai Besar (U. siebersi) (Flannery, 1995a, 1995b; Musser & Carleton, 2005). Four species are recorded from the Solomon Islands, namely U. imperator, U. porculus, U. rex, and U. vika, all of which are either endangered, critically endangered or presumed recently extinct (Flannery, 1995b; Lavery & Judge, 2017; taxonomic authorities listed below).
In Australia, two species of Uromys are currently recognized (Breed & Ford, 2007). Uromys caudimaculatus has a distribution stretching from Cape York to the most
Taxonomic registration: urn:lsid:zoobank.org:pub:BEE60C62-CA61-48C2-B8EB-E7DAC94FEA09
Corresponding author: Jonathan Cramb j2.cramb@connect.qut.edu.au
Received: 3 February 2020 Accepted: 20 August 2020 Published: 25 November 2020 (in print and online simultaneously) Publisher: The Australian Museum, Sydney, Australia (a statutory authority of, and principally funded by, the NSW State Government)
Citation: Cramb, Jonathan, Scott A. Hocknull, and Gilbert J. Price. 2020. Fossil Uromys (Rodentia: Murinae) from central Queensland, with a description of a new Middle Pleistocene species. In Papers in Honour of Ken Aplin, ed. Julien Louys, Sue O’Connor, and Kristofer M. Helgen. Records of the Australian
Museum 72(5): 175-191. https://doi.org/10.3853/j.2201-4349.72.2020.1731
Copyright: © 2020 Cramb, Hocknull, Price. This is an open access article licensed under a Creative Commons Attribution 4.0 International License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided
the original authors and source are credited.
COLIN
176 Records of the Australian Museum (2020) Vol. 72
U. boeadii U. emmae
U. neobrittanicus
Laura Basin
Black Mountain Corridor
U. sherrini
U. imperator U. porculus U. rex
U. hadrourus F
P |
dekin G rdeki ap
Mt Etna and Capricorn Caves
(U. aplini and U. caudimaculatus)
St Lawrence Gap——]
Figure 1. Map of north-east Sahul and Melanesia showing the location of study sites, the modern distributions of species of Uromys, and barriers to dispersal of mesic taxa in eastern Queensland (after Bryant & Krosch, 2016). Bathymetric depth to 200 m marked in light blue. Distribution data is from Aplin & Flannery (2017), Aplin et al. (2017), Groves & Flannery (1994), Kennerley (2016), Lavery (2019), and Woinarski & Burbidge (2016). Spot distribution of U. sherrini is based on known specimens in the collections of the Queensland Museum, CSIRO National Wildlife Collection, and Natural History Museum (London).
southerly modern occurrence of the genus, just south of Townsville in the Bowling Green National Park (Moore, 2008); QMJM1248 from Atlas of Living Australia website at https://www.ala.org.au/ (accessed 10 January 2020). The taxonomic history of Australian populations of U. caudimaculatus, and extralimital taxa synonymized with it, was summarized by Jackson & Groves (2015). The second, smaller Australian species, U. hadrourus, is restricted to the upland regions of north-east Queensland (Atherton Tableland, Mount Carbine, Thornton Peak, and Mount Bartle Frere). A third taxon, U. sherrini, described originally by Thomas (1923a), is currently considered to be a junior synonym of U. caudimaculatus (Tate, 1951), but Kristofer Helgen and Ken Aplin (pers. comm. November 2009) considered U. sherrini to be distinct from U. caudimaculatus on the basis of unpublished morphological and molecular comparisons. We therefore treat it as a separate species in this study.
The evolutionary history of Australian rodents has been investigated in recent decades using several lines of morphological (e.g., craniodental, phallic, and spermatozoan morphology) and molecular evidence to assess phylogeny
(e.g., Lidicker & Brylski, 1987; Groves & Flannery, 1994; Breed & Aplin, 1995; Rowe et al., 2008; Robins et al., 2010; Steppan & Schenk, 2017). Molecular sampling of hydromyin taxa is incomplete, and meta-analyses that include broad taxonomic sampling have recovered specific or generic level relationships that are questionable. For example, Upham et al. (2019), in their meta-analysis of mammalian phylogenies recovered Pithecheir as the sister taxon of Uromys, despite the placement of these genera in different divisions by other authors (Musser & Carleton, 2005). Bryant et al. (2011) and Lavery & Judge (2017) both conducted molecular analyses of Uromys division taxa, but unfortunately did not include the majority of species of Uromys. Bryant et al. (2011) did, however, recover Paramelomys as the sister taxon to a clade containing Melomys, Solomys, and Uromys, providing a potentially useful outgroup for any morphological assessment of phylogeny. Morphological phylogenetic methods are obviously of vital importance to palaeontological studies, but we are aware of only one published example that included Australian species of Uromys: Groves & Flannery (1994) in their revision of the genus.
Cramb et al.: New Middle Pleistocene Uromys species 177
M; Mi anterior buccal length M> M2 XE breadth M3 Ms
Figure 2. Molar cusp terminology. In Uromys and closely related genera the cusps in each molar loph are fused, so individual cusps may be difficult to distinguish in worn specimens. Molar terminology follows Musser (1981), Aplin & Helgen (2010), and Lazzari et al. (2010). (A) upper molars, left side in occlusal outline; (B) lower molars, right side, in occlusal outline. Abbreviations: a-buc, antero-buccal cuspid; a-/in, antero-lingual cuspid; ed, entoconid; hd, hypoconid; md, metaconid; pd, protoconid; pi, posterior indent; psc, posteroconid.
With the majority of species of Uromys found in New Guinea and Melanesia, it has long been assumed that Uromys had its phylogenetic origin in these regions; Watts & Aslin (1981) posited that Uromys was a relatively recent arrival in Australia, having crossed the Torres Strait during the Last Glacial Maximum. This view was not held by all researchers with Tate (1951) suggesting that Uromys arrived in Australia during the Middle Pleistocene, and Hand (1984) stating that the timing of arrival was unclear. The recognition that U. hadrourus was a species of Uromys rather than a species of Me/omys (see Jackson & Groves, 2015) hinted that Uromys had been present in Australia for some substantial time, with Aplin (2006) citing Watts and Baverstock’s (1994) molecular data to suggest the possibility that the genus was present before 2.5 Ma. Such a possibility would be supported if fossils of the right age were available. Despite the presence of murines in Sahul since at least 4.18 Ma (Piper et al., 2006), published reports of fossil Uromys are almost all restricted to the Late Pleistocene and Holocene (e.g., O'Connor ef al., 2002; Aplin et al., 1999). The exception is Hocknull (2005), who reported a large Mosaic-tailed Rat from the Mount Etna caves, which was later found to be of Middle Pleistocene age (Hocknull et al., 2007). This taxon is here described as Uromys aplini sp. nov., and is the geologically oldest species of the genus yet recorded.
Materials and methods
All fossil specimens included in this study were excavated as part of ongoing research into the fossils of the caves in the Mount Etna and Capricorn Caves region, eastern central Queensland (Fig. 1). Fossils were compared with specimens of all available species of Uromys in the collections of the Queensland Museum, Australian Museum, and the Australian National Wildlife Collection (Appendix 1). Where specimens of some species were not available in Australian collections, comparisons with published descriptions and images were made. Fossil specimens were measured with digital callipers, and imaged with a Visionary Digital "passport storm" camera system, an Olympus Stylus TG-4 compact digital camera, a Hitachi TM-1000 environmental scanning electron microscope at the Queensland Museum, and a Leica DFC450 C digital microscope camera at the School of Earth and Environmental Sciences, The University of Queensland. All fossils described in this paper are catalogued in the collections of the Queensland Museum, in Brisbane, Australia. Molar cusp terminology is presented in Fig. 2.
Study sites background
Fossil remains described here were collected from cavernous limestone located at the Mount Etna and Limestone Ridge Caves National Park and the Capricorn Caves Tourist Park (Hocknull, 2005, 2009; Price et al., 2015). The bulk of fossils
178 Records of the Australian Museum (2020) Vol. 72
Table 1. Character matrix used in phylogenetic analysis. Modified from Groves & Flannery (1994). Note that the original numbering of characters is retained from Groves & Flannery (1994), although external characters are removed. Additional characters: (50) M'? length: 0 € 7 mm, 1 = 7-8 mm, 2 = 8-9 mm, 3 = 9-10 mm, 4 = 10-11 mm, 5 = 11-12 mm, 6 = 12-13 mm, 7 = 13-14 mm; (57) M? length/M! width: 0 = 3-3.2, 1 = 3.2-3.4, 2 =3.4-3.6, 3 = 3.6-3.8, 4 = 3.84.0.
LL zm rq 123456789012345678
Paramelomys rubex 1 ? ? ?000?7? 1100001? 0 Uromys anak 0-1 (,1.1 d.1 O-1 0 1^0 LI Q0 0.1 -8 U. aplini sp. nov. 1??? 100??110100?? I U. boeadii 010100000000000000 U. caudimaculatus 1 1 11 101011001001 11 U. emmae Lab Det Ol L--0. 1 ee T $-0 U. hadrourus Ld 1.bd9-0 O-[rd.c00.0 9 IT 9 4 U. imperator 101001010100110000 U. neobrittaniens 0 10110101010100000 U. porculus 000001010000110000 U. rex 0010010101 1 1110000 U. sherrini I| hU EA ET E Cu-IT-peg-1 PO ea U. vika 0??7?7010??11?1000?20
from the deposits are most likely derived from the feeding activities of owls. Fossil deposits from Mount Etna were described initially by Hocknull (2005) with biocorrelation of these faunas suggesting a Pliocene age. Subsequent radiometric dating of flowstones associated with the fauna demonstrated, however, that these deposits were in fact Pleistocene in age and restricted to the Middle Pleistocene (Hocknull et al., 2007). Additional sites, descriptions, and dating assessments were also undertaken and available in Hocknull (2009). At Capricorn Caves Tourist Park, Queensland Museum Locality (OML) 1456 is located within the Olsen's Cave system. Faunal remains recovered from this site were described and chronometrically dated using a combination of radiocarbon and uranium-series techniques, resulting in a Late Pleistocene age (Price et al., 2015).
