Search

Peramelidae Gray, 1825 CONTENTS: Echymipera, Isoodon, Microperoryctes, Perameles (fig. 39), Peroryctes, and Rhynchomeles. STEM AGE: 16.5 Mya (95% HPD: 12.4–20.6 Mya). CROWN AGE: 8.8 Mya (95% HPD: 6.3–12.3 Mya). UNAMBIGUOUS CRANIODENTAL SYNAPOMORPHIES: Orbitosphenoid laterally inapparent or absent (char. 15: 0→1; ci = 1.000), and dP3 very small, nonoccluding, and structurally simplified, or absent (char. 120: 1→2; ci = 0.118). COMMENTS: Our dated Bayesian total-evidence analysis recovers monophyly of Peramelidae sensu Van Dyck and Strahan, 2008 (= Echymiperinae + Peramelinae + Peroryctinae), to the exclusion of Thylacomyidae and Chaeropodidae (fig. 33), as do our undated Bayesian total-evidence analysis (fig. 32) and all our molecular analyses (figs. 27–29). As already discussed, a similar topology has been recovered in most recent molecular and total-evidence analyses (Westerman et al., 2012; Kear et al., 2016; Travouillon and Phillips, 2018). Of the two craniodental features that optimize as unambiguous synapomorphies of Peramelidae, a laterally inapparent or absent orbitosphenoid is unique to this clade and shows no homoplasy among our terminals, whereas presence of a structurally simplified dP3 serves to distinguish peramelids from Macrotis and Chaeropus, in which dP3 is better developed. However, the morphology of dP 3 in † Bulungu, † Galadi and † Yarala, which fall outside the crown clade (Perameloidea) in our analyses, is currently unknown. † Crash bandicoot from the middle Miocene AL90 site (which has been radiometrically dated as ~14.8 Mya; Woodhead et al., 2014) at Riversleigh World Heritage Area was described by Travouillon et al. (2014a) as a member of the peramelid crown group. In support of this, † Crash has been recovered in a clade with Perameles and Isoodon in some recent analyses (Travouillon et al., 2014 a, 2015b: fig. 12a; Chamberlain et al., 2015), although not others (Travouillon et al., 2015b: fig. 12b; Kear et al., 2016). However, † Crash considerably predates our late Miocene estimate for the most recent common ancestor of Peramelidae. Apart from † Crash, the oldest known peramelids are probably early Pliocene taxa that have been referred to Perameles (P. † allinghamensis, P. † bowensis, and P. † wilkinsorum; Warburton and Travouillon, 2016; Travouillon et al., 2017)., Published as part of Beck, Robin M. D., Voss, Robert S. & Jansa, Sharon A., 2022, Craniodental Morphology And Phylogeny Of Marsupials, pp. 1-353 in Bulletin of the American Museum of Natural History 2022 (457) on pages 215-217, DOI: 10.1206/0003-0090.457.1.1, http://zenodo.org/record/6971356, {"references":["Gray, J. E. 1825. An outline of an attempt at the disposition of the Mammalia into tribes and families with a list of the genera apparently appertaining to each tribe. Annals of Philosophy 10 (2): 337 - 344.","Van Dyck, S., and R. Strahan (editors). 2008. The mammals of Australia, 3 rd ed. Sydney: New Holland Publishers.","Westerman, M., et al. 2012. Phylogenetic relationships of living and recently extinct bandicoots based on nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 62 (1): 97 - 108.","Kear, B. P., K. P. Aplin, and M. Westerman. 2016. Bandicoot fossils and DNA elucidate lineage antiquity amongst xeric-adapted Australasian marsupials. Scientific Reports 6: 37537.","Travouillon, K. J., and M. J. Phillips. 2018. Total evidence analysis of the phylogenetic relationships of bandicoots and bilbies (Marsupialia: Peramelemorphia): reassessment of two species and description of a new species. Zootaxa 4378 (2): 224 - 256.","Woodhead, J., et al. 2014. Developing a radiometricallydated chronologic sequence for Neogene biotic change in Australia, from the Riversleigh World Heritage Area of Queensland. Gondwana Research 29 (1): 153 - 167.","Travouillon, K. J., S. J. Hand, M. Archer, and K. H. Black. 2014 a. Earliest modern bandicoot and bilby (Marsupialia, Peramelidae and Thylacomyidae) from the Miocene of the Riversleigh World Heritage Area, northwestern Queensland, Australia. Journal of Vertebrate Paleontology 34 (2): 375 - 382.","Yates, A. M. 2015 b. New craniodental remains of Wakaleo alcootaensis (Diprotodontia: Thylacoleonidae) a carnivorous marsupial from the late Miocene Alcoota Local Fauna of the Northern Territory, Australia. PeerJ 3: e 1408.","Chamberlain, P. M., K. J. Travouillon, M. Archer, and S. J. Hand. 2015. Kutjamarcoot brevirostrum gen. et sp. nov., a new short-snouted, early Miocene bandicoot (Marsupialia: Peramelemorphia) from the Kutjamarpu Local Fauna (Wipajiri Formation) in South Australia. Alcheringa: an Australasian Journal of Palaeontology 40 (2): 197 - 206.","Travouillon, K. J., M. Archer, S. J. Hand, and J. Muirhead. 2015 b. Sexually dimorphic bandicoots (Marsupialia: Peramelemorphia) from the Oligo-Miocene of Australia, first cranial ontogeny for fossil bandicoots and new species descriptions. Journal of Mammalian Evolution 22: 141 - 167.","Warburton, N. M., and K. J. Travouillon. 2016. The biology and palaeontology of the Peramelemorphia: a review of current knowledge and future research directions. Australian Journal of Zoology 64 (3): 151 - 181.","Travouillon, K. J., et al. 2017. A review of the Pliocene bandicoots of Australia, and descriptions of new genus and species. Journal of Vertebrate Paleontology: e 1360894."]}

The current literature on marsupial phylogenetics includes numerous studies based on analyses of morphological data with relatively limited sampling of Recent and fossil taxa, and many studies based on analyses of molecular data that include a dense sampling of Recent taxa, but relatively few that combine both data types. Another dichotomy in the marsupial phylogenetic literature is between studies that focus on New World taxa, others that focus on Sahulian taxa. To date, there has been no attempt to assess the phylogenetic relationships of the global marsupial fauna, based on combined analyses of morphology and molecular sequences, for a dense sampling of Recent and fossil taxa. For this report, we compiled morphological and molecular data from an unprecedented number of Recent and fossil marsupials. Our morphological data consist of 180 craniodental characters that we scored for 97 species representing every currently recognized Recent genus, 42 additional ingroup (crown-clade marsupial) taxa represented by well-preserved fossils, and 5 outgroups (non-marsupial metatherians). Our molecular data comprise 24.5 kb of DNA sequences from whole-mitochondrial genomes and six nuclear loci (APOB, BRCA1, GHR, RAG1, RBP3 and VWF) for 97 marsupial terminals (the same Recent taxa scored for craniodental morphology) and several placental and monotreme outgroups. The results of separate and combined analyses of these data using a wide range of phylogenetic methods support many currently accepted hypotheses of ingroup (marsupial) relationships, but they also underscore the difficulty of placing fossils with key missing data (e.g., †Evolestes), and the unique difficulty of placing others that exhibit mosaics of plesiomorphic and autapomorphic traits (e.g., †Yalkaparidon). Unique contributions of our study are (1) critical discussions and illustrations of marsupial craniodental morphology, including descriptions and illustrations of some features never previously coded for phylogenetic analysis; (2) critical assessments of relative support for many suprageneric clades; (3) estimates of divergence times derived from tip-and-node dating based on uniquely taxon-dense analyses; and (4) a revised, higher-order classification of marsupials accompanied by lists of supporting craniodental synapomorphies. Far from the last word on these topics, this report lays the foundation for future research that may be enabled by the discovery of new fossil taxa, better-preserved material of previously described taxa, novel morphological characters, and improved methods of phylogenetic analysis.

International audience; The expedition commanded by the Frenchman Nicolas Thomas Baudin aboard the ships Le Géographe and Le Naturaliste (and Le Casuarina for the return journey) to the southern hemisphere between 1800-1804 collected specimens from numerous locations including the Canary Islands (Tenerife), Île de France (Mauritius), Cape Town (South Africa), Australia and Timor. Additionally, specimens were donated or purchased from locations not visited including the Comoros, Madagascar and Sumatra. Unfortunately, Baudin died at Île de France on the return trip so the responsibility of the account of the voyage was given to other members of the expedition. Responsibility for writing up the primary account of the voyage was granted to François Péron, who published the first volume of the narrative of the Voyage de Découvertes aux Terres Australes in 1807. Following his death in 1810, the second volume of the narrative was completed by Louis de Freycinet and published in 1816. The other four volumes of the Voyage included three atlases (the first by expedition artists Charles-Alexandre Lesueur and Nicolas-Martin Petit in 1807 and the second and third by Freycinet in 1811 and 1812), and a volume entitled Navigation et Géographie by Freycinet in 1815. Based on recent and on-going research, a review of many of the original documents is presented here, revealing hitherto unpublished details about who collected and donated mammals to the expedition. Research was conducted mainly in the collections of the Muséum national d'Histoire naturelle (MNHN) in Paris and their associated acquisition books. The Baudin expedition was responsible for an unprecedented collection of over 100 000 specimens of natural history, which remains the single largest collection of natural history specimens from Australia. A total of 101 mammal taxa relating to the Baudin expedition were identified during this study, which included 51 species described as a result of the expedition and 50 species that were described either before or subsequently, but not associated with the expedition. Of the taxa described, 20 species and three subspecies are currently recognised valid. During this study 43 museum specimens that were referable to 29 taxa were identified and at least five specimens seem to have been misplaced based on the information available. These specimens were derived from 24 holotypes, four paratypes, three syntypes, five lectotypes, and six paralectotypes and one topotype that are currently held at the MNHN. Some of these specimens are part of the estimated 51 mammals that were brought back alive to France on the boats. Charles-Alexandre Lesueur's illustrations complemented this study; of the 177 that are held at the Muséum d'Histoire naturelle du Havre, 149 could be attributed to one or more species and 28 were of unidentified species.

Perameles nasuta É. Geoffroy Saint-Hilaire, 1804 (Figs 10; 78; 82) Perameles nasuta É. Geoffroy Saint-Hilaire, 1804b: 62; pl. 44. COMMON NAME. — Southern Long-nosed Bandicoot. CURRENT NAME. — Perameles nasuta É. Geoffroy Saint-Hilaire, 1804. See Jackson & Groves (2015: 88). COLLECTOR/ S. — Charles-Alexandre Lesueur, contra Julien-Laferrière (1994: 11) who mentioned both Péron and Lesueur as its collectors. COLLECTION LOCALITY. — Near Port Jackson (Sydney), New South Wales, Australia (Julien-Laferrière 1994: 11). COLLECTION DATE. — Between 3 July and 18 November 1802. SPECIMEN NUMBER/S. — MNHN-ZM-MO-1990-417 (Fig. 10), holotype, skin mount (de Beaufort 1966: 524; Julien-Laferrière 1994: 11). OTHER NUMBER/S. — Anc. Cat. Gal. 215, Nouv. Cat. Gal. 327 (de Beaufort 1966: 518-519). ILLUSTRATIONS. — Illustrated by Charles-Alexandre Lesueur – Muséum d’Histoire naturelle du Havre, Inv. no. 80276 (Fig. 78; second from left - skull) and Inv. no. 80080 (Fig. 82; two living animals)., Published as part of Jackson, Stephen M., Jansen, Justin J. F. J., Baglione, Gabrielle & Callou, Cécile, 2021, Mammals collected and illustrated by the Baudin Expedition to Australia and Timor (1800 - 1804): A review of the current taxonomy of specimens in the Muséum national d'Histoire naturelle de Paris and the illustrations in the Muséum d'Histoire naturelle du Havre, pp. 387-548 in Zoosystema 43 (21) on page 402, DOI: 10.5252/zoosystema2021v43a21, http://zenodo.org/record/5142972, {"references":["JACKSON S. M. & GROVES S. M. 2015. - Taxonomy of Australian Mammals. CSIRO Publishing, Melbourne, 529 p.","BEAUFORT F. DE 1966. - Catalogue des types de mammiferes du Museum national d'Histoire naturelle. Paris. VI. Monotremata. - VII Marsupialia. Bulletin du Museum national d'Histoire naturelle. Section B. Botanique, Biologie et EcologieVegetables Phytochimie (2) 38: 509 - 553. https: // www. biodiversitylibrary. org / page / 55121975"]}