At Mount Etna, Middle Pleistocene faunal assemblages dated to 72500 ka to 7280 ka are interpreted as having occupied closed rainforest palaeoenvironments (QML1311H, QML1313) including taxa or lineages now only found in rainforests of northern Queensland and New Guinea (Hocknull, 2005; Hocknull et al., 2007; Price & Hocknull, 2011; Cramb & Hocknull, 2010). A younger Middle Pleistocene fauna (OML1312) dated to 205-170 ka is interpreted as having occupied a xeric environment and includes species or lineages found in arid habitats today. The Late Pleistocene fauna (QML 1456) from Capricorn Caves is interpreted to be more mesic in comparison to the xeric Middle Pleistocene fauna, but still drier-adapted than the older Middle Pleistocene rainforest fauna. Together, these three periods show major faunal transitions typified by local extinction and replacement of species with new more dry- adapted forms (Hocknull et al., 2007; Price, 2012).
Phylogenetic analysis
A preliminary attempt was made to ascertain the phylo- genetic position of the new fossil species of Uromys by scoring craniodental characters using a character state matrix first developed by Groves & Flannery (1994).
1 9
— =- O OF D me mæ me e =~ e -—
22222222333444444445 5 01234567679012467890 1 2702700000110111001000 1 1010000000011101000 {67} {123} 120 2 E000 07] 93-4. X9 3 X3 0.442 00100000000111010005 2 10000000111 1110110041456) {1234} 10000000010111011015 2 1000000011111101110 £12} £0123} 01010011001000100006 1 1010000000011101000 {67} {23} 01010011001000000000 2 0111111100100000000 145 {12} 270270000011111101100 £56} {12} 25 541 0 043 2n OLD Or 0% 9 0 0-9.2 1
We restricted our assessment to the craniodental characters used by Groves & Flannery (1994) with the addition of one measurement character (Character 50: M!= length) and one measurement ratio (Character 51: M'? length/M! width). These continuous data were binned and scored as multi-states for variable taxa. The length measurement was ordered in the analysis. All other characters were unordered. Three additional species, including the fossil taxon U. aplini sp. nov., U. vika (based on the description published by Lavery & Judge, 2017) and specimens considered to represent U. sherrini were used to augment the phylogenetic analysis (see Appendix 1). Only U. siebersi was not able to be scored, due to the rarity of specimens. Some character states were not able to be scored due to either their lacking in preservation in the fossils and extant craniodental remains, or obscurity in determining the state. Character states for 2-4, 8-9, 17, 20, 22, 44 could not be ascertained from comparison of specimens with character descriptions provided by Groves & Flannery (1994), so were given a “?” and considered uncertain. We have amended the character state of character 11 for U. hadrourus because it was incorrectly scored in Groves & Flannery (1994). All characters were weighted equally.
Molecular analysis by Bryant et al. (2011) found that Melomys is the sister taxon to Uromys, while Paramelomys is the sister clade to both genera. For this reason, we included Paramelomys rubex as the outgroup for the analysis to polarize the character states within Uromys. The modified matrix is shown in Table 1. The analysis was conducted using Mesquite version 3.61 (Maddison & Maddison, 2019) and PAUP 4.0 (Swofford, 2001).
Our phylogenetic assessment is only considered to be preliminary using standard parsimony, where the characters are polarized by an outgroup (Paramelomys rubex). Multi- state characters are considered to be polymorphic, whilst those characters with “?”s are considered to be uncertain. The tree-searching algorithm used was tree-bisection and reconnection (TBR) from 100 random additions. Bootstrap values were calculated using 1000 replicates, with the resulting nodes with values greater than 50% retained.
Abbreviations
QMF—Queensland Museum fossil specimen; QML— Queensland Museum fossil locality; QMJ, QMJM— Queensland Museum modern mammal specimen; CM— CSIRO Australian National Wildlife Collection mammal specimen; AM M.—Australian Museum mammal specimen; NMVC—Museum Victoria mammal specimen; ka (kilo annum)—thousands of years ago; Ma (mega annum)— millions of years ago.
Results
Two species of Uromys were identified from fossils in the study region, the extant U. caudimaculatus and the extinct U. aplini sp. nov. Uromys caudimaculatus was recovered from Capricorn Caves (QML 1456) in excavation spits 142-147 cm, 152-157 cm, and 177-182 cm (inferred as dating to the Late Pleistocene), and U. aplini sp. nov. was recovered from multiple Middle Pleistocene deposits at Mount Etna.
Systematic palaeontology
Class Mammalia Linnaeus, 1758
Subclass Theria Parker & Haswell, 1897
Supercohort Placentalia Bonaparte, 1838
Order Rodentia Bowdich, 1821
Family Muridae Illiger, 1811
Subfamily Murinae Illiger, 1811
Tribe Hydromyini Alston, 1876 sensu Lecompte et al., 2008
A B
Cramb et al.: New Middle Pleistocene Uromys species 179
Uromys Peters, 1867
Synonyms: Gymnomys Gray, 1867; Cyromys Thomas, 1910; Melanomys Winter, 1983 (but see Jackson & Groves, 2015, for explanation).
Included species:
Uromys caudimaculatus (Krefft, 1867) Uromys imperator (Thomas, 1888) Uromys rex (Thomas, 1888)
Uromys porculus (Thomas, 1904)
Uromys anak Thomas, 1907
Uromys sherrini Thomas, 1923a
Uromys siebersi Thomas, 1923b
Uromys neobrittanicus Tate & Archbold, 1935 Uromys hadrourus (Winter, 1984) Uromys boeadii Groves & Flannery, 1994 Uromys emmae Groves & Flannery, 1994 Uromys vika Lavery & Judge, 2017
Generic diagnosis: Groves & Flannery (1994) considered three cranial characters (with the addition of one soft-tissue character) to be diagnostic of species of Uromys: a hard palate that extends posterior of the posterior margin of M^, I, is much deeper than it is wide, and a greatly expanded anterolateral spine on the auditory bulla.
Uromys caudimaculatus (Krefft, 1867) Fig. 3A, 3B
Material examined. QML1456: spit 142-147 cm: QMF60126 right M', QMF60127 left M', OMF60128 right M!, QMF60129 left M?, QMF60130 left Mj, QMF60131 right M». Additional specimens were also recovered from spits 152-157 cm, and 177-182 cm.
C
Figure 3. Succession of Uromys spp. in the Mt Etna area. (A—B) Uromys caudimaculatus, (A) QMF60126 right M', QML1456 spit 142-147, c. 50 ka; (B) QMF60127 left M', deposit and age as for A. (C-D) Uromys aplini, (C) QMF55340 left M', QML1312, 205-170 ka; (D) QMF60125 right M', QML1311 H, > 450 ka. Scale bar = 1 mm.
180 Records of the Australian Museum (2020) Vol. 72
Figure 4. Comparison of skulls of Uromys sherrini and U. caudimaculatus in ventral outline. (4) U. sherrini (CM10822); (B) U. caudimaculatus (CM705). The larger degree of deflection in the zygomatic plate, seen in U. caudimaculatus, is indicated with an arrow.
Scale bar = 5 mm.
Remarks. Isolated molars of this species are distinguished by a combination of characters including very large size; crescentic lophs on M'?; deep posterior indent present on M'7; long, variably bifurcated lingual root on M'?; crescentic lophids on M, 5; lingual root present on M}; large posteroconid on M, »; and a relatively shallow cleft between the antero-buccal cuspid and protoconid on M}.
Uromys caudimaculatus was previously considered to include U. sherrini, so it is pertinent to include a list of characters that distinguish these species. These are: the margins of the interorbital area above the orbits, which are almost parallel in U. sherrini but divergent in U. caudimaculatus (Thomas, 1923a); the anterior palatal foramina are commonly broader in U. sherrini; the M!? of U. sherrini have shallower posterior indents on the T8-9 complex; the posterior loph on M?/, is commonly narrower; the nasals are shorter, not projecting anterior of the premaxillae as in U. caudimaculatus; the anterior edge of the zygomatic plate is directed antero-lingually in dorsal or ventral outline, while that of U. caudimaculatus 1s deflected, making it parallel with the rostrum (Fig. 4).
Specimens of U. caudimaculatus from QML 1456 have only slightly worn tooth crowns, indicating that the owls thought to be the accumulating agents of the deposit were preying on young individuals. The excavation spit that yielded the stratigraphically youngest U. caudimaculatus specimens (1.e., 142-147 cm) is probably slightly younger than 50 kyr (see Price et al., 2015 for a full discussion of the age of the deposit). The older spits (1.e., 152-157 cm and 177—182 cm) are undated, but are likely to be Late Pleistocene (c. 80—60 ka) based on the age model presented in Price et al. (2015). Deposition in QML 1456 is thought to have been continuous during the late Quaternary, with no evidence of depositional hiatuses.
Uromys aplini sp. nov.
urn:Isid:zoobank.org:act:C52317A8-D118-4E10-AA9C-21DF62C8EECA
Figs 3C, 3D, 5-7
Holotype. QMF52014 (Queensland Museum fossil specimen) partial skull, QML1313 (Queensland Museum fossil locality) Speaking Tube Cave, Mount Etna, eastern central Queensland. Deposit has a minimum age of c. 280 ka (Hocknull ef al., 2007). Paratypes. QMF55753 partial skull; QMF55542 right mandible with M,; both specimens have same locality as holotype, QML 1313.
Material examined. QML1311H: QMF55547 right M', QMF55548 right M?, QMF55549 right MP, QMF55550 left M,, QMF55551 right M,, QMF55552 left M,, QMF60125 right M'; QML1313: QMF52014 partial skull, QMF55522 left M', QMF55523 left M', QMF55524 right M!, QMF55525 left M?, QMF55526 right VP, QMF55527 right M?, QMF55528 right MP, QMF55529 left M?, QMF55530 left VP, QMF55531 left Mj, QMF55532 left M}, QMF55533 left Mj, QMF55534 right M,, QMF55535 left M», QMF55536 left Mj, QMF55537 right Mj, QMF55538 left Mj, QMF55539 left M, QMF55540 left I', QMF55541 left maxilla fragment, QMF55543 right mandible with M, and M;, QMF55544 right M?; QML1313A: QMF55545 left M!, QMF55546 right M1; QML1312: QMF55340 left M!. Additional specimens were also recovered from QML 1284, QML1284A, QML1311C/D, QML1311J, QML1383, QML1384LU, and QML1385.
Age Range. Chibanian (Middle Pleistocene), chronometric- ally dated to >500 ka to c. 205 ka.
Diagnosis. Large Uromys, but smaller than most species of Uromys (Uromys) with the exception of U. hadrourus (Fig.
Cramb et al.: New Middle Pleistocene Uromys species 181
Table 2. Craniodental measurements of Uromys aplini sp. nov. All measurements in millimetres. SD — standard deviation; CV = coefficient of variation; APF = anterior palatal foramen; QML = Queensland Museum Locality.
n I! width 3 zygomatic plate length 2 diastema length 2 hard palate width 2
I, width
N
M! width
oo -NA =
M? width
mean SD
1.64 0.05 7.40 0.79 14.55 0.07 10.03 0.07 1.38 0.35
2.57 na 2.84 0.22 2.82 0.12 2.85 na 2.80 0.18
2.78 na
range 1.58-1.68
6.84-7.96
14.50-14.60
9.98—10.08 1.13-1.63
na 2.53-3.02 2.73—2.90 na 2.53-3.02
na
CV
na
na
na
QML n mean SD range CV I' depth 1313 3 3.05 007 298-31] — interorbital width 1313 2 858 0.33 834-881 — APF length 1313 2 629 041 6.00-6.58 — hard palate length 1313 2 2597 0.79 25.40-26.53 — I, depth 1313 1 1.98 na na na M! length I311H 1 482 na na na 1313 3 509 020 486-542 — 1313A — — — — — 1312 1 527 na na na all 5 5.08 026 4.82-5.42 — M? length 1311H 1 333 na na na 1313 6 371 021 346-400 — 1313A — — — — — all 7 3.66 024 333-400 — M? length 1311H 2 238 007 23343 — 1313 6 226 0.15 208-252 — all 8 229 0.14 208252 — M? length 1313 2 1017 0.29 29.96-10.37 — M, length 13131H 14 415 017 396-433 — 1313 6 443 013 419-454 — 1313A 2 417 0.09 410-423 — all 12 429 0.19 3.96454 4.46 M, length 1311H 4 351 O14 342-372 — 1313 3 345 0.28 319-374 — 1313A 3 33] 0.14 323-347 — all 10 343 0.19 319-374 5.51 Mj length 13431H 2 301 026 282-319 — 1313 4 265 0.16 245-278 — all 6 277 025 245-319 —
8; Table 2); it is distinguished on the following combination of characters: posterior indent on T8—9 of M! poorly developed; molar enamel ornament moderately developed; anterior palatal foramina short, shared equally between premaxilla and maxilla; rostrum proportionally short and robust; supraorbital ridges and postorbital processes absent. Features that further distinguish U. aplini from all other species of Uromys are listed in the Remarks section.
Groves & Flannery (1994) divided Uromys into two subgenera: U. (Uromys) and U. (Cyromys). Uromys aplini is placed in U. (Uromys) on the basis of the following diagnostic characters identified by Groves & Flannery (1994): short, slit-like anterior palatal foramina; simplified, elongate molars; reduced M?/4; posteriorly lengthened bony palate; reduced anterior lophid on M,, which fuses to middle lophid after moderate wear; zygomatic arches swing posteriorly and ventrally to level of molar alveoli; and orthodont incisors.
2.83 0.12 2.71-3.04 — 2.72 na na na 2.81 0.11 2.71-3.04 —
Orne
M? width
N
2.13 0 2.08 0.12 2.09 0.10
2.13-2.13 — 1.92-2.21 — 1.92-2.21 —
oo ON
M, , length 3 10.740.24 10.48-10.95 —
4 2.62 0.08 6 2.67 0.8 2 0.12 1
0.14
2.55-2.73 — 2.37-2.93 — 2.46-2.63 — 2.37-2.93
M, width
2.55 2 2.63
M, width 5 285 3 281 3 268
11 2.79
0.11 0.09 0.08 0.12
2.69-2.97 — 2.70-2.89 — 2.61-2.77 — 2.61-2.97
M; width 2 248 4 229
6 235
0.01 0.12 0.13
2.47-2.48 — 2.18-2.43 — 2.18-2.48 —
Etymology: Named for Kenneth Peter Aplin (1958-2019), for his contribution to Australian palaeontology and the taxonomy and systematics of Australasian murids.
Description
Skull. Two partial skulls are known (QMF52014 and 55753, Fig. 5A,B). The lacrimals, jugals, and much of the posterior of the skull and basicranium are missing from both specimens.
The nasals appear to be consistent in width along preserved length, tapering sharply at posterior contact with frontals.
Premaxilla short and robust. Anterior palatal foramen short, narrow, tapering abruptly at extremities, occupying similar area of premaxilla and maxilla. Anterior palatal foramen roughly half of length anterior of M!. Narrow crest on ventral surface of maxilla between junction with premaxilla and anterior margin of M! variably
182 Records of the Australian Museum (2020) Vol. 72
Figure 5. Cranial elements of Uromys aplini sp. nov. (4) QMF52014 partial skull in (top to bottom) dorsal, right lateral, and ventral view; (B) QMF55753 partial skull in dorsal, left lateral, and ventral view; (C) QMF55541 left maxilla fragment, showing a narrow crest
on the diastema. Scale bar = 5 mm.
developed, likely associated with age (some specimens, e.g., QMF55541, have it developed to an extreme degree, forming a blade. Fig. 5C). Zygomatic plate long, anterior edge straight, evenly curving posteriorly at dorsal end into zygomatic arch. Maxillary portion of zygomatic arch slopes posteroventrally at approximately 45? angle, almost reaching level of molar alveoli.
Palatine contacts maxilla level with posterior margin of M!. Posterior palatal foramen level with M?. Palate terminates in small, blunt postpalatal spine, approximately level with most posterior point of maxilla.
Frontals with sharp corner between temporal and orbital faces. Very small postorbital processes on squamosals, not associated with sutures. Supraorbital ridges not evident on frontals; parietal crests weakly developed on dorsal margin of squamosals and parietals. Braincase not greatly inflated, relatively flat dorsally, width exaggerated on QMF52014 by parting of parietals at midline.
I'. Proportionally deep (I! depth/width of QMF52014 =
2.98/1.66 mm), orthodont. One paratype (QMF55753, Fig. 5B) appears to retain orange pigment in enamel, although this may instead be diagenetic iron staining.
M!. Crown elongate, rounded anterior margin and angular posterior margin. Lophs sloped posteriorly. Accessory cusp and anterior cingulum absent. Lingual cusps bulge lingually at bases, giving lingual margin of crown an irregular appearance. Buccal cusps do not bulge at bases. T1 oval-shaped in occlusal outline, oriented antero-buccally postero-lingually. T1 postero-lingual of T2. T1 separated from T2 by shallow cleft; T1 and T2 join after moderate wear. T2 broad and robust. T3 directly buccal of T2, posterior margins of T2—3 form straight line. T3 small, fused to T2. T3 discernible from T2 by shallow, poorly defined groove on anterior face of T2-3 complex. T4 subcircular in occlusal outline when unworn, becomes subtriangular after wear. In occlusal outline, T4 projects to a point anterior of the junction with T5; tapers posteriorly. T4 tapers towards T5, separated by a shallow cleft, joined after moderate wear. T5
Cramb et al.: New Middle Pleistocene Uromys species 183
B
Figure 6. Mandibles of Uromys aplini sp. nov. (A) QMF55542 right mandible with M,; (B) QMF55543 right mandible with M,, partial
M,, and M,. Scale bar = 5 mm.
broad, boomerang-shaped in occlusal outline, with bulk of T5 antero-buccal of T4 and antero-lingual of T6. T6 poorly defined, variably separated from T5 by shallow groove on anterior face of T5—6 complex. T6 broad, oriented antero- lingually postero-buccally, continuous with buccal half of T5. T5-6 complex roughly parallel with T8—9 complex. T7 appears absent, although one specimen (QMF55522, Fig. 7C) has a bulge in the posterior loph that could be interpreted as a T7 fused to T8. T8—9 complex broad, based between buccal margin of crown and posterior point of T4. T9 fused to T8, poorly defined by change in angle of anterior margin of occlusal surface of T8—9 complex. Very small posterior indent associated with posteroloph, commonly not visible in occlusal view.
Fine enamel ornament present on anterior faces of all lophs. M! has four roots: anterior, two lingual (commonly fused close to crown), and postero-buccal. Molar roots commonly split into multiple rootlets at tips. Alveoli of lingual roots variably fused, creating appearance of a single elongate lingual root.
M". Elongate, tapering posteriorly. Lingual cusps bulge lingually at bases, buccal cusps do not. T1 forms antero- lingual corner of crown. T1 subcircular in occlusal outline when slightly worn, becomes subtriangular (tapering buccally and posteriorly) after wear. T2—3 absent. Position of T3 variably marked by shallow depression on anterior face of T5-6 complex. T4 directly posterior of T1. T4—6 loph essentially identical to that on M!. T8-9 complex based between buccal margin of T6 and posterior point of T4. T8—9 tapers slightly but does not form a point. T8 and T9 not differentiated.
A ridge on the lingual side of T8 may represent a T7. Very small posterior indent associated with posteroloph, commonly not visible in occlusal view. Fine enamel ornament on anterior faces of both lophs, possibly less developed than that on M!. M? has four main roots: antero- buccal, postero-buccal, postero-lingual, and antero-lingual. The antero-lingual and postero-lingual roots are variably joined. The antero-buccal root is variably bifurcated at the tip into two small rootlets.
MP. Compact and simplified, moderately reduced. Some specimens (e.g., QMF55528, Fig. 7H) subcircular in occlusal outline. T1 well defined, rounded, oval-shaped in occlusal outline. T2—3 absent. Individual cusps of T4—6 loph not discernible. T4—6 loph gently curved, most anterior point at presumed location of T5. T4—6 loph sloped posteriorly. Posterior cusp broad, slightly narrower than T4—6 loph. Posterior cusp upright, very close to T4—6 loph. Some specimens (e.g., QMF55530, Fig. 7G) have posterior cusp very close to “T4” but larger gap separating posterior cusp from “T6”. Posterior cusp oval-shaped in occlusal outline. One specimen (QMF55544) has a small posterior cingulum cusp. M? has four roots: antero-buccal, posterior, and joined antero-lingual and lingual.