Didelphis obesula Shaw, 1797 (Figs 78; 81) Didelphis obesula Shaw, 1797: 298. COMMON NAME. — Southern Brown Bandicoot. CURRENT NAME. — Isoodon obesulus (Shaw, 1797). See Jackson & Groves (2015: 85). COLLECTOR/S. — Specimen apparently not collected. COLLECTION LOCALITY. — No data. COMMENT. — Occurs in southern Australia. COLLECTION DATE. — No data. SPECIMEN NUMBER/S. — No data. ILLUSTRATIONS. — Illustrated by Charles-Alexandre Lesueur – Muséum d’Histoire naturelle du Havre, Inv. no. 80276 (Fig. 78; skull, second from right) and Inv. no. 80081 (Fig. 81; two living animals of which one has a pouch young., Published as part of Jackson, Stephen M., Jansen, Justin J. F. J., Baglione, Gabrielle & Callou, Cécile, 2021, Mammals collected and illustrated by the Baudin Expedition to Australia and Timor (1800 - 1804): A review of the current taxonomy of specimens in the Muséum national d'Histoire naturelle de Paris and the illustrations in the Muséum d'Histoire naturelle du Havre, pp. 387-548 in Zoosystema 43 (21) on page 402, DOI: 10.5252/zoosystema2021v43a21, http://zenodo.org/record/5142972, {"references":["SHAW G. 1797. - The Naturalist's Miscellany: or coloured figures of natural objects, drawn and described immediately from nature. F. P. Nodder & Co., London. Volume 8: Text to plate 298. https: // www. biodiversitylibrary. org / page / 59277553","JACKSON S. M. & GROVES S. M. 2015. - Taxonomy of Australian Mammals. CSIRO Publishing, Melbourne, 529 p."]}

Perameles auratus Ramsay, 1887 Proc. Linn. Soc. N.S.W. (ser. 2) 2 (3), 551. (30th November 1887). Description read at the 31st August 1887 meeting of the LSNSW. Current name. Golden Bandicoot Isoodon auratus (Ramsay, 1887) following Jackson & Groves (2015). Taxonomic status. Recognised as either a species or subspecies of I. obesulus (Shaw, 1797) throughout most of the past century, but often accepted as a valid species in recent decades, see Jackson & Groves (2015). Syntype. M.468, adult male, skull extracted, puppet skin with large scrotum and tail stub (Fig. 3). In 2016, an old label, evidently Ramsay’s original, was found attached to skin M.468 and states “type of the species” and is initialed “Ed.P.R.” [= E. P. Ramsay], see Fig. 1c. That tag has subsequently been detached from the skin. A skull and mandibles labeled M.468 currently associated with skin M.468 is a possible mismatch (Fig. 4), see Comments below. The skull marked M.468 has fully erupted and worn dentition and is intact other than missing the right zygomatic arch and is the skull illustrated in figs 62–65 of Lyne & Mort (1981) as being the “type”. Mahoney & Ride (1988) refer to M.468, study skin and skull as “the holotype ”, which could be interpreted as lectotype designation by inference of a holotype (Article 74.6, the Code), given the possibility that syntypes exist. We leave lectotype designation to those undertaking taxonomic research. We do not recognise the skull currently associated with skin M.468 as being the syntype skull. Its association with the skin requires confirmation by a specialist in Isoodon taxonomy or might be resolved using DNA sequencing. Other material. M.469, female, study skin. Skin tag has “skin (). skull () on M368?”, the latter presumably an error for M.468. A skull labelled M.469 (Fig. 4), fourth molars not fully erupted, is thus doubtfully associated with skin M.469. The MAMU Register entry for M.469, done in the 1960s, states, “skin only, skull attached to M.468?”. Since that entry, a second skull has apparently been found in the Collection and doubtfully matched with skin M.469 but the basis of this match, when it was done, and by whom is not recorded in the database. The type status of skin M.469 and associated skull remains undetermined. It might be the female skin from Derby listed in the MAMU 1890s Catalogue but it might be a third, previously overlooked female for which no collector or date are recorded, see Comments below. Type locality. “The neighbourhood of Derby, N.W. Australia ” (Ramsay), Western Australia. Comments. Ramsay’s account focused entirely on a specimen with a tail stub for which he does not indicate the sex. Study skin M.468 (a male) is almost certainly that specimen. Ramsay’s original tag was still attached to skin M. 468 in 2016 but the skull currently associated with that skin is probably mismatched from another individual, as yet unidentified. Two main issues impede identification of type material of I. auratus in the Collection. First, is male skin M.468 a holotype or syntype?; second, which skull currently in the Collection, if any, belongs with type skin M.468? We have not resolved the second issue but identify key aspects for further investigation. The 1890s Catalogue lists only two specimens of I. auratus, adult skins of a male and female from Derby (the collector is not given). The male skin is assumed to be M.468 but the identity of the female skin now in the Collection is less clear. It might be the female listed in the Catalogue or an undocumented third individual, as discussed later. The female skin listed in the Catalogue was either examined by Ramsay or received by Macleay after Ramsay’s presentation of the account at the August 1887 meeting of the LSNSW. It might have arrived in the second consignment of specimens from Derby at an undermined date in 1887, or in 1888 when Froggatt returned to Sydney. We have not found documentation of the number and dates of shipments of natural history material sent from Derby by Froggatt to Macleay. The two references that we have found are both vague and ambiguous. Mammals collected by Froggatt are mentioned at two meetings of the LSNSW held in 1887. Anonymous (1888: 981) records that at the 26th November 1887 meeting, Masters exhibited mammals recently received that Froggatt had sent from Derby, that included “Two specimens of Hapalotis Boweri Ramsay; Perameles auratus, Ramsay; Perameles n. sp. (?); ….”. Crucially, we cannot be certain that Masters exhibited only those mammals obtained from the second shipment, as he might have exhibited a mixture from both shipments. It is also unclear whether “ Perameles auratus ” refers to one or more specimens. Further, whether “ Perameles n. sp. (?)” refers to M.469 or an entirely different specimen of another species, perhaps later deemed not be a new species. In a paper presented at the December 1887 meeting, Macleay (1888) noted that he had so far received two shipments of Froggatt’s Derby material. Although he stated that the first shipment from Froggatt was noted in the June meeting of the LSNSW, the first mention of Froggatt’s Derby material in the Proceedings was at the August meeting. At the December 1887 meeting, Macleay (1888) stated that he had “only just” received the second shipment and that “mammals of this collection were exhibited at our last meeting”, referring to the meeting of 30th November 1887. This might imply, though not necessarily, that additional material of I. auratus arrived after Ramsay’s description was presented at the meeting of 31st August 1887 but it is also possible that Ramsay initially received several specimens of I. auratus. The fact that Ramsay did not provide a description of an intact tail implies that he did not have another specimen with a tail. Alternatively, he might have focused on one specimen and ignored others. Ramsay has a track record of abbreviated mammal descriptions that fixate on describing one specimen and little else, even for species later determined to involve a type series. In any case, the possibility remains that Ramsay also examined an adult female. Doubt about whether Ramsay had a syntype series is resolved by adoption of Recommendation 73F of the Code, i.e. an assumption of syntypes when more than one specimen is suspected. The only practical consequence is whether skin M.468 is recognised as a holotype or lectotype. Locating the skull belonging to type skin M.468 remains a key issue that we have not resolved. The MAMU 1890s Catalogue does not indicate whether the skulls were extracted from the two skins. It is not known if Ramsay extracted a skull for his description but it seems likely that he did. He did not provide a description or measurements of the skull but was able to examine the rear molars. The jaws of bandicoots do not open wide enough for inspection of the rear molars unless the cheeks are cut or the skull extracted (Dr Kenny Trevouillon pers. comm.). Establishing the correct skull match with type skin M.468 and the identity of the second skin is complicated by the prospect, overlooked prior to our study, that a third I. auratus specimen in alcohol was in the Collection prior to the 1920s. The MAMU Donations Book entry for 27th September 1926 lists one I. auratus sent to the AM for skull extraction and identification (skin and skull returned 12th November 1929), along with two I. macrourus. The annotation next to the I. auratus entry in September 1926 states “Parcel from Tank”, suggesting that the specimen was removed from an alcohol storage tank. We interpret “parcel” as a specimen wrapped in cloth, making it more likely to have been overlooked when the Catalogue was prepared in the 1890s. This suggests that either the entry for the female skin in the MAMU 1890s Catalogue had neglected to indicate “spirit”, or that an additional female specimen was in the Collection. If there was a third specimen in alcohol, the collector and locality have not been recorded. Two skins but only one skull of I. auratus were registered in the 1960s and were probably assumed to be the two individuals listed in the 1890s Catalogue. A second unlabeled and unnumbered skull was found in the Collection at an unknown date after registration. If three specimens (a male and two females) were in the Collection prior to the 1920s, then one skin and one skull are now missing. The missing skin and skull could have been from two different individuals, including the skull belonging to male type skin M.468, given that skull morphology of I. auratus is not sexually dimorphic (Lyne & Mort 1981) and skulls cannot be assigned to sex. Skin M. 469 might be the one from Derby listed in the 1890s Catalogue, or the alcoholic female turned into a skin at the AM but for which the locality was not recorded. The specimen might have been collected by Froggatt, but other scenarios, though less likely, cannot be excluded given the poor documentation of specimens to and from the Collection after the early 1880s. The specimen might have entered the Collection at any time from the 1880s to the early 1920s. Froggatt donated mammal specimens that he collected in 1887 from north west Australia to the AM in 1932 and might also have donated material to the MAMU, for which the documentation has not been located. Alternatively, it might have been an exchange from another source for which the documentation has been lost. A further consideration is that the two I. auratus specimens now in the Collection possibly represent different taxa. Travouillon & Phillips (2018) recognised two morphologically distinct forms of I. auratus, a “Kimberley” form and an inland form. Froggatt had collected extensively in both inland and subcoastal areas by early September 1887 (Froggatt 1934) and would have collected within the distribution of both suspected forms of I. auratus. Both specimens of I. auratus in the MAMU were examined by Dr Kenny Travouillon (Western Australian Museum, pers. comm. 2016) as part of ongoing taxonomic research. He suggests, based on skin and skull morphology, that the smaller skull currently labelled M.469 could belong with skin M.468, but this requires further research into morphological variation in populations of I. auratus. Skull measurements of the “typical specimen” (= type specimen) of I. auratus reported by Thomas (1904: 228) were provided by E.R. Waite (then of the Australian Museum) and these dimensions are consistent with skull M.469, see Table 2. Although the measurements given in Table 2 suggest a trivial size difference between the skulls, M.469 is clearly the smaller skull and a good match to those of Waite (Fig. 4). It is clear that Waite measured skull M.469 but whether that skull was from the male or female skin(s) remains unknown. Crucially, this depends on whether Waite encountered one or two skulls. The “typical specimen” cited by Thomas (1904) does not necessarily equate to the tail-less skin M.468 examined by Ramsay. If two skins were present when Waite took the measurements, both could have been treated as “typical” specimens, i.e. “co-types”. It is possible that Ramsay labelled each as “type of the species”, a practice evident from his original labels still attached to mammal syntype skins in the AM. In conclusion, three likely scenarios emerge from the quagmire of uncertainty, depending on whether Waite encountered one or two dry skins. First, a simple situation emerges if the 1890s Catalogue entry had simply neglected to add “skin in spirit” against the female skin. In that case, Waite could only have encountered one dry skin and one skull, those of the tail-less male described by Ramsay (assuming no other undocumented specimens were in the Collection at that time). This would support the preliminary conclusion of Travouillon that the skull currently marked M.469 belongs to skin M.468. Second, Waite encountered two dry skins, which generates many uncertainties. Under this scenario, a female skin and skull have been lost or exchanged from the Collection. Waite might have assumed that both skins were co-types but if only one skull had been extracted, from which skin? If both skulls had been extracted, their correct association with the skins might have been muddled before Waite’s visit in c. 1903, or subsequently. It seems that neither of the skulls would have been marked with names or numbers and Ramsay did not publish skull or dental measurements to enable comparisons. Third, Waite encountered two dry skins, one now lost, and a third specimen, a female in alcohol, was converted to a skin and skull in the 1920s but the locality and collector remain unknown. A simple path through such uncertainty for a taxonomic specialist would be to assume that Ramsay had a syntype series; designate the male skin as lectotype; and designate one of the two skulls in the Collection as the lectotype skull once the taxonomic status of the two suspected forms of I. auratus is resolved., Published as part of Parnaby, Harry & Gill, Anthony C., 2021, Mammal type specimens in the Macleay Collections, University of Sydney, pp. 201-252 in Zootaxa 4975 (2) on pages 211-214, DOI: 10.11646/zootaxa.4975.2.1, http://zenodo.org/record/4806677, {"references":["Ramsay, E. P. (1887) Description of two new species of marsupials (Perameles and Antechinus), and of a new species of Mus (M. Burtoni), from the neighbourhood of Derby, N. W. A. Proceedings of the Linnean Society of New South Wales, Series 2, 2 (3), 551 - 553, pl. xvii. https: // doi. org / 10.5962 / bhl. part. 29199","Jackson, S. M. & Groves, C. P. (2015) Taxonomy of Australian Mammals. CSIRO Publishing, Melbourne, Victoria, 529 pp.","Shaw, G. (1797) The Naturalist's Miscellany: or coloured figures of natural objects, drawn and described immediately from nature. Vol. 8. E. Nodder, London, text to plate 298 pp. https: // doi. org / 10.5962 / bhl. title. 79941","Lyne, A. G. & Mort, P. A. (1981) A comparison of skull morphology in the marsupial bandicoot genus Isoodon: its taxonomic implications and notes on a new species, Isoodon arnhemensis. Australian Mammalogy, 4 (2), 107 - 133.","Mahoney, J. A. & Ride, W. D. L. (1988) Peramelidae. In: Walton, D. W. (Ed.), Zoological Catalogue of Australia. Vol. 5. Mammalia. Australian Government Publishing Service, Canberra, pp. 36 - 42.","Anonymous. (1888) Notes and Exhibits. Proceedings of the Linnean Society of New South Wales, Series 2, 2 (4), 978 - 982.","Macleay, W. (1888) Notes on Mr. Froggatt's collections made in the year 1887, in the vicinity of Derby, King's Sound, N. W. Australia. Proceedings of the Linnean Society of New South Wales, Series 2, 2 (3), 1017 - 1020. https: // doi. org / 10.5962 / bhl. part. 29226","Travouillon, K. J. & Phillips, M. J. (2018) Total evidence analysis of the phylogenetic relationships of bandicoots and bilbies (Marsupialia: Peramelemorphia): reassessment of two species and description of a new species. Zootaxa, 4378 (2), 224 - 256. https: // doi. org / 10.11646 / zootaxa. 4378.2.3","Froggatt, W. W. (1934) A naturalist in Kimberley in 1887. Australian Naturalist, 9 (4), 69 - 82.","Thomas, O. (1904) On a collection of mammals made by Mr. J. T. Tunney in Arnhem Land, Northern Territory of South Australia. Novitates Zoologicae, 11, 222 - 229. https: // doi. org / 10.5962 / bhl. part. 26833"]}