Mandible. No specimens are completely intact, with all displaying degrees of damage to the posterior processes and incisor alveolus. Mandible deep and robust, with deepest point ventral of M,. M, longer than M5, but similar width. M, smaller than M», but not heavily reduced. Coronoid process damaged or missing on all specimens, but appears to be taller than articular process. Articular process projects slightly posterior of angular process. Angular process damaged on
184 Records of the Australian Museum (2020) Vol. 72
A B
Figure 7. Isolated molars of Uromys aplini sp. nov. (A) QMF55524 right M!; (B) QMF55523 left M'; (C) QMF55522 left M'; (D) QMF55527 right M?; (E) QMF55525 right M?; (F) QMF55526 right M?; (G) QMF55530 right M?; (H) QMF 55528 left M?; (7) QMF55529 left M3; (J) QMF55531 left M,; (K) QMF55533 left M,; (L) QMF55532 left M,; (M) QMF55534 right M,; (N) QMF 55536 left Mz; (O) QMF55535 left M5; (P) QMF55537 right M4; (Q) QMF55539 left M4; (R) QMF55538 left M,. Scale bar = 1 mm.
all specimens, but appears to be rounded. Mental foramen ventral of dorsal inflection of diastema. Superior masseteric crest very poorly defined below molars; inferior masseteric crest well developed, terminates anteriorly posterior of mental foramen. Mandibular symphysis marked by dorsal crest in anterior part of diastema; symphysis ends ventrally of anterior root of M,. Incisor alveolus forms prominent tubercle on buccal surface of ascending ramus, although this is damaged in all specimens. Postalveolar ridge sharply defined below and posterior of M;, less defined posterior of retromandibular fossa. Retromandibular fossa small in young adult individuals, greatly expanded in mature individuals (assessed on the basis of molar wear).
I,. Proportionally deep (I, depth/width of QMF55542 [subadult] = 1.98/1.13, adults proportionally deeper). One specimen (QMF55543, Fig. 6B) may retain orange pigment in enamel, but lost in majority of specimens.
M,. Rounded anteriorly, subrectangular posteriorly. Anterior and middle lophids crowded together. Anterior lophid upright, middle, and posterior lophids sloped anteriorly. Anterior lophid narrower than middle lophid. Middle and posterior lophids of similar width. Antero- buccal cuspid small, subcircular in occlusal outline, fused to antero-lingual cuspid. Antero-buccal and antero-lingual cuspids only distinguishable when unworn, form single anterior lophid after moderate wear. Antero-lingual cuspid much larger than antero-buccal cuspid, forms much of the anterior lophid. Unworn specimens show antero- lingual cuspid with two buccal extensions: one joining the antero-buccal cuspid, the other directly posterior and postero-lingual of the antero-buccal cuspid between the main body of the anterior lophid and the middle lophid. Antero-buccal cuspid separated from protoconid by shallow cleft, eliminated by wear on some specimens; antero-lingual cuspid separated from metaconid by relatively deeper cleft, more resistant to wear.
Protoconid subtriangular in occlusal outline, tapering lingually to join metaconid and posteriorly along buccal margin of crown. Metaconid subequal in size to protoconid. Metaconid subtriangular in occlusal outline, tapering buccally to join protoconid, tapering slightly posteriorly and anteriorly. Anterior margin of middle lophid buccally perpendicular to long axis of crown, curves antero-lingually to most anterior point of metaconid. Posterior face of middle lophid curved, bowing anteriorly between most posterior points of protoconid and metaconid.
Entoconid directly posterior of metaconid. Entoconid subtriangular in occlusal outline, tapering buccally to join hypoconid and posteriorly to a lesser degree. Hypoconid directly posterior of protoconid, buccal and slightly posterior of entoconid. Hypoconid subtriangular in occlusal outline, tapering lingually to join entoconid, and posteriorly to a lesser degree. Hypoconid and entoconid variably have small anterior extensions. Hypoconid projects slightly further posteriorly than entoconid. Anterior edge of occlusal surface of posterior lophid commonly straight, but some specimens (e.g., QMF55533, Fig. 7K) have a slight bulge, at approximately the midline of the crown. Posterior margin of occlusal surface curved, bowed anteriorly with most anterior point directly posterior of midline junction between hypoconid and entoconid. Posteroconid tolerably well
Cramb et al.: New Middle Pleistocene Uromys species 185
developed, lenticular in occlusal outline, bound by bases of hypoconid and entoconid. Posteroconid does not project beyond posterior margin of crown. Fine enamel ornament on posterior faces of middle and posterior lophids, not visible on anterior lophid due to close proximity of middle lophid. M, has three roots: anterior, a broad posterior, and a small lingual root.
M,. Crown roughly square in occlusal outline, with rounded corners. Both lophids sloped anteriorly. Protoconid larger than metaconid, both at apex and base. Protoconid tear-shaped in occlusal outline, tapering lingually to join metaconid at midline of crown. Metaconid directly lingual of protoconid, tear-shaped in occlusal outline, tapering buccally. Unworn specimens (e.g., QMF55536, Fig. 7N) have no cleft separating protoconid and metaconid. Hypoconid directly posterior of protoconid. Hypoconid tear-shaped when unworn, becomes subtriangular after light wear. Hypoconid tapers antero-lingually to join entoconid at midline of crown. Hypoconid tapers posteriorly further than entoconid. Entoconid slightly less robust than hypoconid. Entoconid tear-shaped in occlusal outline, tapering directly buccally, meeting hypoconid at an angle. No separation between hypoconid and entoconid.
Posteroconid well developed, lenticular in occlusal outline. Posteroconid commonly centred on midline of crown, although one specimen (QMF55535, Fig. 7 O) has it centred slightly buccal of the midline. Posteroconid projects slightly beyond posterior margin of crown. Fine enamel ornament on posterior faces of lophids. M, has two broad roots: anterior and posterior.
Mi. Almost triangular in occlusal outline, with heavily rounded corners. Protoconid slightly larger than metaconid. Protoconid tear-shaped in occlusal outline, tapering lingually to join metaconid. Metaconid tear-shaped in occlusal outline, tapering buccally to join protoconid. Protoconid and metaconid joined by narrow ridge. Posterior lophid broad, commonly supplemented by small cuspid on buccal side. Posterior lophid shaped like an elongate oval in occlusal outline, supplementary cuspid subcircular. Supplementary buccal cuspid variably separated from posterior lophid by shallow cleft or fused. M, has three roots: posterior, and fused antero-buccal and antero-lingual.
Remarks
Uromys aplini can be distinguished from other members of Uromys (Uromys) as follows: Uromys aplini differs from U. caudimaculatus by being smaller; having a less elongate rostrum; having a smaller posterior indent in T8—9 on M'?; and having shorter anterior palatal foramina. Uromys aplini differs from U. sherrini by being smaller; having a less elongate rostrum; and having a more reduced M?/4. Uromys aplini differs from U. hadrourus by being larger; having proportionally shorter anterior palatal foramina; having a proportionally shorter rostrum; by commonly possessing a crest on the maxilla between the maxilla/premaxilla contact and M!: and having a zygomatic arch that plunges further ventrally, reaching the level of the molar alveoli. The molars of Uromys aplini could not be effectively compared to those of U. hadrourus, as all examined specimens of the latter were heavily worn. Uromys aplini differs from
186 Records of the Australian Museum (2020) Vol. 72
M+? length (mm)
1.75 2 2.25 2.5 2.75 3
M! width (mm)
. (Uromys) neobrittanicus . (Uromys) anak
. (Uromys) caudimaculatus . (Cyromys) imperator
. (Uromys) sherrini
. (Uromys) siebersi
. (Uromys) boeadii
. (Cyromys) rex
. (Uromys) emmae
. (Uromys) aplini
. (Cyromys) vika
. (Uromys) hadrourus
> T c c c c c c c c c c c c
. (Cyromys) porculus
3.25 3.5 3.75
Figure 8. Bivariate plot of molar proportions (M' width vs M'? length, in mm) of species of Uromys. Additional data provided by Tate (1951), Winter (1984), Groves & Flannery (1994) and Lavery & Judge (2017). Plot generated in PAST 2.12 (Hammer ef al., 2001).
U. anak by being smaller; having a less elongate rostrum; lacking postorbital processes; having a smaller posterior indent in T8—9 on M'?; and having the anterior palatal foramina shared equally between the premaxilla and maxilla. Uromys aplini differs from U. neobrittanicus by being smaller; lacking large postorbital processes; having the skull relatively flat dorsally; and having parietals that are roughly rectangular in dorsal outline. Uromys aplini differs from U. emmae by being smaller; having a deeper zygomatic arch; having the zygomatic plate not projecting as far anterior of the zygomatic arch; and having the anterior palatal foramina shared equally between the premaxilla and maxilla. Uromys aplini differs from U. boeadii by being smaller; having smaller postorbital processes; and lacking supraorbital ridges.
Only one skull of U. siebersi is known, and this specimen was not available for the current study. But a measurement of the molar row (13.3 mm) provided by Groves & Flannery (1994) shows that U. siebersi 1s larger than U. aplini in this aspect (Table 2). Thomas (1923b) also provided measurements, though these are less precise than currently obtainable with modern precision measuring tools. The interorbital width and length of the “palatal foramina" (presumably the anterior palatal foramina) are both larger (10.3 mm and 7 mm, respectively, versus 8.34 mm and 6.00 mm for U. aplini).
Uromys aplini is hitherto known mostly from deposits at Mount Etna that are dominated by taxa that had ecological affinities to rainforest environments. The oldest deposits that yield the species are 7500 ka, whilst the youngest is 205-170 ka.
Phylogenetic analysis
Our phylogenetic analysis returned topological features similar to that recovered by Groves & Flannery (1994). We used Paramelomys rubex as the most appropriate outgroup taxon to polarize the character-states within Uromys. Thirty- three characters were parsimony informative with seven uninformative and considered to be autapomorphies of these taxa. The derived character states for characters 12, 24 and 25 are considered to be autapomorphies of U. rex, so are uninformative in relation to U. aplini. The derived character states for characters 34 and 38 are considered to be autapomorphies of U. imperator and U. emmae respectively, also uninformative for the fossil taxon. Finally, uninformative characters 15 and 37 are restricted to U. hadrourus, with the derived state of character 15 an autapomorphy and 37 ambiguous due to the missing states in the fossil taxon (U. aplini) and in U. sherrini.
The parsimony analysis returned two most parsimonious trees (MPT) of 93 steps (Fig. 9). Both MPTs consistently returned a basal split with one clade solely composed of species within the subgenus Cyromys and found today in the Solomon Islands group (U. imperator, U. rex, U. porculus, and U. vika). The Cyromys clade is strongly supported by bootstrap value of 91%. The other clade is composed solely of species within the subgenus Uromys and includes our fossil taxon, U. aplini. Although this clade is poorly supported, it is likely that the large amount of missing data and morphological variability of U. caudimaculatus have created internal instability within this clade. Further characterization of U. caudimaculatus subspecies and better
P. rubex
U. rex 51
81 U. imperator
91 U. porculus
U. vika
U. anak 87
U. neobrittanicus
— U. boeadii
U. emmae
U. caudimaculatus
U. sherrini
U. aplini
METUS U. hadrourus
Cramb et al.: New Middle Pleistocene Uromys species 187
North -«—
Mt Etna and Capricorn Caves
Uromys (Cyromys)
Uromys (Uromys)
: HK >
m £- A d
Figure 9. Results of preliminary phylogenetic analysis using parsimony. Bootstrap values > 50% provided showing monophyly of the Solomon Islands Uromys (Cyromys) and Australopapuan clade as sister taxon with Australian species basal to New Guinean species.
resolution of missing data may increase the support for the monophyly of Uromys (Uromys) and Uromys (Cyromys).
Resolution within the Uromys (Uromys) clade 1s poor, although the New Guinean U. anak and New Britain U. neobrittanicus are strongly supported (87%) as sister taxa. The positions of the remaining taxa are poorly supported by bootstrap values, but both MPTs return identical positions of all species, suggesting that the overall topology is valid. At the base of the clade lie the Australian Uromys hadrourus, the fossil taxon U. aplini, and U. sherrini. In a more derived position, sister to these Australian endemic species, is U. caudimaculatus, which 1s then sister taxon to a clade containing the northern New Guinean island endemics (U. emmae and U. boeadii), the mainland New Guinea U. anak, and U. neobrittanicus from New Britain.
The two basal clades, comprising Uromys (Cyromys) and Uromys (Uromys), were supported by Groves & Flannery (1994), so this result is not surprising. But our analysis, using Paramelomys as the outgroup, suggests that the Australian Uromys are basal to the Uromys (Uromys) clade.
The Middle Pleistocene age of our phylogenetically basal extinct taxon (U. aplini) is younger than the divergence time estimates (e.g., Early Pleistocene) for the more derived extant species within the clade (Watts & Baverstock, 1994; Bryant et al., 2011). This would probably preclude U. aplini from being a chronospecies of the extant Australian species of Uromys (Uromys).
Discussion Phylogeny and biogeography of Uromys
Our phylogenetic analysis supports three extant species of Uromys in northern Queensland, including two that are geographically restricted (U. hadrourus and U. sherrini) and one that is more broadly distributed across the northern region of Cape York (U. caudimaculatus). Fossils described here demonstrate that species of Uromys were previously more widespread in northeastern Australia than would be expected on the basis of their present distribution. Importantly, they show that Uromys occurred in regions during the Pleistocene that are today south of major modern biogeographic barriers for mesic taxa (e.g., the Burdekin and St Lawrence Gaps, see Bryant & Krosch, 2016).
The Middle Pleistocene extinct species U. aplini 1s phylogenetically positioned near the base of the Uromys (Uromys) clade between the geographically and ecologically restricted U. hadrourus and U. sherinni (Fig. 9). All taxa are found in northern Queensland, along the eastern seaboard. These taxa do not, however, form a resolved clade to the exclusion of species from New Guinea and its surrounding islands; therefore, it is hard to determine whether the Australian species are a monophyletic clade suggesting a single arrival and diversification. A reading of the current preliminary phylogenetic hypothesis would have the ancestor of Uromys dispersing from the Indo-New Guinea region
188 Records of the Australian Museum (2020) Vol. 72
and founding two, possibly parallel radiations that derive Uromys (Uromys) and Uromys (Cyromys). One lineage either diversified from isolation within, or has become restricted to, the Solomon Islands group, producing all of the members of Uromys (Cyromys) and their current biogeographic distribution. The other, arriving on mainland Australia, diversified first and then dispersed to the islands and mainland New Guinea with the most derived taxon reaching New Britain.
The Pleistocene record of U. aplini demonstrates that Uromys was present in Australia over 500,000 years ago, and occurred well south of the current biogeographical range of the genus, reaching at least the Mount Etna region by the Middle Pleistocene. Molecular-based data estimate that a U. hadrourus/U. caudimaculatus lineage extends back at least 1 million years, and possibly 2.5 million years ago to the Early Pleistocene (Watts & Baverstock, 1994; Bryant et al., 2011). Currently no Early Pleistocene fossil sites are known from north-east Queensland, while in north-west Queensland, the Early Pleistocene Rackham's Roost fauna from Riversleigh, though rich in xeric-adapted rodents, understandably lacks Uromys (Godthelp, 1999). Therefore, the mesic-adapted habitats were probably already restricted to the wetter eastern seaboard by the Early Pleistocene. Thus, arrival, speciation, isolation, and extinction of species of Uromys in eastern Queensland potentially occurred all within the last two million years.
Based on current palaeoclimatic and palaeoenvironmental proxies across continental Australia (Christensen et al., 2017) and more local Neogene records (Henderson & Nind, 2014), it 1s likely that corridors of mesic habitat were restricted to the eastern seaboard of Australia, including central-eastern and north-eastern Queensland, during the Quaternary. Therefore, connectivity of these mesic habitats would have been needed for ancestral Uromys to disperse southwards along the eastern seaboard to at least the Mount Etna region, subsequently producing U. aplini.
Local extinction of U. aplini occurred at Mount Etna sometime after 205-170 ka as the environment transitioned from closed wet rainforest to dry-adapted habitats (Hocknull et al., 2007). Sometime after this, U. caudimaculatus arrived in the region for the first time, likely dispersing southward along a route similar to that taken by the ancestor of U. aplini. The age of the U. caudimaculatus lineage is considered to be >1 Ma (Bryant et al., 2011) but the species has not been detected in the ^» 500—280 ka deposits at Mount Etna. This dispersal may have occurred sometime after the extinction of U. aplini (c. 205—170 ka), during a period of mesic return. Therefore, corridors of habitat must have existed to allow the dispersal of this taxon south to the Mount Etna region. Until its local extinction, Uromys caudimaculatus existed in this region after 50 ka but prior to the onset of the Last Glacial Maximum. The exact timing of the local extinction of U. caudimaculatus remains unresolved. Additional dating of layers containing this taxon could potentially refine this local extinction timeline.
Together, these two records of Uromys demonstrate multiple southern dispersals and subsequent local extinctions during the Pleistocene, with the likelihood that these dispersals required the crossing of several biogeographical barriers identified (Bryant & Krosch, 2016) along the eastern seaboard in north-east and central-eastern Queensland (Fig. 1).
Combining the spatio-temporal record of Uromys along with our preliminary phylogenetic hypothesis suggests that the mesic regions of the Australian mainland supported the initial radiation of Uromys (Uromys), with a separate earlier lineage diversifying into the taxa contained within Uromys (Cyromys) that possibly occupied the emergent Solomon Islands. Subsequent dispersal of the Australian clade throughout New Guinea is contrary to what would be expected on the basis of the species richness of Uromys currently found today throughout the New Guinea to Solomon Islands region, compared to that of mainland Australia. It is, however, recognized that throughout much of the Cenozoic, bias of mesic faunal extinction resulted in an overall shift of mainland Australian biomes toward more xeric-adaptation, thus mesic biomes are now significantly under-represented (Byrne ef al., 2011). The timing of these extinctions, and the effect of these on our understanding of present-day biogeography and phylogeography remains poor, without further study of the fossil record. Establishing the fossil record of these mesic biome lineages is crucial to understanding the timing and tempo of these biogeographical changes. Uromys represents just one group that can provide data on the evolution of this significant biome.
Palaeoecology of Uromys
Living species of Uromys are semiarboreal omnivores (Breed & Ford, 2007). The ability to access food resources in the canopy (e.g., fruits, before they fall to the forest floor) has been suggested as a competitive advantage for species of Uromys (Rader & Krockenberger, 2006); this probably played a role in resource partitioning in the species-rich Mount Etna Middle Pleistocene rainforest. The larger size of most species (U. hadrourus and U. porculus being exceptions) allows them to utilize food resources that are inaccessible to smaller rodents. For example, large species of Uromys in north Queensland are known to gnaw through the hard, thick shells of coconuts (Watts & Aslin, 1981) and are also infamous for opening metal traps (Elliot traps) to steal bait or prey upon smaller mammals (Laurance et al., 1993; Eric Vanderduys, pers. comm. January 2020). Furthermore, there is evidence that smaller murines actively avoid large species of Uromys (Leung, 2008) suggesting that an “ecology of fear" (Brown et al., 1999) may have a role in structuring small mammal assemblages, at least on a local scale. Uromys aplini is the largest murine in the Mount Etna deposits, and may have behaved much like its extant relatives, robbing large seeds, consuming fruits and insects, and generally terrorizing the smaller vertebrates.
Extinction of Uromys in central Queensland
The majority of rainforest-inhabiting species at Mount Etna became extinct after 280 ka (minimum age of site QML1313). But a small number of rainforest-adapted species, e.g., Dendrolagus sp. (Hocknull et al., 2007) and Antechinus yuna (Cramb & Hocknull, 2010) persisted for some tens of thousands of years, and appear in low numbers in QML1312, dated to 205—170 ka (Hocknull et al., 2007). Uromys aplini is one of these, and is represented by a single specimen in QML 1312. The possibility of this specimen being derived from faunal mixing (e.g., a time-averaged or reworked deposit) can be discounted as the assemblage of surviving rainforest taxa shows clear selection of certain
species. For example, multiple specimens of Antechinus yuna are present, yet Antechinus yammal is absent, despite these two species being ubiquitous in older rainforest assemblages (Cramb & Hocknull, 2010).
The late survival of U. aplini implies some degree of ecological flexibility, a reasonable proposition in light of the apparent ability of extant U. caudimaculatus to make use of a variety of habitats in north Queensland (Moore, 2008). Despite this adaptability, U. aplini disappeared from the local record prior to deposition of site QML1456 (<80 ka, Price et al., 2015). Uromys caudimaculatus appears intermittently in the lower, older spits of QML 1456, before apparently becoming locally extinct soon after 50 ka. The loss of both species may be explicable by an increasingly dry regional climate during the latter part of the Pleistocene and associated replacement of closed-canopy forests by open habitats. Despite a return to more mesic conditions during the Holocene, and deposits representing Holocene-aged accumulations, there is no evidence of Uromys returning to the Mount Etna area.
ACKNOWLEDGEMENTS. The authors wish to thank Kristen Spring and QM geosciences staff for curation of specimens, Heather Janetzki, Sandy Ingleby, Karen Roberts, Ken Aplin, and Fred Ford for access to comparative material, the Willi Hennig Society for providing phylogenetic software, Tyrone Lavery for providing an additional datum, Noel and Jeanette Sands and family for assistance in the field, all staff at Capricorn Caves including the Augusteyn family, for their support of palaeontological research, all researchers, honoraries, and volunteers involved in the Mount Etna project, and Liz Cramb for supporting her husband's palaeontology habit. Collection of material for this project was supported by the Ian Potter Foundation and ARC Linkage Grant (LP0453664).