Perameles moresbyensis Ramsay, 1877a Proc. Linn. Soc. N.S.W. (ser. 1) 2 (1), 14. (July 1877). Description read at the 29th January 1877 meeting of the LSNSW. Current name. Papuan Brown Bandicoot Isoodon macrourus moresbyensis (Ramsay, 1877), following Jackson & Groves (2015). Taxonomic status. Often viewed as a subspecies of Isoodon macrourus but further research is necessary to resolve the status of the subspecies of I. macrourus (Jackson & Groves 2015). The validity of this entity has remained in doubt since its publication. Although accepted as a full species in the decades immediately following publication of the description, since the early 20th century taxonomists have remained divided as to whether it is a subspecies or not a valid form. Type locality. No locality given in the original account. Cited in the taxonomic literature as the Port Moresby district, probably because Ramsay used the vernacular “ Port Moresby Bandicoot” in his original account and specific epithet. Material. M.474, (possible syntype) male, skin mount on stand, skull in situ. An old loose paper tag states: “ Perameles moresbyensis, Rams. Port Moresby ♂ ”. A subadult skull with associated label of “temp M.474” has been mistakenly matched with the skin, given that the skull is in situ. That skull is possibly male based on large canines, and is I. macrourus, but the subspecies has not been determined (Dr Kenny Travouillon, Western Australian Museum, pers. comm. 2016), i.e. it is not yet clear if the skull was from Papua New Guinea or Australia. M.475, (likely syntype) adult female, skull with study skin missing tail tip, associated Masters label reads “ Perameles Moresbyensis Ramsay Port Moresby”. Comments. A holotype has usually been assumed in the taxonomic literature and the prospect of a type series has not been discussed. An adult male in the AM (M.2554) is usually cited as Ramsay’s “type” (e.g., Thomas 1888; Lyne & Mort 1981) but is considered a syntype by Parnaby et al. (2017). Ramsay very likely based his description on several specimens, although his account does not reveal the number, sex, or the collections in which they were lodged. Three considerations imply a type series. First, his description of ear colour states “... the ears brown (in one yellow)” [his emphasis]. This implies that he examined several specimens, or less likely, that one ear was brown, the other yellow. Second, his statement “The animal examined is full grown, but young...” contrasts with AM M.2554, which is an old animal (see Parnaby et al. 2017). His statement “the animal examined” is not necessarily inconsistent with a type series if he had selected one specimen for detailed description. Third, the skull of AM M.2554 was not extracted until the 1980s, yet Ramsay appears to have examined a skull. Although he did not describe the skull and his dental measurements could have been made with the skull in situ, his remark that the fourth molars are fully erupted could not have been made without an extracted skull. Possible type material of this taxon in the MAMU has been overlooked, probably because collector and collection date have not been recorded for any of the relevant specimens. The MAMU 1890s Catalogue lists only three specimens under “ Perameles moresbyensis ”: an adult male and female listed as “stuffed” and an adult female “skin”, the latter perhaps a flat skin. Skulls are not listed nor are alcohol specimens. Skulls were usually not listed separately in the Catalogue unless the specimen consisted of a skull only. Currently there are two dry skins in the Collection, along with four specimens in alcohol (M.896–M.899). Although all four alcoholic skins are marked “Port Moresby” in the original entries to the 1960s Mammal Register, locality attribution of associated skulls requires verification due to possible incorrect association of I. macrourus skulls from northern Australian. Reliable diagnostic cranial or dental characters that discriminate I. macrourus from Australia and Papua New Guinea have not yet been defined (Dr Kenny Travouillon, pers. comm. 2016). While it is possible that the 1890s Catalogue had overlooked the four alcohol specimens, the data and skulls might be mis-match to all four, and two of the specimens were registered twice. The one specimen for which Ramsay gave external measurements is a close approximation with skin MAMU M.475. We were able to measure eight of the 14 measurements given by Ramsay, five of which are comparable to his, see Table 3. The main discrepancy is “tip of nose to anterior margin of the eye”, which is c. 1.6 inches (40.6 mm) vs. Ramsay’s measurement of 1.7 inches (43.2 mm), a distance that falls nearly to the centre of the eye in M.475. Distortion or subsequent shrinkage of the skin might account for this discrepancy. Body length was not given by Ramsay but can be derived from other measurements in Table 3, which indicates that linear body length of M.475 (c. 11.5 inches) also approximates the specimen measured by Ramsay (11.5 inches). Consequently there are no major inconsistencies in the measurements that would exclude this skin as possibly being the one examined by Ramsay. Measurements of male skin mount M.474 are larger than Ramsay’s, e.g., nose tip to tail tip measured in inches is c. 18 (45.7 cm) vs. 16.5 (41.9 cm); tail length is c. 6 (15.2 cm) vs. 5 (12.7 cm), and hindfoot length c. 3 (76.2 mm) vs. 2.5 (63.5 mm). As previously indicated, the main obstacle to identifying syntypes in the MAMU is that collector and collection date are not recorded for any of the New Guinea specimens of this taxon. Some of these could have been purchased from Goldie or his associated collectors during 1877–1882, after publication of the account. However, it is likely that several specimens were collected by Petterd and Broadbent and sold to Macleay in 1876, prior to the 29th January 1877 meeting of the LSNSW at which Ramsay read his description. In his paper on mammals of the Chevert Expedition, Ramsay named several species using material from Macleay’s Collection that were almost certainly obtained after the Chevert Expedition, given that the Expedition did not reach Port Moresby (Macleay 1875a; Fulton 2016a). Several members of the Expedition (including Petterd and Broadbent) left the Chevert in August 1875 and joined Andrew Goldie to collect in the region of Port Moresby (see Fulton 2017; 2018). Petterd and Broadbent returned to Australia in 1876 and Macleay noted in his private journal for 7th March 1876 “Pettard and Broadbent visited this morning……Stone purchased all of their birds, I purchased all the mammals they had” (Horning 1994). This would surely be the origin of some of the type specimens of Perameles moresbyensis. The collecting localities of the specimens examined by Ramsay remain unclear. One possibility is that only specimens from the Port Moresby district were used in the description, on the assumption that Ramsay would have listed Hall Sound and Yule Island because he recognised them as separate localities. The absence of basic information in the description suggests that it is equally likely that Ramsay applied the term “ Port Moresby bandicoot” but simply neglected to list the localities of any type specimens. If so, type material could have included Hall Sound. It is not known if bandicoots were collected during the Chevert Expedition. In his ad lib account of the Expedition written before the return of the collections to Sydney on the Chevert, Macleay (1875b) mentioned that a species of Perameles was encountered but whether any were collected is unclear. Given that Perameles was the only genus of New Guinean bandicoots then recognised, his statement could apply to any of the four bandicoot genera that are now recognised from the lowlands of southern Papua New Guinea (Flannery 1995). It appears that the type series included at least one specimen collected by Andrew Goldie that made its way into the AM, probably via Ramsay’s private collection. Parnaby et al. (2017) did not justify their recognition of the AM specimen as a syntype and the AM material is discussed further in the interests of defining the syntype series of this taxon. A mature male skin mount with extracted skull in the AM (M.2554) believed to be a syntype is an old animal with worn teeth and thus not the specimen that was the primary focus of Ramsay’ description, which he states was a young animal (see Parnaby et al. 2017). The collector of M.2554 was not recorded in the original M register entry made in 1915, which states: “ Perameles moresbyensis, Ramsay. Type. from old collection.” An old pencil annotation indicates skin with skull inside. It is possible that AM M.2554 is the one obtained from Andrew Goldie. In a paper read at the 27 December 1876 meeting of the LSNSW, Ramsay (1877b) described a collection of birds and mammals recently obtained from the Port Moresby area by Goldie. In that paper, Ramsay has a line entry “ Perameles sp. ? Perameles Novae-Guineae ”, without further comment and without indicating the number of specimens. This most likely refers to material that he later described as P. moresbyensis at the January 1877 meeting of the LSNSW, his account of Chevert mammals. The only other bandicoot that he named from the New Guinea mainland (Perameles broadbenti Ramsay, 1879) was based on a specimen obtained after that meeting. Ramsay (1877b) stated that he examined Goldie’s entire New Guinea collection. We have not established whether he purchased material of P. moresbyensis for the AM or his private collection. A purchase slip might exist in AM Archives but in case one is not found, the provenance of M.2554 is discussed further. Three new mammals described by Ramsay in 1876 are listed under their published names as “ sp. nov. ” in the AM annual report for the year ending 31st December 1876 (Ramsay 1877c) but P. moresbyensis is not listed in the AM annual reports for 1876 or 1877. This might have been an oversight but the annual reports often did not detail purchased specimens. Regardless of whether Ramsay had purchased material of P. moresbyensis for his private collection, a specimen was in the AM Collection after Ramsay retired. AM M.2554 was registered in March 1915 and listed as being from the “old collection”. Birds purchased from the Ramsay family Dobroyde Collection in 1898 were registered at the AM in 1912 and it is not clear whether the unregistered bandicoot found in the Mammal Collection was incorrectly assumed to be an unregistered specimen from the old collection, but instead arrived with the Dobroyde purchase., Published as part of Parnaby, Harry & Gill, Anthony C., 2021, Mammal type specimens in the Macleay Collections, University of Sydney, pp. 201-252 in Zootaxa 4975 (2) on pages 217-219, DOI: 10.11646/zootaxa.4975.2.1, http://zenodo.org/record/4806677, {"references":["Ramsay, E. P. (1877 a) Zoology of the \" Chevert \". Mammals. Part 1. Proceedings of the Linnean Society of New South Wales, Series 1, 2 (1), 7 - 15.","Jackson, S. M. & Groves, C. P. (2015) Taxonomy of Australian Mammals. CSIRO Publishing, Melbourne, Victoria, 529 pp.","Thomas, O. (1888) Catalogue of the Marsupialia and Monotremata in the Collection of the British Museum (Natural History). British Museum (Natural History), London, xii + 401 pp. https: // doi. org / 10.5962 / bhl. title. 56181","Lyne, A. G. & Mort, P. A. (1981) A comparison of skull morphology in the marsupial bandicoot genus Isoodon: its taxonomic implications and notes on a new species, Isoodon arnhemensis. Australian Mammalogy, 4 (2), 107 - 133.","Parnaby, H., Ingleby, S. & Divljan, A. (2017) Type specimens of non-fossil mammals in the Australian Museum, Sydney. Records of the Australian Museum, 69 (5), 277 - 420. https: // doi. org / 10.3853 / j. 2201 - 4349.69.2017.1653","Macleay, W. (1875 a) The voyage of the Chevert. The Sydney Morning Herald, 11 October, pg. 5.","Fulton, G. R. (2016 a) Thomas Reedy 1842 - 1929: gardener for Sir William Macarthur and plant collector on the Chevert Expedition to New Guinea in 1875. Cunninghamia, 16, 51 - 63.","Fulton, G. R. (2017) Dr William H. James 1852 - 76: medical doctor and naturalist. Bulletin of the British Ornithologists' Club, 137 (1), 71 - 87. https: // doi. org / 10.25226 / bboc. v 137 i 1.2017. a 5","Fulton, G. R. (2018) Notes on the mammals collected on the Chevert Expedition, to New Guinea, in 1875. Proceedings of the Linnean Society of New South Wales, 140, 1 - 6.","Horning, D. (1994) Transcription of the William John Macleay journal, from original located at Mitchell Library MSS 2009 Linnean Society of New South Wales records, 1826 - 1970. [unknown pagination, unpublished manuscript at Macleay Museum, University of Sydney]","Macleay, W. (1875 b) Notes on the zoological collections made in Torres Straits and New Guinea during the cruise of the \" Chevert \". Proceedings of the Linnean Society of New South Wales, Series 1, 1, 36 - 40. https: // doi. org / 10.5962 / bhl. part. 12383","Flannery, T. F. (1995) Mammals of New Guinea. Revised Edition. Australian Museum / Reed Books, Chatswood, New South Wales, 440 pp.","Ramsay, E. P. (1877 b) Notes on a collection of birds from Port Moresby; with descriptions of some new species. Proceedings of the Linnean Society of New South Wales, Series 1, 1 (4), 386 - 395. https: // doi. org / 10.5962 / bhl. part. 12426","Ramsay, E. P. (1879) Contributions to the zoology of New Guinea. Part III. Description of a new marsupial allied to the genus Perameles, Geoff. Proceedings of the Linnean Society of New South Wales, Series 1, 3 (4), 402 - 405, pl. 27. https: // doi. org / 10.5962 / bhl. part. 22249","Ramsay, E. P. (1877 c) Appendix 7, Taxidermist's Department. In: Scott, A. W. (Ed.), Report of the Trustees of the Australian Museum for the year ending 31 December, 1876. Charles Potter, Acting Government Printer, Sydney, New South Wales, pp. 12 - 13."]}

Perameles macroura var. torosus Ramsay, 1877a Proc. Linn. Soc. N.S.W. (ser. 1) 2 (1), 12. (July 1877). Description read at the 29th January 1877 meeting of the LSNSW. Current name. Northern Brown Bandicoot Isoodon macrourus torosus (Ramsay, 1877), following Jackson & Groves (2015). Taxonomic status. Considered to be a valid species by some taxonomists of the late 19th and early 20th century while others doubted its validity, even as a subspecies. Currently, usually recognised as a subspecies of I. macrourus (Gould, 1842a), but further research is needed and the biological reality of I. macrourus torosus has been questioned by some recent authors (see Jackson & Groves 2015). Ramsay published the name Perameles macroura var. torosus, amended to Perameles macroura torosa by Mahoney & Ride (1988). Syntype. M.471, adult male, skin mount from Endeavour River (Fig. 5). Registered sometime between 1963 to 1965 and listed in the MAMU Mammal Register as a male skin mount, with associated number M1 (Fig. 6), but no mention of a skull. A skull and mandibles subsequently matched to skin M.471 were registered as M.1153 (Fig. 7), probably during 1965. Skull M.1153 is probably correctly matched with skin M.471, but this requires confirmation. Although “M1” is written in pencil on the skull, it is not known if this was done at the AM during original skull extraction, or subsequently. It is clear from entries and annotations during the 1960s registration of the bandicoot collection that the association of skulls with skins was problematic. Skin mount M.471 is currently the only candidate for lectotype designation but that decision should be made by a specialist in Isoodon taxonomy. Type locality. Usually assumed to be near Cooktown, Queensland (e.g., Troughton 1941). In his original description, Ramsay mentioned a specimen from Endeavour River (= Cooktown) but gave the distribution as Rockingham Bay to the Endeavour River, implying syntypes. Comments. The type series has not been identified in the literature until now. Ramsay described an adult female skin. He does not describe a skull and there is no indication that he examined an extracted skull. The few dental measurements given could have been taken from a skull in situ. Although Ramsay’s description focused on a female from Endeavour River, it is likely that it was based on several specimens, although his description is ambiguous. He stated that specimens that he regarded to be P. m. torosus were from Rockingham Bay north to the Endeavour River. Consequently, syntypes are assumed here although previous literature references imply that Ramsay based the description on one specimen. For example, Mahoney & Ride (1988) concluded that the type series of I. m. torosus consisted of one specimen (the holotype), whereabouts unknown. Although this taxon was described in his paper reporting mammals obtained on the Chevert Expedition, the specimens were probably not obtained during the expedition. The Chevert party did not collect on the mainland in the region of Endeavour River en route to Papua New Guinea (Macleay 1875a; see Chevert itinerary of Fulton (2016a)). On the return trip, during which Brazier appeared to be the only zoologist remaining on board, the Chevert crew remained offshore at Endeavour River for a few hours while detained by Customs for cargo inspection (MacMillan 1957: 153). The syntypes originally in Macleay’s Collection probably included several specimens collected by Edward Spalding in July 1874 from the Endeavour River region. An entry by Macleay in his private diary dated 31st July 1874 states that he received a shipment of specimens from Endeavour River sent by Spalding, that included “skins of a new Bandicoot” (Horning 1994). The only specimen from Endeavour River listed in the MAMU 1890s Catalogue does not use the name P. m. torosus. Instead, the Catalogue lists Queensland specimens from three localities under “ Perameles macrura [sic] Gould the North-Australian Bandicoot” (Fig. 2). The author of the 1890s Catalogue, who we assume to be Masters, evidently followed the mammal nomenclature of Thomas (1888) and Ogilby (1892), neither of whom recognised torosus as a valid race of P. macroura. The Catalogue lists only two dry skins from Queensland: an adult male stuffed skin from Endeavour River and a female skin from “ Queensland ”. Perhaps the female skin from Queensland is the one from Endeavour River described by Ramsay but we have not compared its measurements against those of the description. The Catalogue also lists a series of adult and subadult P. macroura in alcohol from a third Queensland locality of Herbert River but the collectors are not specified. Macleay could have obtained material from a range of collectors in addition to Spalding but after publication of P. m. torosus. One possibility is John Archibald Boyd, who sent Macleay several shipments of mammals in alcohol during the 1880s from his home at Ripple Creek, Ingham, on the Herbert River (e.g., Macleay 1885). Although Cardwell is not mentioned in the original description, reference to Rockingham Bay probably included Cardwell, a well-known collecting locality from which Ramsay and Macleay obtained mammal specimens from collectors in the first half of the 1870s. For example, Kendall Broadbent collected around Cardwell in 1873 (Whittell 1954) and sent mammals to Ramsay. Ramsay also made several collecting trips to the Rockingham Bay district before publication of his description of P. m. torosus. It is therefore possible that additional syntypes might originally have been in Ramsay’s private collection (the Dobroyde Museum) or the AM but we have not located any supporting documentation. Male skin mount M.471 is a likely syntype, and it appears that Troughton had also reached that conclusion. Troughton, then Curator of Mammals at the AM, examined the taxonomy of I. macrourus during 1926 and 1927. Having extracted and examined the skulls of two specimens in the MAMU, which we believe were one I. macrourus from Endeavour River, and another from Port Darwin, he expressed the opinion (Anonymous 1927) that the Endeavour River specimen was Ramsay’s type and that torosus was probably not a valid race, a view that he maintained over ensuring decades (e.g., Troughton 1941). We are confident that skin mount M.471 is the specimen considered to be the “ type ” by Troughton. Although unregistered at that time, we believe that M.471 (associated tag M1) was one of two “ Perameles macrourus ” skins sent to the AM in 1926 for skull extraction and identification, the other likely to be male skin mount M.470 (associated tag M2) from Port Darwin. The MAMU Donations Book entry for 27th September 1926 lists “ one P. auratus and 2 P. macrourus” being sent to the AM. The Donations Book entry for 12th November 1929 indicates that these specimens were returned and “restuffed after removal of skulls for sp. identification”. The metal tags and hand-written watch tags marked M1 and M2 now associated with these skins are characteristic of similar hand-writing on tags attached to AM specimens from that period. Tags M1 and M2 were presumably attached by Troughton to distinguish each I. macrourus skin when skulls were extracted at the AM. A third skin (P. auratus, presumably M.469) sent for skull extraction at the same time does not have an associated M tag. There appear to be no previously published images or skull measurements of type material of P. m. torosus. Measurements (mm) of M.1153. Cranium: GL, 85.2; CONL, -; GL from nasals, -; ZYG, 35.9; POC, 11.4; Anterior palatal vacuity length, 7.2; Nasal Length, 34.1; Maximum breadth across both nasals, 5.4; Rostral width at infraorbital foramina, 15.7; Skull breadth at lachrymals, 20.7; Brain case height, 21.1; Basisphenoid bone length, 13.2; Alisphenoid bulla width, 8.8; Maximum breadth across bullae, 28.1; I 1–4 (at base of enamel), 6.35; I 1–5 (at base of enamel), 9.1; I 1 –M 4 (cingl.), 46.6; M 1 –M 3 (crowns), 14.0; M 1 –M 4 (crowns), 17.15; C 1 –C 1 breadth (alveoli), 11.8; M 3 –M 3 breadth (crowns), 24.6. Mandible: Dentary length (from condyl.), 65.2; P 1 –P 3 (cingl.), 11.5; M 1 –M 4 (crowns), 18.3., Published as part of Parnaby, Harry & Gill, Anthony C., 2021, Mammal type specimens in the Macleay Collections, University of Sydney, pp. 201-252 in Zootaxa 4975 (2) on pages 215-217, DOI: 10.11646/zootaxa.4975.2.1, http://zenodo.org/record/4806677, {"references":["Ramsay, E. P. (1877 a) Zoology of the \" Chevert \". Mammals. Part 1. Proceedings of the Linnean Society of New South Wales, Series 1, 2 (1), 7 - 15.","Jackson, S. M. & Groves, C. P. (2015) Taxonomy of Australian Mammals. CSIRO Publishing, Melbourne, Victoria, 529 pp.","Gould, J. (1842 a) Characters of a new species of Perameles, and a new species of Dasyurus. Proceedings of the Zoological Society of London, 10 (1842), 41 - 42.","Mahoney, J. A. & Ride, W. D. L. (1988) Peramelidae. In: Walton, D. W. (Ed.), Zoological Catalogue of Australia. Vol. 5. Mammalia. Australian Government Publishing Service, Canberra, pp. 36 - 42.","Troughton, E. L. G. (1941) Furred Animals of Australia. First edition. Angus & Robertson, Sydney, xxvii + 374 pp.","Macleay, W. (1875 a) The voyage of the Chevert. The Sydney Morning Herald, 11 October, pg. 5.","Fulton, G. R. (2016 a) Thomas Reedy 1842 - 1929: gardener for Sir William Macarthur and plant collector on the Chevert Expedition to New Guinea in 1875. Cunninghamia, 16, 51 - 63.","MacMillan, D. S. (1957) A Squatter went to Sea: the Story of Sir William Macleay's New Guinea Expedition (1875) and his Life in Sydney. Currawong Publishing Co., Sydney, x + 165 pp.","Horning, D. (1994) Transcription of the William John Macleay journal, from original located at Mitchell Library MSS 2009 Linnean Society of New South Wales records, 1826 - 1970. [unknown pagination, unpublished manuscript at Macleay Museum, University of Sydney]","Thomas, O. (1888) Catalogue of the Marsupialia and Monotremata in the Collection of the British Museum (Natural History). British Museum (Natural History), London, xii + 401 pp. https: // doi. org / 10.5962 / bhl. title. 56181","Ogilby, J. D. (1892) Catalogue of Australian Mammals, with Introductory Notes on General Mammalogy. Trustees, Australian Museum, Sydney, xvi + 144 pp. https: // doi. org / 10.5962 / bhl. title. 1130","Macleay, W. (1885) On some Reptilia lately received from the Herbert River district, Queensland. Proceedings of the Linnean Society of New South Wales, Series 1, 10 (1), 64 - 68. https: // doi. org / 10.5962 / bhl. part. 17898","Whittell, H. M. (1954) The Literature of Australian Birds: a History and a Bibliography of Australian Ornithology. Paterson Brokensha, Perth, Western Australia, xi + 140 + 788 pp. [limited edition reprinted by Maurizio Martino, Mansfield, Connecticut, USA]","Anonymous. (1927) Notes and Exhibits meeting of 27 th October, 1926. Proceedings of the Linnean Society of New South Wales, 51, xli - xlii. [Troughton exhibits a bandicoot]"]}

Parnaby, Harry, Gill, Anthony C. (2021): Mammal type specimens in the Macleay Collections, University of Sydney. Zootaxa 4975 (2): 201-252, DOI: https://doi.org/10.11646/zootaxa.4975.2.1

Many mammals dig, either during foraging to access subsurface food resources, or in creating burrows for shelter. Digging requires large forces produced by muscles and transmitted to the soil via the skeletal system; thus fossorial mammals tend to have characteristic modifications of the musculoskeletal system that reflect their digging ability. Bandicoots (Marsupialia: Peramelidae) scratch-dig mainly to source food, searching for subterranean food items including invertebrates, seeds, and fungi. They have musculoskeletal features for digging, including shortened, robust forelimb bones, large muscles, and enlarged muscle attachment areas. Here, we compared changes in the ontogenetic development of muscles associated with digging in the Quenda (Isoodon fusciventer). We measured muscle mass (m m ), pennation angle, and fiber length (FL) to calculate physiological cross-sectional area (PCSA; a proxy of maximum isometric force) as well as estimate the maximum isometric force (Fmax) for 34 individuals ranging in body size from 124 to 2,390 g. Males grow larger than females in this bandicoot species, however, we found negligible sex differences in mass-specific m m , PCSA or FL for our sample. Majority of the forelimb muscles PCSA showed a positive allometric relationship with total body mass, while m m and FL in the majority of forelimb muscles showed isometry. Mechanical similarity was tested, and two thirds of forelimb muscles maximum isometric forces (Fmax) scaled with isometry; therefore the forelimb is primarily mechanical similar throughout ontogeny. PCSA showed a significant difference between scaling slopes between main movers in the power stroke, and main movers of the recovery stroke of scratch-digging. This suggests that some forelimb muscles grow with positive allometry, specially these associated with the power stroke of digging. Intraspecific variation in PCSA is rarely considered in the literature, and thus this is an important study quantifying changes in muscle architectural properties with growth in a mammalian model of scratch-digging.