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Cramb et al.: New Middle Pleistocene Uromys species 191
Appendix 1. List of modern specimens examined.
Uromys anak: AM M.15633, AM M.15634, AM M.15645, AM M.15646, AM M.15647, AM M.15753, AM M.15754, AM M.15854, AM M.16695, AM M.32337, AM M.38676, AM M.38702, AM M.38864, QMJM3838.
Uromys caudimaculatus: CM705, QMJ2344, QMJ5907, QMJ5908, QMJ6349, QMJ9304, QMJ9386, QMJ9387, QMJ9388, QMJ9389, QMJ9390, QMJ9454, QMJ9455, QMJ10131, QMJ10133, QMJ10134, OMJ11512, QMJ16181, QMJ16187, QMJ16450, QMJ16725, QMJ16768, QMJ16772, QMJM17309, QMJ17609, QMJ17610, QMJ17611, QMJ20347, QMJ22127, QMJ22538, QMJ22540, QMJ22606, QMJ22607, QMJ23023, QMJM1001, QMJM8738, QMJM10038, QMJM18470, QMJM21138.
Uromys hadrourus: QMJM504, QMJM21 73, QMJM8146. Uromys neobrittanicus. AM M.20689, NMVC6890. Uromys rex: AM M.13594.
Uromys sherrini: CM10822, QMJ5907, QMJ5908, QMJ8000, QMJ16725, QMJ17612, QMJ17613, QMJ17614, QMJ21272, QMJ22539, QMJ22606.
Records of the Australian Museum (2020) vol. 72, issue no. 5, pp. 193—206 https://doi.org/10.3853/j.2201-4349.72.2020.1723
Records of the Australian Museum
a peer-reviewed open-access journal
published by the Australian Museum, Sydney communicating knowledge derived from our collections ISSN 0067-1975 (print), 2201-4349 (online)
Late Quaternary Fossil Vertebrates of the Broken River Karst Area, Northern Queensland, Australia
GILBERT J. PRice'(®, JONATHAN CRAMB!@®, JULIEN Louvs?(9, KENNY J. TRAVOUILLON°*®, ELEANOR M. A. PEASE, YUE-XING FENG!@®, JIAN-XIN ZHAO!(®, AND DOUGLAS IRVIN*
' School of Earth and Environmental Science, The University of Queensland, Brisbane Qld 4072, Australia ? Australian Research Centre for Human Evolution, Griffith University, Brisbane Qld 4111, Australia ? Western Australian Museum, Locked Bag 49, Welshpool DC WA 6986, Australia
^ Hills Speleology Club Limited, Sydney, Australia
ABSTRACT. Two new fossil deposits from caves of the Broken River area, northeast Queensland, provide the first regional records of vertebrate species turnover and extinction through the late Quaternary. Fossil assemblages from Big Ho and Beehive Caves are dominated by small-bodied vertebrates, especially mammals. They represent owl roost deposits, although limited presence of larger-bodied taxa such as macropodids may be the result of occasional pitfall trapping. U-series dating demonstrates that Big Ho dates to the penultimate glacial cycle (c. 165 ka) and Beehive to the early Holocene (c. 8.5 ka). A total of 34 mammalian taxa were identified; within the two deposits, seven taxa are unique to Big Ho and another seven are found only in Beehive. The deposits also preserve five extinct fossil taxa (bandicoots and rodents) that add to a growing list of small-bodied species known to have suffered extinction in the late Quaternary. The deposits further yield the remains of four species of bandicoots and rodents (Chaeropus yirratji, Notomys longicaudatus, Conilurus albipes, and Pseudomys gouldii) that suffered extinction post- European colonization. These new fossil records represent significant increases in the known geographic and temporal range of several species and begin to fill an important gap in our understanding of the faunal history of tropical northeast Australia.
Introduction
Modern Australian ecosystems emerged during the Quaternary under a backdrop of major fluctuations in atmospheric carbon dioxide concentration, sea levels, and temperature, with a long-term trend towards progressively drier climates (Martin, 2006; Kershaw et al., 2003; Price, 2013). The period was marked not only by significant evolutionary events, but also major extinctions and geographic range shifts of many flora and fauna (e.g., Kershaw, 1994; Jordan et al., 1995; Reed & Bourne, 2000,
2009; Hocknull et a/., 2007; Prideaux et al., 2007; Price, 2012; Price et al., 2005; Black et al., 2014). Today, at a time of widespread awareness over detrimental anthropogenic and climatic impacts on Australian ecosystems, it has become critical to understand the history of ecosystem origins and responses to similar past events. The Quaternary fossil record has a significant role to play in yielding that crucial information (Reisinger et al., 2014).
While many vertebrate fossil deposits of Quaternary age have been recognized in Australia, the record is patchy and geographic coverage is strongly biased towards southern
Keywords: Quaternary; Pleistocene; mammal; marsupial; extinction; range shifts
Corresponding author: Gilbert J. Price g.price1@uq.edu.au
Received: 3 February 2020 Accepted: 27 August 2020 Published: 25 November 2020 (in print and online simultaneously) Publisher: The Australian Museum, Sydney, Australia (a statutory authority of, and principally funded by, the NSW State Government)
Citation: Price, Gilbert J., Jonathan Cramb, Julien Louys, Kenny J. Travouillon, Eleanor M. A. Pease, Yue-xing Feng, Jian-xin Zhao, and Douglas Irvin. 2020. Late Quaternary fossil vertebrates of the Broken River karst area, northern Queensland, Australia. In Papers in Honour of Ken Aplin, ed. Julien Louys, Sue O'Connor, and Kristofer M. Helgen. Records of the Australian Museum 72(5): 193—206.
https://doi.org/10.3853/j.2201-4349.72.2020.1723
Copyright: © 2020 Price, Cramb, Louys, Travouillon, Pease, Feng, Zhao, Irvin. This is an open access article licensed under a Creative Commons Attribution 4.0 International License (CC BY 4.0), which permits unrestricted use, distribution, and
e
reproduction in any medium, provided the original authors and source are credited.
194 Records of the Australian Museum (2020) Vol. 72
Netherwood Tonalite
Judea Formation
Study sites
Broken River Group
Charters Towers _ Formation ——
Figure 1. Map showing study sites and regional geological provinces of the Broken River karst area.
Australia. Very little is known about the Quaternary history of ecosystems in northern Australia (Price et al., 2017; Hocknull et al., 2020). This situation is not necessarily due to a paucity of fossil sites in the region, but more likely results from a dearth of field investigations; when these have been conducted, they have yielded records comparable to the south. Areas such as the Darling Downs (southeast Queensland) and Mt Etna (central eastern Queensland), for example, have produced some of the most extensive records of Quaternary vertebrates north of the Queensland— New South Wales border. There, records show waves of extinction of both megafauna and micro-fauna (e.g., rodents, bandicoots) alongside progressive decreases in precipitation and expansion of more open habitats through the late Quaternary (Hocknull, 2005a; Hocknull et al., 2007; Cramb & Hocknull, 2010a; Price & Hocknull, 2011; Price ef al., 2009; Price & Sobbe, 2005; Price & Webb, 2006; Price et al., 2015). While some Quaternary vertebrate fossils have been recovered from northern Australia (e.g., Archer ef
al., 1978; Molnar, 1981; Klinkhammer & Godthelp, 2015; Cramb et al., 2018), the records remain patchy, are mostly undated, and are usually one-off collections or reports of single species.
Here we describe new Quaternary fossil faunas from two limestone caves in the Broken River area, Greenvale, northeast Queensland (Fig. 1). Although Palaeozoic marine fossils have long been known in the area (see Henderson & Withnall, 2013 and references therein), Quaternary vertebrate fossils within the cavernous limestones were only reported in the 1980s and remained unstudied until the 2000s. Since then, individual reports of bilbies, bandicoots, rodents, and megafaunal taxa such as Diprotodon and Thylacoleo have been produced (Hocknull, 2005b; Cramb & Hocknull, 2010b; Price ef al., 2017; Travouillon et al., 2019). Full mammalian palaeocommunities from the region have yet to be documented. The aim of the present paper is to report on the first two (of several) fossil assemblages excavated from caves of the region.
Price et al.: Late Quaternary vertebrates of Australia 195
Figure 2. Images of the karst, study caves, and fossils of the Broken River karst area. (A) rillenkarren typical of the Broken River limestone karst; (B) Beehive fossil deposit (arrow indicating fossil-bearing breccia); (C) Big Ho fossil deposit (arrow indicating fossil-bearing breccia exposed as a false floor); (D) partially acid-digested breccia from Beehive showing high concentration of vertebrate fossils.
Geographic and geological settings
The Broken River area has been the subject of explorations by the Chillagoe Caving Club who are working on mapping many of the region's caves. The two deposits reported here are from Big Ho and Beehive, caves that formed in the main outcrop of limestone in the southern part of the Jack Formation, Graveyard Creek Group, part of the larger Broken River Province (Fig. 1). Asthis part ofthe outcrop is located on private property, specific locality details remain confidential but are available to bona fide researchers upon request to the Chillagoe Caving Club, Chillagoe, Queensland.
The Graveyard Creek Group is 150 m to > 5000 m thick and contains folded Silurian-Mississippian siliciclastic and carbonate sedimentary rocks, with the contained Jack Formation being around 580 m thick (Henderson & Withnall, 2012). The Jack Formation is dominated by limestone and mudstone rich in autochthonous fossils that include corals, molluscs, brachiopods, conodonts, and fish remains, among other taxa (Henderson & Withnall, 2012). The Jack Formation outcrop is heavily karstified and contains extensive and well-developed rillenkarren (Fig. 2A) making access to the caves particularly challenging. The formation is tilted to c. 90? and the caves are largely joint controlled, thus, contain many narrow but tall passages and caverns, including Big Ho and Beehive.
Materials and methods
Collection and curation
Fossil breccias were collected during a short fieldtrip in conjunction with the Chillagoe Caving Club in May 2012. The aim of the trip was to conduct a general survey of selected caves to assess their palaeontological significance. Both ofthe caves visited contain heavily lithified, fossil-rich breccias. Due to the high degree of lithification, the breccias could only be removed by breaking them into smaller blocks for transport out of the cave; more traditional excavation techniques (e.g., top-down excavations with small hand tools such as trowels) were not possible. The stratigraphic depths of the collected breccias varied from 50—70 mm for both deposits, with the breccia blocks weighing a total of approximately 12 kg. Stratification within both deposits was not evident, thus for the purpose of this study, are considered as two single, discrete accumulation phases. It is likely that the two assemblages are only minimally time-averaged and do not represent a large amount of time in terms of their depositional accumulation, respectively.