Ixodes acer n. sp., Ixodes giluwensis n. sp., Ixodes mirzai n. sp., Ixodes planiscutatus n. sp. and Ixodes stellae n. sp. (Acari: Ixodidae), are described based on females ex various marsupials (Dasyuromorphia: Dasyuridae, Diprotodontia: Petauridae, Peramelemorphia: Peramelidae), rodents (Rodentia: Muridae) and a bird (Galliformes: Megapodiidae) from Indonesia (Papua Province) and Papua New Guinea. Females of all new species are similar to those of I. luxuriosus Schulze, 1935, I. steini Schulze, 1935 and I. zaglossi Kohls, 1960, but can be distinguished from them and each other by the distinctness of lateral carinae and cervical grooves on the scutum, length of setae on the idiosoma, sculpture of the basis capituli and palpal segment I dorsally, size of the dorsal cornua, shape and sculpture of the palpal segment I ventrally, size of ventral spur on the palpal segment I, dental formula on the hypostome and measurements of various structures and their proportions. For comparative purposes the female of I. luxuriosus, I. steini and I. zaglossi is redescribed and lectotypes for I. luxuriosus and I. steini have been designated. Studied females of I. luxuriosus, I. steini and I. zaglossi were found on marsupials (Dasyuromorphia: Dasyuridae; Diprotodontia: Macropodidae, Petauridae, Phalangeridae, Pseudocheiridae; Peramelemorphia: Peramelidae), rodents (Rodentia: Muridae) and echidnas (Monotremata: Tachyglossidae) in Indonesia (Papua Province) and Papua New Guinea.

Novel ecosystems are increasingly common worldwide, particularly in areas heavily impacted by humans such as urban and peri-urban landscapes. Consequently, interest in their potential contribution to biodiversity conservation is growing, including their ability to sustain populations of threatened species. However, few studies have explored whether novel habitats can support viable populations over time and how they compare to less modified, remnant habitats. We investigated the capacity for novel habitats to support an endangered mammal, the southern brown bandicoot (Isoodon obesulus obesulus: Peramelidae), in a highly-modified landscape near Australia's second largest city, Melbourne. We compared bandicoot abundance and body condition between five novel and two remnant sites, and examined whether novel sites support residency and key demographic processes necessary for bandicoot population persistence. We found that bandicoot abundance was higher at novel than remnant sites, with the highest abundance at the novel site with the most urbanised surroundings. Female body condition was similar between novel and remnant sites. The majority of bandicoots at novel sites were resident, and breeding activity, recruitment of first-year adults, and survival of mature adults were observed at all novel sites. Our results demonstrate the potential significance of novel habitats for conserving threatened species within heavily-modified landscapes, and encourage us not to judge the quality of habitats on their novelty alone. Broadening our appreciation of the potential value of novel ecosystems could increase off-reserve species conservation opportunities, a key priority within the context of the Anthropocene and unprecedented global change and biodiversity loss.

Animals that forage for food or dig burrows by biopedturbation can alter the biotic and abiotic characteristics of their habitat. The digging activities of such ecosystem engineers, although small at a local scale, may be important for broader scale landscape processes by influencing soil and litter properties, trapping organic matter and seeds, and subsequently altering seedling recruitment. We examined environmental characteristics (soil moisture content, hydrophobicity and litter composition) of foraging pits created by the southern brown bandicoot (Isoodon obesulus; Peramelidae), a digging Australian marsupial, over a 6-month period. Fresh diggings typically contained a higher moisture content and lower hydrophobicity than undisturbed soil. A month later, foraging pits contained greater amounts of fine litter and lower amounts of coarse litter than adjacent undug surfaces, indicating that foraging pits may provide a conducive microhabitat for litter decomposition, potentially reducing litter loads and enhancing nutrient decomposition. We tested whether diggings might affect seedling recruitment (seed removal by seed harvesters and seed germination rates) by artificially mimicking diggings. Although there were no differences in the removal of seeds, seedling recruitment for three native plant species (Acacia saligna, Kennedia prostrata and Eucalyptus gomphocephala) was higher in plots containing artificial diggings compared with undug sites. The digging actions of bandicoots influenced soil moisture and hydrophobicity, the size distribution of litter and seedling recruitment at a local scale. The majority of Australian digging mammals are threatened, with many suffering substantial population and range contraction. However, their persistence in landscapes plays an important role in maintaining the health and function of ecosystems.

Perameles papillon sp. nov. Figs. 5–7 Holotype. WAM M571, skin and skull, female. Type locality. Ooldea, South Australia (SA). Paratypes. WAM M126, skull, sex unknown, Eucla Pass, Western Australia (WA); WAM M570, skin and skull, female, Ooldea (SA); WAM M572, skin and skull (juvenile), male, Ooldea (SA); WAM M574, skin and skull (Juvenile), male, Ooldea (SA); WAM M576, skin and skull, female, Ooldea (SA); WAM M577, skin and skull, female, Ooldea (SA); AM M4352, Skin and skull, female, Rawlinna (WA); AM M31547, alcohol specimen, male, Ooldea (SA); BMNH 1925.10.8.26, skin, male, Ooldea (SA); BMNH 1925.10.8.27, skin, female, Ooldea (SA); SAM M3973, alcohol specimen and skull, female, collected between Ooldea and Tallaringa (SA); SAM M4640, skin and skull, male, Ooldea (SA). Referred specimens. AMNH 196371, 2 left subfossil dentaries, Horseshoe Cave, Nullarbor (WA); AMNH 220153 and AMNH 220154, juvenile subfossil skulls, south west Loongana, Nullarbor (WA); AM S1821, skull, juvenile, Rawlinna (WA); AM M2986, skin and skull, female, Fisher, Nullarbor (SA); AM M4353, skin and skull, juvenile female, Rawlinna (WA); AM M4850, skin and skull, male, Ooldea (SA); AM M4851 and M4852, skin and skull, female, 9 miles west of Ooldea (SA); AM M3049, alcohol specimen, juvenile female, Ooldea (SA); AM M4368–M4369, alcohol specimen, juvenile female, Rawlinna (WA); AM M4370–M4371, alcohol specimen, juvenile male, Rawlinna (WA); AM M4939 and M4941, alcohol specimen, pouch young male, Ooldea (SA); AM M4940, alcohol specimen, pouch young female, Ooldea (SA); AM M4942–4943, alcohol specimen, male, East West Line (SA); AM M4944–4945, alcohol specimen, female, East West Line (SA); AM M4978–4979, alcohol specimen, pouch young female of M4852, 9 miles west of Ooldea (SA); AM M31545, alcohol specimen, juvenile male, Ooldea (SA); AM M31548, alcohol specimen, male, Ooldea? (SA); MAGNT U7605, skull, specimen bred in Adelaide in 1926 (original locality unknown); NMV C7173 and C7174, subfossil skulls, N11 Cave, Nullarbor East (SA); NMV C7175 and C7176, subfossil skulls, Weebubbie Cave, Nullarbor (WA); NMV C7177 and C7178, subfossil skulls, Weekes Cave, Nullarbor (SA); NMV C7180, 2 subfossil dentaries, Koonalda Cave, Nullarbor (SA); QM J5028, alcohol specimen, female, Nullarbor (SA); SAM M846, alcohol specimen and skull, juvenile female, Ooldea (SA); SAM M1394, mount, male, Ooldea (SA); SAM M1395, mount, Ooldea (SA); SAM M1396, mount, female, Ooldea (SA); SAM M1397, skin, male Ooldea (SA); SAM M1398, skin, female Ooldea (SA); SAM M3926–3927, M3976, M3981–3982, M3984, M3987–3988, alcohol specimen and skull, female, no location data; SAM M3977, M3983, M3985–3986, alcohol specimen, juvenile, no location data; SAM M3980, M3989, alcohol specimen and skull, male, no location data; SAM M4639, skin and skull, juvenile female, Ooldea (SA); SAM M5226, mount, no data; SAM M6302, many subfossil skulls, Weebubbie Cave, Nullarbor (WA); SAM M6303, many subfossil skulls, Weekes Cave, Nullarbor (SA); SAM M6304, many subfossil skulls, N11 Cave, Nullarbor East (SA); SAM M6305, many subfossil skulls, Koonalda Cave, Nullarbor (SA); WAM 76.4.21, skull, Abrakurrie Cave, Nullarbor (WA); WAM 67.4.114–116, 67.10.305–307, skulls, Homestead Cave, Nullarbor (WA); WAM 66.6.45–53, 66.1.54, 68.3.63–67, 69.5.17, skulls, Murra-el-elevyn Cave, Nullarbor (WA); WAM 69.7.557– 559, skulls, Kjeldahl Cave, Cocklebiddy Roadhouse, Nullarbor (WA); WAM 73.1.107, 74.5.39, skulls, Webbs Cave, Nullarbor (WA); WAM 67.5.128, 67.5.130, skulls, Warbla Cave, Nullarbor (SA); WAM 67.10.420–421, 68.3.118, skulls, Blowhole, Eucla, Nullarbor (WA); WAM 67.4.250, 72.1.8, 72.1.1088–1089, skulls, Horseshoe Cave, Nullarbor (WA); WAM 66.6.45–53, skulls, Graham’s Cave, Nullarbor (WA); WAM 68.3.51, skull, cave 3 3/ 4 mile South of Cocklebiddy Cave, Nullarbor (WA); WAM 68.3.33, skull, Mullamullang Cave, Nullarbor (WA); WAM 67.11.32, skull, Capstan Cave, Nullarbor (WA); WAM 67.4.343, skull, cave 23 miles west of Eucla, Nullarbor (WA); WAM 68.3.20, skull, cave 6 miles south of Madura, Nullarbor (WA); WAM 67.9.134, cave near Madura, in doline, Dingo Douga, Nullarbor (WA); WAM 66.3.14–16, 67.10.177–182, skulls, Firestick Cave, Nullarbor (WA); Nullarbor (WA). Diagnosis. Perameles papillon sp. nov. differs from other species of Perameles in having a dark horizontal bar on its ears, a bracelet of light brown fur around its wrists and covering the heel of the foot and the posterior edges of the footpad, a dark butterfly pattern on its rump, shorter pes and tail (see results), enlarged lacrimal crest (except for P. gunnii), enlarged bullae (largest of any Perameles), accessory fenestrae extending anteriorly to the canine, maxillopalatine fenestrae extending posteriorly to the M3, well-developed squamosal epitympanic sinus. Perameles papillon sp. nov. differs from all members of Peramelemorphia (except P. myosuros) in having females distinctively larger than males. Etymology. Papillon, French for butterfly, in reference to the butterfly shaped pattern on its rump. Remarks. Westerman et al. (2012) subsampled specimen AM M31545 from the Australian Museum, which was attributed to Perameles b. notina, from Ooldea in South Australia. After scoring specimens from this region, including the rest of the Nullarbor Plain in Western Australia, and comparing them to the holotype of Perameles b. notina (then named Perameles myosura notina) BMNH 43.8.12.21 from the head of St Vincent Gulf in South Australia, it became evident that they did not match in morphology, the type being much larger and several features on the skin and skull did not match any of the specimens recovered from the Nullarbor region. As a result, AM M31545 and all specimens found in the Nullarbor regions (present in collections in various museums) are considered here a new species. The results of our analysis also show that Perameles papillon sp. nov. is distinct morphologically from Perameles bougainville and P. eremiana, but also distinct to the currently unrecognised P. myosuros, P. notina and P. fasciata. Our results further suggest that the latter three taxa are distinct taxa and should no longer be subsumed in P. bougainville (see discussion below). Distribution. Collected only in the Nullarbor Plain, on either side of the Western Australian and South Australian border (triangles in Fig. 8). Description. The description of the skin is a comparison of Perameles papillon sp. nov., represented by Holotype WAM M571 (Fig. 5), to P. b. bougainville, represented by WAM M3641 (Suppl. Fig. 2), and P. b. notina, represented by the holotype BMNH 43.8.12.21, unless specified otherwise (Suppl. Fig. 3). P. b. fasciata and P. b. myosuros are not used in the comparison here as they were not sampled by Westerman et al. (2012), though we do suspect they are distinct taxa and will be investigated in future research. P. papillon sp. nov. is similar in size to P. b. bougainville but much smaller than P. b. notina, showing no marked sexual dimorphism between males and females, though females seem to be a little larger (see Table S2 –3 for measurements, Fig. 3 for morphometric analysis). The fur on the dorsal side of the head is a little darker and browner than P. b. bougainville, but lighter than P. b. notina. The ventral side is cream in colour as in P. b. bougainville whereas it is greyer in P. b. notina. The ears have a noticeable dark horizontal bar across them, absent in both P. b. bougainville and P. b. notina or any other known peramelemorphian. Note that none of the wet specimens of P. papillon sp. nov. show this feature (dark bar on ear) but it is present in all dry skins, and this is suspected to be as a result of accelerated fading in alcohol specimens. The ears are long as is P. b. bougainville, but shorter than P. b. notina. There are long dark vibrissae present at the front of the snout, above the eye and on the cheek below the eye. These are also present in P. b. bougainville and P. b. notina, but less obvious in P. b. bougainville. The body is also darker and browner on the dorsal side than in P. b. bougainville but lighter than P. b. notina. Both P. papillon sp. nov. and P. b. bougainville are cream coloured on the ventral side, while P. b. notina is mostly grey except for patches of cream. A complex pattern of different coloured bars is present on the rump (juveniles WAM M574 and M572 show the early stages of the development of this pattern on the rump), which is absent in P. b. bougainville (P. b. bougainville has a single very faint darkish bar across its rump that tapers laterally) but present in P. b. notina, though the holotype is faded, making this pattern less obvious. The pattern resembles that of P. gunnii and P. b. notina but it is less well defined. There isn’t a central dark dorsal bar as in P. gunnii, and most likely in P. b. notina, and the light bars are bicoloured, cream and golden brown (golden brown only in P. gunnii and P. b. notina). The anterior most two dark bars are almost completely black or at least dark brown (specimens are likely faded), and looks like butterfly wings in shape in dorsal view. The posterior darkest bars are much lighter in colour and less well defined, distinguishing P. papillon sp. nov. from P. gunnii and P. b. notina, which have a well-defined posterior bar. The tail is short as in P. b. bougainville (longer in P. b. notina), but it is tricoloured (dark brown dorsally, beige laterally and cream ventrally) instead of bicoloured as in P. b. bougainville (the tail is dark brown dorsally and cream ventrally) or unicoloured as in P. b. notina (the tail is dark grey). The forelimbs are light brown in colour on the dorsal side up to the wrist, where it surrounds the wrist like a bracelet. The manus and the ventral side of the forelimbs are cream in colour. This pattern is different to P. b. bougainville, which lacks the bracelet of brown fur around the wrist, and is lighter in colour dorsally. The forelimbs of P. b. notina are completely grey. The manus has the three large toes (digit II, III and IV), and the typically reduced digits I and V as in all Peramelidae. Compared to P. b. bougainville and P. b. notina, the digits are shorter but have more pronounced toepads as in P. b. notina (the toes of P. b. bougainville more slender). The hindlimbs, in the region of the tibia, are cream in colour anteriorly and greyish brown on the posterior side. The darker colour invades the heel of the cream coloured pes, and the posterior edges of the footpad, unlike in P. b. bougainville where the pes is completely cream in colour and P. b. notina where the pes is completely grey in colour. The footpad is hairless posterior to digit I to the heel as in P. b. notina, while in P. b. bougainville, this area is covered in fur. As in the manus, the pes has shorter toes with thicker toepads than P. b. bougainville and P. b. notina. The description of the skull and teeth is based on paratype WAM M577 for Perameles papillon sp. nov. (Fig. 6), and describes only features that differ from other Perameles, using WAM M6166 for P. b. bougainville, BMNH 43.8.12.21 for P. b. notina, WAM M2629 for P. eremiana, WAM M16590 for P. gunnii (Suppl. Fig. 4), QM JM18575 for P. pallescens and WAM M44421 for P. nasuta (see Travouillon 2016a), unless specified otherwise. The skull of P. papillon sp. nov. is similar in size to P. b. bougainville and P. eremiana (much smaller than P. b. notina, P. gunnii, P. pallescens and P. nasuta) but it is shorter than P. eremiana (nasals shorter in length), and much wider than both P. b. bougainville and P. eremiana. In dorsal view, the lacrimal crest is highly developed, as seen in P. gunnii and P. b. notina, and is visible as a distinct posteriorly orientated process in the orbit. The lacrimal crest is less developed in all other species of Perameles. No sagittal crest is present, as in all Perameles except in males of P. nasuta and P. pallescens. In ventral and lateral view, the alisphenoid tympanic process, or bullae, is highly distinctive in being the most developed (the second largest bullae) within the genus. P. b. bougainville has the smallest bullae, followed by P. nasuta, P. gunnii, P. eremiana, P. papillon sp. nov. and P. b. notina, in increasing order of size. On the palate, accessory fenestrae are present between the incisive fenestrae and the maxillopalatine fenestrae as in all Perameles except P. nasuta and P. pallescens, but they extend anteriorly to the canine, unlike other species of Perameles except P. b. notina where the anterior extension of the accessory fenestrae is posterior to the canines. The maxillopalatine fenestrae also extend further posteriorly than in other Perameles except P. b. notina, from the anterior of the M3 to level with the middle of M3, instead of level with M2. The dividing septa of the maxillopalatine fenestrae are thick anteriorly and posteriorly, thinning centrally, as in Perameles nasuta (very thin in all other Perameles). Palatine fenestrae are present and large as in P. b. bougainville (broken in P. b. notina). In lateral view the orbitosphenoid is identified as a very reduced bone, as in P. nasuta, P. pallescens and P. gunnii (much larger in P. b. bougainville and P. eremiana, broken in P. b. notina). The antorbital fossa is very deep as in all Perameles except in P. nasuta and P. pallescens. The squamosal epitympanic sinus of P. papillon sp. nov. is the most developed of any Perameles (most visible in paratype WAM M126), being wide with a high posterior wall. The I5 is pointed and canine-like as in P. b. bougainville, P. b. notina, P. nasuta and P. pallescens. The canines are small with accessory cusps, in both males and females, as in P. b. bougainville, P. b. notina and P. eremiana (males of P. gunnii, P. nasuta and P. pallescens have an enlarged canine, and males of P. nasuta and P. pallescens generally lack accessory cusps). The canines are entirely bordered by the maxilla as in all Perameles, except P. eremiana (bordered by premaxilla anteriorly). The diastema between the canine and P1 is long (longer than P1) as in all Perameles except P. nasuta and P. pallescens. The P1 is as long as the P2 as in P. gunnii, P. nasuta and P. pallescens (shorter in P. b. bougainville, longer in P. eremiana and P. b. notina), and the P2 is noticeably shorter than the P3, as in P. b. notina, P. nasuta and P. pallescens (subequal in length in all other Perameles). The development of the lingual shelf of P3 is small as in P. b. bougainville and P. eremiana. The major cusp of P3 is similar in shape to all Perameles except P. gunnii (more conical). The upper molars of P. b. notina are completely worn down and the holotype lacks a dentary and therefore these are not used in the comparison following. On the M1, a short stylar crest is present anterior to the metastyle (best seen on juvenile WAM M572) with an associated StE, while StE is a conical cusp with no associated crest in P. b. bougainville (best seen in juvenile WAM M16086). StA is variable in size on M1–2. StB and StC are fused and indistinguishable as in P. b. bougainville and P. eremiana. The preparacrista reconnects with the postparacrista posteriorly on the M1, as in P. eremiana. The postprotocrista on M1 terminates posterior to the metacone as in P. b. bougainville and P. gunnii with no distinct posterior cingulum present (terminates more buccally in all other Perameles, as a posterior cingulum). On the M2, StB is conical as in all Perameles except P. gunnii (oval in shape). No stylar crest is present posterior to the conical StD as seen in P. gunnii and P. b. bougainville (small crest present in all other taxa). The postprotocrista ends posterior to the metacone as in all Perameles, except P. nasuta. StE is present as a distinct large cusp as in P. gunnii (it is reduced in all other species). On the M3, the postprotocrista ends posterior to the metacone as in all Perameles, except P. nasuta. The metaconule is small with only a small shelf separating it from the lingual flank of the metacone, as in P. b. bougainville and P. pallescens (a larger metaconule and shelf is present in all other taxa). StE is absent as in all Perameles except P. gunnii. No anterior cingulum present as in all Perameles except for some specimens of P. pallescens and P. nasuta. On the M4, StB is variable, sometimes absent, sometimes present as a small cusp. This feature is variable across Perameles. The postparacrista is curved, forming a small centrocrista, as in all Perameles, except P. gunnii and P. b. bougainville, where it is straight. The postprotocrista ends anterior to the posterior end of the postparacrista as in all Perameles except P. nasuta. The metacone is small as in all Perameles except P. gunnii and P. nasuta, which have an enlarged metacone. StC/D is absent (sometimes present in P. gunnii and P. nasuta). The lower canine is small and has accessory cuspids as in all Perameles except in males of P. nasuta and P. pallescens, which have enlarged unicuspid canines. The diastema between the canine and p1 is shorter than the length of p1, as in P. pallescens (longer that p1 length in all other taxa). The p1 is as long as the p2 as in P. b. bougainville and P. eremiana (it is shorter in all other taxa). The p2 is longer than the p3 as in P. b. bougainville and P. eremiana (subequal in all other taxa). No cuspid is present on the hypoflexid of the lower molars as in all Perameles except P. nasuta and P. eremiana. On the m1, the paraconid-metaconid distance is equal to the metaconid-protoconid distance as in all Perameles except P. pallescens, which has a longer paraconid-metaconid distance. The posthypocristid is oblique to the tooth row and connects to the hypoconulid as in all Perameles (sometimes perpendicular in P. b. bougainville and connects to entoconid). On the m2, the posthypocristid is also oblique and connects to a large hypoconulid (this feature varies across Perameles). On the m3, the hypoconid is buccal to the protoconid as in all Perameles except P. b. bougainville and some specimens of P. pallescens, which have the two cuspid level with one another.

Peroryctes aruensis Kear et al. 2016 new comb. Lemdubuoryctes aruensis Kear et al. 2016 Holotype. WAM 14.9.6, left maxilla with P3, M1–2. Referred material. WAM 14.9.1–14.9.5 and WAM 14.9.7– 14.9.20, WAM 14.9.51 – 14.9.53. Type locality. Liang Lemdubu cave, Pulau Kobroor, Aru Islands group, Eastern Indonesia. Remarks. Kear et al. (2016) described this taxon, and assigned it to a new genus based on the result of their phylogenetic analysis. This taxon is only represented by heavily worn specimens, and so Kear et al. (2016) scored worn teeth in a matrix that was designed to be scored only on unworn teeth. Their principal character for placing this taxon in a new genus is the presence of a centrocrista in M1–3. This is only seen in unworn Oligo-Miocene bandicoot teeth, but almost every single peroryctid (Peroryctes, Echymipera, Rhnynchomeles and Microperoryctes) do wear their teeth to the point that a centrocrista is formed on M1–3 as a result of wear. The holotype of Peroryctes broadbenti illustrates this morphology, being an adult with worn teeth, and centrocristae are present on M1–3. This does not happen in Peramelidae, Chaeropodidae or Thylacomyidae. As a result, the assessment of this taxon is dubious, and was rescored here, and re-analysed. All characters that can be scored for L. aruensis were identical to that of Peroryctes broadbenti. This does not necessarily suggest that they are the same species. Our analysis suggests that it belongs to the genus Peroryctes, and so we reassigned it here, and provide a new diagnosis, based on features that can be seen on those worn specimens and the results of the morphometric analysis. New diagnosis. Peroryctes aruensis differs from Peroryctes raffrayana in having dimorphic upper and lower third premolars. This feature only occurs in Peroryctes broadbenti, and some Echymipera (most noticeable in E. clara). The third premolars are as long and wide as in P. broadbenti, which are narrower but longer than E. clara. It differs from P. raffrayana in having much larger teeth. It differs from P. broadbenti in having a shorter and less wide M4, having a reduced protocone; longer P2 and longer p1., Published as part of Travouillon, Kenny J. & Phillips, Matthew J., 2018, Total evidence analysis of the phylogenetic relationships of bandicoots and bilbies (Marsupialia: Peramelemorphia): reassessment of two species and description of a new species, pp. 224-256 in Zootaxa 4378 (2) on page 241, DOI: 10.11646/zootaxa.4378.2.3, http://zenodo.org/record/1169497, {"references":["Kear, B. P., Aplin, K. P. & Westerman, M. (2016) Bandicoot fossils and DNA elucidate lineage antiquity amongst xeric-adapted. Scientific Reports, 6, 37537. https: // doi. org / 10.1038 / srep 37537"]}

Isoodon fusciventer Gray 1841 new stat. Isoodon obesulus fusciventer Gray 1841 Holotype. BMNH 41.1176, skin and skull. Type locality. King George Sound, Western Australia, Australia Distribution. South-west Western Australia. Remarks. The results of our analysis show that the current subspecies of Isoodon obesulus do not form a monophyletic clade. This was also recovered in the analysis of Westerman et al. (2012), where Isoodon obesulus fusciventer does not form a clade with Isoodon obesulus obesulus suggesting that it is a distinct taxon. As a result, we propose to raise Isoodon obesulus fusciventer to species status, and provide a new diagnosis. Diagnosis. Isoodon fusciventer differs from all other Isoodon species in having a large StA on M1, with a long crest running anteroposteriorly (this cusp and crest are small and short in all other taxa). Isoodon fusciventer differs form I. obesulus and I. auratus in having a well-developed lingual shelf that extends to the buccal side on P3; no stylar crest on M1; retaining a StC/D on M4. Isoodon fusciventer differs from I. obesulus and I. macrourus in having fused StB and StC on M1 (these cusps are identifiable as distinct cusps in other taxa); posthypocristid is oblique to the tooth row on m1and m2; preentocristid on m4 is absent. Isoodon fusciventer differs from I. auratus in having an enlarged anterior cingulum on M1 which connects to the talon; having P1 as long as P2 (shorter in I. auratus). Isoodon fusciventer differs from I. obesulus in having a preparacrista posterobuccally orientated to connect to a posteriorly located StB on M1; having a ventrally angular alisphenoid tympanic process which terminates anterior to the transverse foramen. Isoodon fusciventer differs from I. macrourus in having P2 and P3 subequal in length (P2 is shorter in I. macrourus); retaining a stylar crest on M2; cristid obliqua on m1 terminates lingual to the protocone but buccal to the midpoint of the tooth width (buccal to the protocone in I. macrourus)., Published as part of Travouillon, Kenny J. & Phillips, Matthew J., 2018, Total evidence analysis of the phylogenetic relationships of bandicoots and bilbies (Marsupialia: Peramelemorphia): reassessment of two species and description of a new species, pp. 224-256 in Zootaxa 4378 (2) on page 235, DOI: 10.11646/zootaxa.4378.2.3, http://zenodo.org/record/1169497, {"references":["Gray, J. E. (1841) Contributions towards the geographical distribution of the Mammalia of Australia, with notes on some recently discovered species, in a letter addressed to the Author. Appendix C. In: Grey, G. (Ed.), Journals of Two Expeditions of Discovery in North-west and Western Australia, During the Years 1837, 38 and 39, Under the Authority of Her Majesty's Government. Describing many newly discovered, important, and fertile districts, with observations on the moral and physical condition of the aboriginal inhabitants, & c., & c. T. & W. Boone, London, pp. 397 - 414.","Westerman, M., Kear, B. P., Aplin, K., Meredith, R. W., Emerling, C. & Springer, M. S. (2012) Phylogenetic relationships of living and recently extinct bandicoots based on nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 62, 97 - 108. https: // doi. org / 10.1016 / j. ympev. 2011.09.009"]}

The phylogenetic relationships of bandicoots and bilbies have been somewhat problematic, with conflicting results between morphological work and molecular data. This conflict makes it difficult to assess the taxonomic status of species and subspecies within this order, and also prevents accurate evolutionary assessments. Here, we present a new total evidence analysis, combining the latest cranio-dental morphological matrix containing both modern and fossil taxa, with molecular data from GenBank. Several subspecies were scored in the morphological dataset to match the molecular data available. Both parsimony and Bayesian analyses were performed, giving similar topologies except for the position of four fossil taxa. Total evidence dating places the peramelemorphian crown origin close to the Oligocene/Miocene boundary, and the radiations of most modern genera beginning in the Late Miocene or Early Pliocene. Our results show that some species and subspecies require taxonomic reassessment, and are revised here. We also describe a new, extinct species from the Nullarbor region. This suggests that the number of recently extinct peramelemorphian species is likely to further increase.