Breccia blocks from the two deposits were taken to The University of Queensland for digestion using weak (2—3%) acetic acid. The acid dissolved the carbonate cements and caused the blocks to break down, allowing the vertebrate
196 Records of the Australian Museum (2020) Vol. 72
fossils to be recovered. Higher concentrations of acid were initially trialled (S—10%) for small, single blocks, but caused too much damage to the contained fossils. Even at the lowest concentrations of acid, gastropods within the breccias were dissolved and thus could not be reported in this study. Following digestion, the loose sediments were wet sieved with 1 mm mesh and fossilized skeletal remains then sorted under microscopes and magnifier lamps. Two c. 500 g breccia blocks, one from each site, remain unprocessed and are retained in the collection as representative material of the original deposits.
Fossils were identified using typical comparative morphological techniques; minor but pertinent remarks concerning the taxonomic identifications are given in the results. We concentrate largely on the identification of mammals due to their abundance, degree of preservation, and ease of identification, although other vertebrates including frogs, squamates, and birds are represented in the deposits. These will be detailed in future reports.
We calculated the number of identified specimens present (NISP) and minimum number of individuals (MNI) for each taxon identified at the lowest taxonomic level possible. The skeletal element used for such calculations varied for each taxon. For frogs, we used pelves; monitor lizards used osteoderms; snakes used vertebrae; birds used humeri; dragon lizards, skinks, and mammals used teeth/maxillae/mandibular elements).
Fossils figured in this study are accessioned in the collec- tions of the Palaeontology Laboratory at The University of Queensland, Brisbane, Australia (accession abbreviation: UQPL, University of Queensland Palaeontology Lab- oratory). Additional fossil specimen accession abbreviation: QMF (Queensland Museum Fossil; Queensland Museum, Brisbane, Australia).
Dating
Analytical dating of the fossil-rich breccias is difficult as they are heavily lithified and thus not suited to common Quaternary methods such as luminescence dating. They also lack dateable charcoals that might be amenable for radiocarbon dating. The deposits also lack interbedded and capping flowstones and cannot be dated using U-series stratigraphic bracketing approaches. Thus, the only viable option was direct dating of in situ fossils using U-series. Fresh skeletal tissues typically lack U. However, post- burial diagenesis means that U is commonly taken up from burial sediments by apatite that scavenges U but excludes Thorium. Subsequent radioactive decay of the original U via alpha and beta emissions eventually leads to stable lead isotopes. However, the early part of the decay sequence to form ?*?Th (1.e., 25U—9*U—??Th) has a half-life of 500—600 ka, thus, making U-series a viable option for dating late Quaternary vertebrate fossils. The U-series age is calculated by determining the amount of daughter ?""Th to parent ?5U. In ideal situations, U is taken up rapidly from the sedimentary environment post-burial, and assuming that U has not subsequently leached from the bone, the resulting U-Th age will represent a minimum age for the fossil, but one that is close to the true age. Loss of U from the system post-uptake can lead to age overestimation making the fossil tissues unsuitable for dating. The mode of U uptake and potential of loss from the fossils can easily be determined by constructing ?*"Th and U-concentration profiles through the dated tissues.
Most of the fossils are from small-bodied vertebrate species (e.g., rodents) that required extraction from the breccias using dilute acetic acid. It was determined that such tissues may not be suitable for dating for two reasons: (a) it is difficult to construct U-profiles and thus test the reliability of the ages using such small specimens; and (5) there is a risk that acetic acid might strip the fossils of U rendering the specimens unsuitable for dating. Consequently, dating focused on broken cross-sections of bones from in situ larger-bodied species (macropodids) within the breccias that were not processed in acid baths. Multiple dating samples (powders) were drilled from one bone each in both deposits using a 1 mm diameter bit following procedures described in Price et al. (2013). The sampled bones are from approximately the middle of each of the sampled portion of the deposits. These samples were then prepared for U-series dating following techniques described in Zhao et al. (2009) and measured using a Nu Plasma multi-collector inductively coupled plasma mass spectrometer at the Radiogenic Isotope Facility, The University of Queensland, following the analytical procedures of Clark ef al. (2012).
Results and interpretation
Geology
The fossils were preserved in haematite-rich clay matrix- supported breccias. Both Big Ho and Beehive are particularly fossiliferous with clasts dominated by fragmentary small- bodied vertebrate remains. Larger clasts were rarely observed. The breccias are massive with no obvious sedimentary structures, including evidence of stratification, in hand samples collected.
The deposit in Beehive is from a small chamber known to cavers as the ^Greenroom". The deposit is at floor level, adjacent to a limestone wall and lacks associated speleothems (Fig. 2B). It is predominately preserved in phreatic niches along the cave wall. Only the top c. 70 mm of the deposit was collected considering the degree of exposure, lithification, and difficulty in excavating with only hand tools. The outcrop runs around 3 m horizontally and no more than c. 30 cm wide when measured from the wall. The total depth of the deposit is unknown.
The Big Ho breccia is located approximately 4 m above ground level in a large open chamber (Fig. 2C). Enough sunlight enters the chamber to allow an extensive groundcover of maidenhair ferns and mosses to grow. Like Beehive, deposits were concentrated in phreatic niches along the cave wall. Breccia scars at the same level on either side of the chamber indicates that the original deposit was extensive and likely formed the entire floor of the chamber. Remains of the in situ breccia now consist only of a false floor that juts out of the limestone wall by no more than 30 cm, and extends horizontally along the wall for approximately 1 m. The most likely mode of deposit formation is as follows: (a) sediment and bone entered the chamber and began to create the deposit; (5) the deposit then lithified into a breccia; and (c) subsequent erosion of the breccia from above and especially underneath created the false floor. The Big Ho breccia was more heavily cemented than the Beehive breccia, less extensively preserved, and topographically higher relative to current floor levels. Thus, on geological evidence and field observations we considered the Big Ho breccia to be substantially older than the Beehive breccia.
taxon
anuran gen. et sp. indet. 1 anuran gen. et sp. indet. 2 scincid gen. et sp. indet. agamid gen. et sp. indet. Varanus sp. indet.
pythonid gen. et sp. indet. elapid gen. et sp. indet. Aves gen. et sp. indet. Dasyurus sp.
Antechinus sp.
Phascogale tapoatafa Planigale sp. cf. ingrami/tenuirostris Sminthopsis macroura Sminthopsis sp. cf. murina Chaeropus yirratji
Isoodon peninsulae Isoodon sp. 2
Perameles sp.
Petaurus norfolcensis Trichosurus sp. ?hypsiprymnodontid macropodid indet. macropodid “small” Conilurus albipes Conilurus capricornensis Leggadina forresti Notomys longicaudatus Notomys sp. 2
Pseudomys australis Pseudomys sp. cf. P. delicatulus Pseudomys desertor Pseudomys gouldii Pseudomys gracilicaudatus Zyzomys sp. 1
Hydromys chrysogaster Melomys cervinipes
Rattus spp.
Rattus lutreolus “Microchiroptera” sp. indet. Miniopterus orianae
Price et al.: Late Quaternary vertebrates of Australia
common name
Frog
Frog
Skink
Dragon Lizard
Monitor Lizard
Python
Venomous snake
Bird
Quoll
Antechinus
Brush-tailed Phascogale Planigale
Striped-faced Dunnart Slender-tailed Dunnart Northern pig-footed Bandicoot Cape York Brown Bandicoot Short-nosed Bandicoot Long-nosed Bandicoot Squirrel Glider Brushtail Possum
Rat Kangaroo Macropod
Macropod
White-footed Rabbit Rat Capricorn Rabbit Rat Forrest’s Mouse Long-tailed Hopping Mouse Hopping Mouse
Plains Mouse
Delicate Mouse
Desert Mouse
Gould's Mouse
Eastern Chestnut Mouse Rock Rat
Water Rat
Fawn-footed Melomys Native Rat
Swamp Rat
Microbat
Bent-wing Bat
Table 1. Taxonomic list for vertebrates of the fossil deposits of the Broken River karst area.
Beehive
NISP
U —
oo MWe Wn Oo
ea
MNI
JI —
—
Big Ho
NISP
| dem oo oo
l I
N N
Nn Nn
A —
197
MNI
Taphonomy
Both deposits preserve very similar taphonomic signatures in that the breccias are heavily lithified, lack obvious stratification, and are dominated by the remains of small- bodied vertebrates. Prior to acid digestion (or at least in the early stage of acid digestion), bones within the breccias were mostly complete in situ (Fig. 2D). Where fragmentation was observed on skeletal elements, the broken ends lacked evidence of abrasion suggestive of re-working (e.g., Fig. 2D). Subsequent acid processing of the breccias led to higher fragmentation of the bones.
The breccias have a high concentration of skeletal remains (e.g., Fig. 2D). The majority of the fauna are nocturnal, non-cavernous species. The fossils lack tooth markings suggestive of predation from carnivorous mammals. They
bear the appearance of typical owl deposits like those of modern roosts observed elsewhere (e.g., Walton, 1990). Thus, the most parsimonious interpretation is that these assemblages were produced predominately by owls. Owls typically hunt at night either consuming their prey in the surrounding region or back in the caves. They will tear the prey apart to consume before regurgitating pellets that contain difficult to digest elements such as bones and teeth, fur and feathers. More robust elements (1.e., bone and teeth) are more resilient to post-burial diagenesis and readily make their way into the fossil record (Walton, 1990). Skeletal remains of larger-bodied taxa preserved within the breccias are unlikely to have been brought into the caves by owls, but rather, are likely victims of the caves acting as pitfall traps. Today, it is not unusual to see the remains of modern skeletons of macropodids (especially rock wallabies and
198 Records of the Australian Museum (2020) Vol. 72
wallaroos) lying on the floors of other caves in the area.