Peramelemorphia comprises four families: the extant Peramelidae (bandicoots), and Thylacomyidae (bilbies); the recently extinct Chaeropodidae (pig-footed bandicoot); and the extinct Yaralidae; with at least ten fossil species of uncertain familial affinity designated as Perameloidea incertae sedis. Extant taxa (18 species) are characteristically omnivorous, small to medium sized (0.1-4.9 kg) semi-fossorial/fossorial marsupials with a quadrupedal bounding gait. They occupy varied habitats from desert to rainforest in Australia and New Guinea. Fourteen pre-Pliocene taxa are currently described on the basis of cranial and/or dental material, yet none is known from its postcranial skeleton. Here we use qualitative morphological and morphometric data to analyse a partial skeleton of a new species of bandicoot from a middle Miocene cave deposit, AL90 Site, in the Riversleigh World Heritage Area. AL90 has been radiometrically dated at 14.7-15.1 Ma, however the species is abundant in numerous early to middle Miocene deposits at Riversleigh The AL90 specimen preserves the skull and dentaries, the fore-and hindlimbs, and elements of the manus, pes and axial skeleton. The species is estimated to have weighed approximately 300g (based on predictive marsupial craniodental regression equations) which is comparable among extant bandicoots to Perameles bougainville (Western barred bandicoot). Unlike modern species, the appendicular skeleton of the fossil taxon is less well-adapted for scratch-digging and fossorial behaviours. A striking contrast is the relatively elongate, gracile bones of the antebrachium of the fossil taxon compared with the generally short, robust forearm of modern bandicoots. Collectively, our data indicate a more generalized niche for this species than crown group peramelemorphians and may support suggestions that archaic bandicoots filled an ecological niche later dominated by small dasyurids during the late Cainozoic.

Echymipera davidi Flannery, 1990 Seebeck, J. H. et al. (ed.), Bandicoots and Bilbies, p. 29, figs 1–3, table 1. (December 1990). Common name. David’s Echymipera. Current name. Echymipera davidi Flannery, 1990; following Groves (2005c). Holotype. M.7154 by original designation. Adult male, list No. 100, skull, study skin, collected by E. Le G. Troughton on 28 April 1945, registered 14 October 1945. Condition. Incomplete cranium, missing 1st, 2nd, 3rd incisors on both sides, and left upper 1st and 2nd post-canine teeth. Both dentaries missing all incisors, left dentary missing 4th and 5th post-canine teeth. Study skin: skin missing from near the tail tip, otherwise in good condition. Type locality. Kiriwina Island (8°30'S 151°00'E), Trobriand Island Group, Milne Bay Province, Papua New Guinea. The exact collecting locality on Kiriwina Island is not given in the register. Comments. The three paratypes are in AMNH. Good quality photographs of holotype and one paratype skull are given in the original description., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on page 314, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Groves, C. P. 2005 c. Order Peramelemorphia. In Mammal Species of the World: A Taxonomic and Geographic Reference, ed. D. E. Wilson and D. M. Reeder, pp. 38 - 42. Baltimore, USA: The Johns Hopkins University Press."]}

Didephilus obesula Shaw, 1797 Nat. Misc. 8: plate 298 and text. (August 1797). Common name. Southern Brown Bandicoot. Current name. Isoodon obesulus (Shaw, 1797), following Jackson & Groves (2015). Neotype. M.11821, designation by Dixon (1981). Young male [associated No. W4288], skull, study skin, intact hyoid bones, skinned body in alc., one intact flea in alc. Collected by A. B. Rose on 18 September 1977 and registered on 27 February 1981. Condition. Cranium complete; both dentaries have broken crowns of 2nd post-canine teeth. Study skin: bald patch on lower abdominal area, otherwise in good condition. Type locality. Ku-ring-gai Chase National Park, north of Sydney, NSW. Dixon (1981) gives the trap site of the neotype as “a point 1.5 km south of the Hawkesbury River, 7.5 km from the Coal and Candle Creek Road turnoff on the West Head Point Road, Kur-ring-gai Chase National Park, 33°36'S 151°16'E.” Comments. Dixon (1981) provides external and cranial measurements and a detailed description of the specimen and photographs of the skull and study skin., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on page 314, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Shaw, G. 1797. Text to plate 298. In The Naturalist's Miscellany: or coloured figures of natural objects, drawn and described immediately from nature. Volume 8, London: E. Nodder. https: // doi. org / 10.5962 / bhl. title. 79941","Dixon, J. M. 1981. Selection of a neotype for the southern short-nosed (brown) bandicoot, Isodon [sic] obesulus (Shaw and Nodder, 1797). Victorian Naturalist 98 (3): 130 - 135."]}

Rhynchomeles prattorum Thomas, 1920 Ann. Mag. Nat. Hist. (ser. 9) 6: 430. (1 October 1920). Common name. Seram Bandicoot. Current name. Rhynchomeles prattorum Thomas, 1920; following Groves (2005c). Paratype. M.29415, collectors no. 7, male, skull, study skin, collected by Pratt brothers in January 1920 from Mount Manusela, Seram Island, Maluku Province, Indonesia. Skin tag states “Alt. 6000 ft, trapped in heavy jungle”. Exchanged from BMNH August 1993, registered 12 October 1993, formally BMNH 1920.7.26.33. Comments. This specimen is from the original series of seven, all collected from the type locality by the Pratt brothers in 1920. The species does not seem to have been reported since. The specimen is considered to be a paratype rather than a referred specimen because Thomas based his species description on more than one individual, e.g., “a strongly contrasted white patch of very variable size on the chest”., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on page 318, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Thomas, O. 1920. On mammals from Ceram. Annals and Magazine of Natural History (series 9) 6: 422 - 431.","Groves, C. P. 2005 c. Order Peramelemorphia. In Mammal Species of the World: A Taxonomic and Geographic Reference, ed. D. E. Wilson and D. M. Reeder, pp. 38 - 42. Baltimore, USA: The Johns Hopkins University Press."]}

Isoodon arnhemensis Lyne & Mort, 1981 Aust. Mammal. 4(2): 128, figs 66–68. (cover date, 13 May 1981). Common name. Golden Bandicoot. Current name. Isoodon auratus arnhemensis Lyne & Mort, 1981; following Jackson & Groves (2015), who highlight the need for taxonomic clarification of this form. Holotype. M.6609 by original designation. Unknown sex, skull, study skin (Figs 15–16), presented by Rev. W. S. Chaseling, collection date not given, registered June 1939. Condition. Cranium missing right upper canine; left dentary missing last post-canine tooth; right dentary complete. Study skin: bald patch on rump, bald area on left side of abdomen. Study skin in good condition but excessive elongation does not reflect correct body proportions. Cranial measurements (mm). M.6609: ConL, 63.67; BasL, 60.56; NasL, 25.29; NasB, 5.34; UC1–C1 (alv.), 9.69; APV, 4.10; PPV, 6.48; PAL, 37.06; UPM (alv.), 2.80; UMR (alv.), 11.79; ZB, 30.31; POC, 11.18; BUL, 13.76; MB, 25.27; DL (angl.), 50.55; DL (condyl.), 48.68; LPM (alv.), 2.81; LMR (alv.), 13.70. Type locality. Melville Bay and Cape Arnhem area, Northern Territory, Australia. The type locality in the original description is given as “Melville Bay, Cape Arnhem area, Northern Territory” but the locality given in the register for each of the three specimens is actually two combined localities “Melville Bay and Cape Arnhem area”, which are separated by c. 25 km. Paratypes. (2, by original designation). Details for both as per holotype: M.6606, male, skull, study skin; M.6610, indeterminate sex, skull, study skin. Comments. Three specimens in the type series.

Parnaby, Harry E., Ingleby, Sandy, Divljan, Anja (2017): Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney. Records of the Australian Museum 69 (5): 277-420, DOI: 10.3853/j.2201-4349.69.2017.1653, URL: http://dx.doi.org/10.3853/j.2201-4349.69.2017.1653

Perameles broadbenti Ramsay 1879a Proc. Linn. Soc. N.S.W. (ser. 1) 3(4): 402, figs 1–15, plate 27. (May 1879). Common name. Giant Bandicoot. Current name. Peroryctes broadbenti (Ramsay, 1879a), following Aplin et al. (2010). Holotype. A.3238, by subsequent determination. Male, skull, skin mount, originally in alc., purchased from Kendall Broadbent, registered November 1878. Collection date not given. Condition. Cranium has fractured left maxilla, right zygomatic arch glued in place, missing left upper 5th postcanine tooth, missing left upper 1st and 3rd incisors, and right upper 2nd, 3rd and 4th incisors; right dentary missing coronoid process, both dentaries have drilled wire holes through both condyles. Skin mount missing ear tips, bald patch on the left side, fracture on the base of the tail, left manus digits 4 and 5 fractured, sparse fur on the ventral surface (stomach area). Type locality. Port Moresby district, Central Province, Papua New Guinea. Specimen obtained from “a considerable distance inland from Port Moresby, in some of the dense scrubs on the banks of the Goldie River” (Ramsay, 1879a). Comments. Ramsay states that his description is based on a single male. Detailed measurements and high quality photographs of the holotype skull are given by Aplin et al. (2010), who review the limited number of specimens available in world collections., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on page 316, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Ramsay, E. P. 1879 a. Contributions to the zoology of New Guinea. Part III. Description of a new marsupial allied to the genus Perameles, Geoff. Proceedings of the Linnean Society of New South Wales (series 1) 3 (4): 402 - 405, pl. 27.","Aplin, K. P., K. M. Helgen, and D. Lunde. 2010. Review of the morphology, distribution and conservation status of Peroryctes broadbenti (Ramsay, 1879), the giant bandicoot of south-eastern Papua New Guinea. American Museum Novitates 3696: 1 - 41. https: // doi. org / 10.1206 / 3696.2"]}

Perameles ecaudatus Ogilby, 1838a (Ogilby, 1838a) Proc. Zool. Soc. Lond. 1838: 25. (July 1838) Common name. Pig-footed Bandicoot. Current name. Chaeropus ecaudatus (W. Ogilby, 1838a), following Jackson & Groves (2015). Holotype. Specimen number 35 in the catalogue of Bennett (1837), according to Mahoney & Ride (1988c). The original specimen was collected on 16 June 1836 (Ogilby, 1838a) and cited as female by Bennett (1837). Comments. Ogilby (1838a) proposed the species on the basis of drawings and notes supplied by Mitchell, which Ogilby states were based on a specimen that had already been lodged in the AM. Only one specimen was obtained on the Murray River, NSW, by Sir Thomas Mitchell (Mahoney & Ride, 1988c). Krefft (1864a: 35) lists specimen “b”, as a skull of the original specimen obtained by Mitchell, and states (Krefft, 1870) that he found the skull “10 years ago” but that he had never sighted the skin of Mitchell’s specimen, which he assumed had deteriorated through neglect prior to his curatorship. Ogilby (1892) lists the type as being in the AM Collection, but it is not clear if he actually sighted the specimen or drew from Krefft’s catalogue, given that much of his work appeared to be a desktop study (see Iredale & Troughton, 1934). Troughton (1932d: 179) believed that the holotype had disappeared from the collection., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on page 403, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Ogilby, W. 1838 a. On a new species of marsupial animal found by Major Mitchell on the banks of the River Murray in New South Wales. Proceedings of the Zoological Society of London 6 (1838): 25 - 27.","Bennett, G. 1837. Catalogue of the Specimens of Natural History and Miscellaneous Curiosities Deposited in the Australian Museum. Sydney: James Tegg and Co. 71 pp.","Krefft, G. 1864 a. Catalogue of Mammalia in the collection of the Australian Museum. Australian Museum Catalogue No. 2. Sydney: Trustees of the Australian Museum. 2 + 134 + 2 pp.","Krefft, G. 1870. Letter to the Editor (re Choeropodus Ecaudatus). Australian Town and Country Journal II (42): 17, col. 1, 22 October.","Ogilby, J. D. 1892. Catalogue of Australian Mammals, with Introductory Notes on General Mammalogy. Sydney: Trustees, Australian Museum. xvi + 142 pp. https: // doi. org / 10.5962 / bhl. title. 1130","Troughton, E. L. G. 1932 d. Australian furred animals, their past, present and future. Australian Zoologist 7 (3): 173 - 193."]}

Perameles cockerelli Ramsay, 1877c [not 1876] Proc. Linn. Soc. N.S.W. (ser. 1) 1(4): 310. (March 1877). Common name. Common Echymipera. Current name. Echymipera kalubu cockerelli (Ramsay, 1877c), following Groves (2005c). Syntype. PA.471, by subsequent determination. Male, skin mount, skull in situ, purchased from James Cockerell, registered in the Palmer Register in c. 1878. Original AM metal tag with “471” attached to the right hind leg. The former existence of at least one other syntype is inferred but its present whereabouts is unknown. Condition. Skin mount (skull in situ), faded, hole in the left side, torn ear tips and missing tail tip. Type locality. Given as “ New Ireland ” (Papua New Guinea) by Ramsay. Flannery (1995c) considered the type locality to be an error, based on the current absence of bandicoots from New Ireland, and the fact that many natural history specimens obtained by Brown had inaccurate locality data, being either from New Ireland, New Britain or one of the islands of the Duke of York group. Comments. Ramsay did not indicate the number of specimens examined. The taxidermist’s department lists preparation of “1 Perameles cockerelli sp nov” as a mount, in the AM annual report for 1876 (Ramsay, 1877h). We regard PA.471 to be a syntype and not a holotype because Ramsay appears to have based his description on more than one specimen. Ramsay’s account of the species, (which was based on external characters only), includes body measurements for two individuals but it is not clear whether both specimens were in the AM at that time or whether he was using measurements from another specimen, perhaps in the possession of Cockerell or George Brown. Whittell (1954: 155) states that all of Brown’s collection was sent to London, and that only Cockerell’s collection remained in Sydney. However, a few of Brown’s specimens (without data) were recently found in private hands in Sydney (I. McAllen, pers. comm. 2016). Ramsay (1877i) provided further brief notes on colouration and notes on dentition that would have been visible from a skin mount. Miklouho-Maclay (1884) illustrated the head of a specimen of cockerelli and implied (1884: 719) that there was only one specimen, a skin mount, in theAM Collection. His drawing is consistent with the head of the skin mount of PA.471., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on page 318, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Groves, C. P. 2005 c. Order Peramelemorphia. In Mammal Species of the World: A Taxonomic and Geographic Reference, ed. D. E. Wilson and D. M. Reeder, pp. 38 - 42. Baltimore, USA: The Johns Hopkins University Press.","Flannery, T. 1995 c. Mammals of the South-West Pacific and Moluccan Islands. Chatswood, New South Wales: Australian Museum / Reed Books. 464 pp.","Ramsay, E. P. 1877 h. Appendix IX, Taxidermist's Department. In Report of the Trustees for the year ending 31 December, 1876, Australian Museum, pp. 12 - 13. Sydney, NSW: Charles Potter, Acting Government Printer.","Whittell, H. M. 1954. The Literature of Australian Birds: a History and a Bibliography of Australian Ornithology. Perth, Western Australia: Paterson Brokensha. xi + 140 + 788 pp. [Limited edition reprinted by Maurizio Martino, Mansfield, CT, USA.]","Miklouho-Maclay, N. d. 1884. Notes on zoology of the Maclay-Coast in New Guinea. I. On a new sub-genus of Peramelidae: Brachymelis. Proceedings of the Linnean Society of New South Wales (series 1) 9 (3): 713 - 720, pl. 38."]}

Perameles macroura var. torosus Ramsay, 1877a Proc. Linn. Soc. N.S.W. (ser. 1) 2(1): 12. (July 1877). Common name. Northern Brown Bandicoot. Current name. Isoodon macrourus torosus (Ramsay, 1877a), following Jackson & Groves (2015). Syntypes. Type material for this taxon has not been reported. As detailed below, it is likely that Ramsay used more than one specimen for his description and that a syntype series once existed. Syntypes are likely to have originally been in the collections of W. Macleay, so possibly now in the Macleay Museum, in the AM, or in Ramsay’s private collection. It is possible that syntypes were sent to other institutions by Ramsay. Type localities. Include Rockingham Bay [Cardwell] and Endeavour River [Cooktown], Qld, Australia. Comments. Past interpretations of Ramsay’s description have assumed a holotype (e.g., Lyne & Mort, 1981; Mahoney & Ride, 1988c) but it is likely that his description involved syntypes. Ramsay’s description focused on a female specimen obtained near Cooktown but the fact that he gave the distribution as Rockingham Bay [Cardwell] to Endeavour River [Cooktown] suggests that he was aware of other specimens. More ambiguous is his statement that “For the present I prefer to place the specimens, showing these peculiarities, as a variety [Ramsay’s emphasis] of P. macroura …”—this could refer to either existing or future material. Troughton (as Anonymous, 1927) states that “the specimen” of torosus described by Ramsay was collected during the Chevert Expedition and was in the Macleay Museum, although whether this is based on first hand knowledge or surmise is unclear. We are not aware that the Chevert party collected at Endeavour River and it seems more likely that the type or syntypes were obtained on collecting trips to the region around that time by several different collectors. Edward Spalding collected material for Macleay at Endeavour River from June to October 1874 (Whittell, 1954), and in July of that year sent specimens that Macleay termed “skins of a new bandicoot” (Fletcher, 1929: 230). Ramsay also obtained mammals from the Rockingham Bay district and visited the area himself on a number of occasions, e.g., in 1874 (Ramsay, 1876c). It appears that all taxonomic assessments of this taxon have relied on Ramsay’s original description and that the types have remained unlocated., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on page 406, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Ramsay, E. P. 1877 a. Note of a species of Echidna (Tachyglossus) from Port Moresby, New Guinea. Proceedings of the Linnean Society of New South Wales (series 1) 2 (1): 31 - 34, 1 pl.","Anonymous. 1927. Notes and Exhibits, Meeting of 27 October, 1926 [Exhibition of a large bandicoot from the Clarence River]. Proceedings of the Linnean Society of New South Wales 51 (5): xlii.","Whittell, H. M. 1954. The Literature of Australian Birds: a History and a Bibliography of Australian Ornithology. Perth, Western Australia: Paterson Brokensha. xi + 140 + 788 pp. [Limited edition reprinted by Maurizio Martino, Mansfield, CT, USA.]","Fletcher, J. J. 1929. The Society's heritage from the Macleays. Part 2. Proceedings of the Linnean Society of New South Wales 54 (3): 185 - 272.","Ramsay, E. P. 1876 c. Description of a new species of Rat Kangaroo, allied to the genus Hypsiprymnus, proposed to be called Hypsiprynodon moschatus. Proceedings of the Linnean Society of New South Wales (series 1) 1 (1): 33 - 35."]}

Perameles moresbyensis Ramsay, 1877a Proc. Linn. Soc. N.S.W. (ser. 1) 2(1): 14. (July 1877). Common name. Northern Brown Bandicoot. Current name. Isoodon macrourus moresbyensis (Ramsay, 1877a), following Groves (2005c). Syntype. M.2554, by subsequent determination. Male, skull (Fig. 17) and skin mount, re-registered in March 1915 from the “old collection”, but previous registration number (if any) or previous registration date not indicated. Condition. Cranium missing rear of braincase including occipitals and much of the parietals; missing left zygomatic arch, missing posterior end of left auditory bulla, hole in the left maxilla through part of the palate, teeth are reinforced with glue; both dentaries complete. Skin mount missing tail tip, missing right ear tip, hole in the side of left hind leg, fur on the ventral surface appears to have been glued on. Cranial measurements (mm). M.2554: GL, —; NasL, 31.62; NasB, 8.06; UC1–C1 (alv.), 11.30; APV, 7.19; PAL, 46.85; UPM (alv.), 4.01; UMR (alv.), 15.45; ZG, —; POC, 11.60; BUL, 13.29 (right side bulla); MB, 25.07; DL (angl.), 62.05; DL (condyl.), 61.30; LPM (alv.), 3.67; LMR (alv.), 16.21. Type locality. Port Moresby area, Central Province, Papua New Guinea. Comments. Ramsay states that “the animal examined was full grown, but young”. Although a single specimen is implied, it is not possible to determine from the original description if he examined other specimens, perhaps belonging to other collectors, but omitted the collector’s name in the original account, and in the M Register. The dentition of M.2554 is inconsistent with Ramsay’s description in that the third lower premolar appears to be smaller, not larger, than the lower canine. Dental and external measurements of M.2554 broadly concur with those given by Ramsay, taking into account the uncertainty of how some measurements were made and whether or not skin measurements were taken before the specimen was prepared as a mount. However, our impression is that M.2554 is a larger animal than that measured by Ramsay, e.g., hind foot length taken from nail to heel is c. 3 inches (vs 2.5). We have not yet been able to trace the provenance of this specimen prior to its registration in 1915. However, Thomas (1888a: 235) stated that the type was in the “Sydney Museum” (as opposed to the Macleay Museum). Although included in his paper on mammals of the 1875 Chevert Expedition (Ramsay, 1877a), material of this taxon was most likely obtained soon after the Expedition. Macleay (1875) states that several members of the Chevert crew had left the Expedition which ended at Yule Island, with the aim of collecting in the Port Moresby district., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on pages 316-318, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Ramsay, E. P. 1877 a. Note of a species of Echidna (Tachyglossus) from Port Moresby, New Guinea. Proceedings of the Linnean Society of New South Wales (series 1) 2 (1): 31 - 34, 1 pl.","Groves, C. P. 2005 c. Order Peramelemorphia. In Mammal Species of the World: A Taxonomic and Geographic Reference, ed. D. E. Wilson and D. M. Reeder, pp. 38 - 42. Baltimore, USA: The Johns Hopkins University Press.","Thomas, O. 1888 a. Catalogue of the Marsupialia and Monotremata in the Collection of the British Museum (Natural History). London: British Museum (Natural History). xii + 401 pp. https: // doi. org / 10.5962 / bhl. title. 56181","Macleay, W. J. 1875. The voyage of the Chevert. The Sydney Morning Herald LXXII (11,667): 5, col. 2, 11 October."]}

Echymipera philipi Troughton, 1945 Rec. Aust. Mus. 21(6): 373. (25 June 1945). Common name. Common Echymipera. Current name. Echymipera kalubu philipi Troughton, 1945; following Groves (2005c). Holotype. M.6999 by original designation. Male, [Field no. Owi Is. No. 29], skull (Fig. 14), study skin, collected by Lieutenant-Colonel Cornelius B. Philip and Major Glen M. Kohls, collection date not given, registered September 1944. Condition. Cranium missing right upper 1st and 2nd incisors; both dentaries complete. Study skin in good condition. Type locality. Owi Island, Schouten Island Group, Cenderawasih (formerly Geelvink) Bay, Papua Province, Indonesia. Paratypes. (3, 2 by subsequent determination). Same locality and collectors as holotype, all registered February 1945, collection dates not given: M.7028 by original designation, female, skull, study skin (allotype); M.7029 (two specimens), female, body in alc. with unfurred female pouch young in alc.; M.7030, male, skull, body in alc. Comments. Troughton does not indicate the number of paratypes but implies that there are three, without citing registration numbers for two specimens. Paratypes are identified as such in Troughton’s writing on specimen tags for the three specimens listed above and a search of the register database indicates a total of four specimens of this species from Owi Island., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on page 315, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Troughton, E. L. G. 1945. Diagnoses of new mammals from the South-West Pacific. Records of the Australian Museum 21 (6): 373 - 376. https: // doi. org / 10.3853 / j. 0067 - 1975.21.1945.551","Groves, C. P. 2005 c. Order Peramelemorphia. In Mammal Species of the World: A Taxonomic and Geographic Reference, ed. D. E. Wilson and D. M. Reeder, pp. 38 - 42. Baltimore, USA: The Johns Hopkins University Press."]}

The paralysis tick Ixodes holocyclus bites humans, companion animals, and livestock in eastern Australia leading to symptoms that range between negligible and severe. Bandicoots (Family Peramelidae) are commonly cited as the “primary host” of I. holocyclus in the media and blamed for outbreaks of ticks and disease fears, creating conflicts between conservation and tick management. We discuss how evidence for bandicoots being essential to the I. holocyclus life cycle has originated from a small number of papers that were limited in scope. False assumptions of host-specificity have contributed to the extrapolation of studies in one ecosystem, yet no study has sampled the full range of hosts of I. holocyclus to understand the relative role of each species across the entire range of I. holocyclus in relation to health threats. Bandicoots are one of many potential tick hosts but cannot yet be considered the “primary host” of I. holocyclus. Researchers and media should refrain from highlighting bandicoots as th...