Forty taxa were identified from the two deposits, with 26 identified to species level (Table 1). Mammals (32 taxa) are the most abundant in terms of taxonomically identifiable remains. Most mammalian taxa occur in both deposits, although five are unique to Beehive and another seven are found only in Big Ho. Species richness is similar across deposits: Big Ho contains 27 species of mammal versus 25 for Beehive. To test for potential bias introduced by sampling intensity, we conducted a rarefaction analysis (Fig. 3) using PAST 4.0 (Hammer et al., 2001). For mammals, although Big Ho has more specimens identified to lowest taxonomic level possible than Beehive (372 vs 302), rarefaction diversity curves for both deposits tend towards an asymptote. When compared at a common number of specimens, 1.e., 302, both curves predict that 25 species would be present. This further highlights the taphonomic similarities between both deposits. It is likely that more collecting from both sites, especially Beehive, would yield a greater diversity of mammalian species. Pertinent taxonomic remarks for the Broken River taxa are given below.
c 20 g 15 — Big Ho (S 10 — Beehive
0 50 100 150 200 250 300 350 400 Specimens (n) Figure 3. Rarefaction curves comparing Big Ho and Beehive mammalian diversity within respective fossil breccias.
Fauna
Anura
Frogs (Fig. 4A,B) were identified on the basis of postcranial remains. Ilial morphologies indicate that at least two taxa are present.
Sauropsida
Squamates (Fig. 4C—G) were identified to family pre- dominantly on the basis of vertebrae, maxillae, and mandible fragments, revealing the presence of varanid, agamid, and scincid lizards, as well as elapid and pythonid snakes. Osteoderms and jaws were less commonly recovered.
Aves
Birds (Fig. 4H,I) are abundant in both study sites, with a variety of sizes and morphologies hinting at a sizeable diversity of taxa. These specimens await future study.
Mammalia Marsupialia Dasyuridae
Dasyurus sp. indet. (Fig. 5A). A medium-sized quoll is represented by isolated molars. Dimensions of the molars indicate that it is within the size range of D. viverrinus and D. geoffroii, but the available material is insufficient to separate these species.
Antechinus sp. indet. (Fig. 5B). The identification of fossil Antechinus is discussed in Cramb & Hocknull (20102). Antechinus is a rare taxon at Broken River, represented by mandibular fragments and isolated teeth. Most species of Antechinus have posterior cingula on M'?, and this is the distinguishing feature of the Broken River specimens. The exact identity of the Broken River specimens cannot be established on the basis of available material.
Phascogale tapoatafa (Fig. 5C). The Brush-tailed Phascogale is a medium-sized dasyurid with a reduced P}, posterior cingula on the upper molars, commonly buccal cingula on M!, and relatively strong buccal cingulids on the lower molars. It is distinguished from other species (P. pirata, P. calura, and an undescribed species from Mount Etna) by being larger and having stronger buccal cingula on the upper molars.
Planigale sp. cf. P. ingrami/tenuirostris (Fig. 5D). Archer's (1976) revision of Planigale found that the most reliable dental features separating species were size and presence/ absence of a P?/. One specimen from Big Ho has a lower molar row measuring 4.05, within the ranges of P. ingrami and P. tenuirostris, and equidistant between the means for both species as given by Archer (1976).
Sminthopsis macroura (Fig. 5E). The Stripe-faced Dunnart is medium-size (for a species of Sminthopsis), and possesses large entoconids on M, that do not contact the hypocristids (Archer, 1981).
Sminthopsis sp. cf. S. murina (Fig. 5F). The majority of Sminthopsis specimens in the study sites lack entoconids on M,.. They are assigned to S. murina here, although distinguishing that species from other “entoconid-less” species of Sminthopsis is difficult (Cramb et al., 2009).
Chaeropodidae
Chaeropus yirratji (Fig. 5G). Three specimens from Big Ho and four from Beehive represent the Northern Pig-footed Bandicoot and were included in the description of the species in Travouillon et al. (2019). Characters on the M! and M, are diagnostic and help separate it from Chaeropus ecaudatus. These include the paracone connecting to StB (stylar cusp B) only on M! (not StA, in unworn teeth); the metaconule on M!” is well-developed, making molars more rectangular in shape; StD present on M! and the paracristid on M, does not connect to the protoconid (in unworn teeth).
Peramelidae
Isoodon sp. (Fig. 5H). This taxon is recorded only in Big Ho and is represented only by isolated lower molars (M, and M2). All measurements are within the modern range for I. macrourus torosus, but at the smaller end of the range. There are very few diagnostic characters on lower molars of
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Figure 4. Non-mammal vertebrates of the Broken River karst area. (4—5) anuran pelves (UQPL 1-2); (C) scincid dentary (UQPL3); (D) agamid upper jaw (UQPLA); (E) varanid osteoderm (UQPLS5); (F) pythonid vertebra (UQL6; (G) elapid vertebra (UQPL 7); (H—/) avian humeri (UQPL8-9). Scale bars = 1 mm.
Figure 5. Dasyurids and bandicoots of the Broken River karst area. (4) Dasyurus lower molar (UOPL10); (B) Antechinus sp. dentary (UQPL11); (C) Phascogale tapoatafa lower molar (UQPL12); (D) Planigale sp. cf. P. ingrami/tenuirostris dentary (UQPL13); (E) Sminthopsis macroura dentary (UQPL14); (F) Sminthopsis sp. cf. S. murina dentary (UQPL15); (G) Chaeropus yirratji maxilla (OMF58987); (H) Isoodon sp. lower molar (UQPL17); (J) Isoodon peninsulae mandible (UQPL 16); (J) Perameles sp. upper molar (UQPL18). Scale bars = 1 mm.
200 Records of the Australian Museum (2020) Vol. 72
Isoodon. Despite being clearly a specimen of /soodon, and being too large to be any other species than 7. macrourus, there are a number of differences compared to the modern taxon. The paraconid on M, is large and in line with the metaconid (the paraconid is commonly reduced and more buccally positioned in all described species of /soodon). The posthypocristid is not perpendicular to the toothrow on either the M, or M», which is a feature typically seen in Isoodon auratus, I. fusciventer, and I. peninsulae but not in I. macrourus. It is an undescribed species of /soodon. We do not erect a new species here, but rather wait on the discovery of additional, more comprehensively diagnosable material.
Isoodon peninsulae (Fig. 5I). The Cape York Brown Bandicoot is the most common species of bandicoot present in both Big Ho and Beehive. The dental measurements are slightly larger than the modern range for this taxon, but still below that of 7. macrourus. The overall morphology matches that of /. peninsulae. It differs from 1. auratus, I. fusciventer, and 7. macrourus in having a preparacrista of M! buccally orientated then posterobuccally orientated. It differs from 7. fusciventer and I. macrourus in having a stylar crest present on M! and no StC/D on M*. It differs from 7. fusciventer, I. macrourus, and I. obesulus in having a small anterior cingulum on M! not connected to the talon, and a large anterior cingulum of M? not connected to talon. It differs from I. macrourus and I. obesulus in having the posthypocristid of M, and M, oblique to the tooth row.
Perameles sp. (Fig. 5J). This taxon is only represented by two isolated molars, a left M! and a left M5, both from Big Ho. The M! length is within the range of modern Perameles pallescens, but the tooth is wider than in the modern species. The M, width is also within the modern range, but the length is longer. The M, typically has no reliable characters to identify specimens to known species (see Travouillon, 2016). The M! has several characters which are diagnostic: it differs from Perameles bougainville, P. fasciata, P. myosuros, P. notina, and P. gunnii in having a short stylar crest not connected to StD. It differs from Perameles eremiana, P. fasciata, P. myosuros, P. notina, and P. papillon in that the preparacrista not reconnecting to the postparacrista posteriorly. It differs from Perameles bougainville, P. eremiana, P. fasciata, P. myosuros, and P. papillon in having StB and StC distinguishable with StC larger than StB. It differs from all Perameles except P. eremiana and P. nasuta in having a very small StA. It differs from P. nasuta and P. pallescens in having a postprotocrista that ends posteriorly to metacone. It differs from P. fasciata in having no anterior cingulum. While there are enough characters to separate it from all modern taxa, it is not described as a separate species here, as it cannot be compared to the extinct fossil species Perameles sobbei, from which no M! has been recovered to date. The M, matches the morphology of P. sobbei (e.g., Price, 2002), but it also matches that of other Perameles, such as P. pallescens. As a result, we consider this taxon as Perameles sp. until further material is recovered, but it is undoubtedly an extinct species.
Petauridae
Petaurus norfolcensis (Fig. 6A). The Squirrel glider is a medium-sized Petaurus distinguished by having molar rows longer than those of P. breviceps and P. biacensis but shorter than P. australis, P. gracilis, and P. abidi.
Phalangeridae
Trichosurus sp. indet. (Fig. 6B). Phalangerids are easily distinguished from similarly-sized pseudocheirids by their bunodont molars.
Few morphological characters separate the species of Trichosurus, but T. vulpecula 1s extant in the Broken River area and is thus considered to be the most likely identity of the fossil specimens. One specimen from Big Ho is an unerupted molar, indicating that the individual was a juvenile.
Hypsiprymnodontidae
cf. Hypsiprymnodontidae gen. et sp. indet. (Fig. 6C). An isolated premolar fragment has the distinctive “buzz- saw" shape and ridges seen in hypsiprymnodontids, some burramyids, and propleopines. The size of the specimen is similar to Hypsiprymnodon, but its generic and specific identity is unknown.
Figure 6. Diprotodonts from the Broken River karst area. (4) Petaurus norfolcensis upper molar (UQPL19); (B) Trichosurus sp. upper molar (UQPL20); (C) ?hypsiprymnodontid premolar (UQPL21); (D) juvenile macropodid mandible (UQPL22). Scale bars = 1 mm.
Macropodidae
Macropodidae indet. (Fig. 6D). Macropodid remains are uncommon, fragmentary, and appear to be from immature individuals. A partial mandible from Beehive represents a very young individual, most likely from a rock wallaby (Petrogale sp.) although additional material 1s required to confirm the identification.
Placentalia Muridae
Rats and mice are most readily identified to species by their upper molars and maxillae, which form the majority of referred specimens here.
Conilurus albipes (Fig. 7A). Distinguishing features of the species are listed by Cramb & Hocknull (2010b). The Broken River sites are the northern-most records of this species.
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Figure 7. Placentals from the Broken River karst area. (4) Conilurus albipes maxilla (UQPL23); (B) Conilurus capricornensis molar fragment (UQPL24); (C) Leggadina forresti maxilla (UQPL25); (D) Notomys longicaudatus maxilla (UQPL26); (E) Notomys sp. 2 maxilla (UQPL27); (F) Pseudomys australis maxilla (UQPL28); (G) Pseudomys sp. cf. P. delicatulus maxilla (UQPL29); (H) Pseudomys desertor molar (UQPL30); (D) Pseudomys gouldii maxilla (UQPL31); (J) Pseudomys gracilicaudatus maxilla (UQPL32); (K) Zyzomys