The Chinchilla Sand is a formally defined stratigraphic sequence of Pliocene fluviatile deposits that comprise interbedded clay, sand, and conglomerate located in the western Darling Downs, south-east Queensland, Australia. Vertebrate fossils from the deposits are referred to as the Chinchilla Local Fauna. Despite over a century and a half of collection and study, uncertainties concerning the taxa in the Chinchilla Local Fauna continue, largely from the absence of stratigraphically controlled excavations, lost or destroyed specimens, and poorly documented provenance data. Here we present a detailed and updated study of the vertebrate fauna from this site. The Pliocene vertebrate assemblage is represented by at least 63 taxa in 31 families. The Chinchilla Local Fauna is Australia's largest, richest and best preserved Pliocene vertebrate locality, and is eminently suited for palaeoecological and palaeoenvironmental investigations of the late Pliocene.
Introduction
The Pliocene was the time when most modern Australian mammal genera emerged and began to dominate ecosystems (Black et al. 2012b). While the majority of Earth's landmasses were similar to those of today, by the middle Pliocene global mean annual temperatures were approximately 3ºC higher than today (Chandler et al. 1994; Sloan et al. 1996). The late Pliocene (ca. 3.6-2.6 million years ago) represents the last time when the Earth experienced temperatures similar to those predicted for the end of the 21st century (Salzmann et al. 2009). This period represents “an unparalleled paleo-laboratory for testing the sensitivity of models that we rely upon for simulating future climate change” (Salzmann et al. 2009: 190).
In Australia, the Pliocene marked a major turning point in the evolution of terrestrial vertebrates. The late Miocene was characterized by cool and dry conditions (Zachos et al. 2001). Global mean annual temperatures rose during the early to middle Pliocene (Haywood et al. 2009), and several lines of evidence point towards a Pliocene aridification trend in Australia (Krebs et al. 2011). As a result of this shift, woodlands contracted markedly and the first significant grasslands developed (Martin 2006). The transition from closed to more open vegetation saw a major evolutionary change in terrestrial faunas, including not only an increase in the diversity of grazing (grass-eating) species, but also gigantism of several lineages such as the herbivorous diprotodontoids and macropodids (Price and Piper 2009). The Pliocene also saw the emergence of Macropus, the long-faced kangaroos that are the most speciose macropodids in Australia today.
Understanding how mammal species responded to Pliocene warming is critical for predicting how future global warming might impact modern species and for “ground-truthing” climate models that seek to simulate future change. Unfortunately, Pliocene vertebrate fossil deposits are relatively rare in Australia, especially in comparison to Miocene and Pleistocene assemblages, making it difficult to track the trends towards modernity of Australian late Cenozoic faunas (Tedford et al. 1992, 2006). One of the richest deposits with Pliocene vertebrates is the Chinchilla Sand, Darling Downs, southeastern Queensland (Fig. 1). This deposit has produced thousands of fossils, many now housed at the Queensland Museum, making it the largest single collection of Pliocene fossil vertebrates in Australia (Price 2012). Many new species have been described from Chinchilla material, and most previous studies have focussed on the taxonomy of marsupials, such as macropodids (e.g., Bartholomai 1973, 1975).
Fig. 1.
Physiography of the Chinchilla local area, showing the location of Chinchilla relative to the Condamine River System. Redrawn after Reiser (1971).
Despite a voluminous literature on the Pliocene fauna from the Chinchilla Sand, its palaeontological significance is poorly understood. This is because: (i) few studies have provided a detailed overview of the total fauna; (ii) type and referred specimens of some taxa have either not had registration numbers assigned, or numbers have been incorrectly cited; (iii) type material of some taxa has been either destroyed or is missing from collections; and (iv) many taxa supposedly from Chinchilla have poorly constrained collecting information. The result is some uncertainty and probable errors regarding the species present in the fauna (e.g., Mackness and Godthelp 2001). Accurate species lists are critical for making palaeoecological inferences, reconstructing palaeoenvironments and exploring changes in animal communities over time and in response to climatic events. The aim of this paper is to integrate the extensive taxonomic literature of the Pliocene Chinchilla Local Fauna.
Institutional abbreviations.—BMNH, British Museum of Natural History, London, UK; Cat. No., Queensland Museum Category Number (refers to the old Queensland Museum indexing system), Brisbane, Australia; FV, Private collection, Terry Poole, Brisbane, Australia; QMC, Queensland Museum Collection, Brisbane, Australia; QM F, Queensland Museum Fossil, Brisbane, Australia; UCR, Department of Earth Sciences, University of California, Riverside, USA; WPC, Wilkinson Private Collection, Chinchilla, Australia.
Geographic and geological setting
The fossil deposits of the Darling Downs are divided roughly into two major collecting areas: the eastern Darling Downs which contains predominantly Quaternary fossil deposits (Price et al 2005; Webb et al. 2007; Price et al. 2011); and the western Darling Downs (including the township of Chinchilla and surrounds; Fig. 1), which are mostly Pliocene (Bartholomai and Woods 1976). The Chinchilla Sand, previously referred to as the Chinchilla Formation by Woods (1956) and incorporating the Chinchilla Conglomerate of Etheridge (1892), is a series of fluviatile deposits exposed along the Condamine River, extending over 65 km from Nangram Lagoon to Warra (Fig. 1) (Bartholomai and Woods 1976; Price 2012).
The Chinchilla Sand is a thick (∼30 m) sequence of interbedded gravel, sand, and clay exposed in many inter-connected gullies and erosional depressions representing multiple episodes of deposition (Fig. 2). The sand and clay are typically weakly consolidated, whereas the associated gravel conglomerate is heavily cemented by calcium carbonate. The sediments are most likely derived from the Orallo Formation and associated lateritized soil profiles (Bartholomai and Woods 1976). The Chinchilla Sand sits unconformably on Mesozoic rocks (Bartholomai and Woods 1976), and is overlain by Quaternary alluvia in some parts.
Fig. 2.
Part of the Chinchilla gully system in the Chinchilla Rifle Range, showing the dominant sedimentary units present.
Vertebrate fossils from the Chinchilla Sand are typically referred to as belonging to the Chinchilla Local Fauna (Archer and Bartholomai 1978), although this likely time-averages several faunal components. Specific provenance data is missing for most specimens recovered from the Chinchilla region. Where specific locality information is available, this is provided. Where it is not, we have simply listed the specimens as coming from “Chinchilla”, which encompasses the area illustrated in Fig. 1. The Chinchilla Sand has not been directly dated, but the faunal composition is similar to that of the Kanunka and Toolapinna Local Faunas of the Tirari Formation (Tedford et al. 1992), Lake Eyre Basin, central Australia. Combined lithostratigraphic, biostratigraphic and palaeomagnetic analyses suggest that these faunas occur close to the Gilbert-Gauss Boundary (Tedford et al. 1992), at approximately 3.6 Ma (Piacenzian-Zanclean boundary; Ogg 2012). The paleoenvironment of Chinchilla during the period of fossil deposition included tropical forests, wetlands, and grasslands (Montanari et al. 2013).
Associated with the vertebrate fauna are fossil plants of putative Tertiary age. Silicified woods, including Mesembrioxylon fusiforme and M. fluviale, were described from the Condamine River, west of Fairymeadow, Chinchilla (Sahni and Dunstan 1920). In a foreword of their paper, Sahni and Dunstan (1920) noted that while the silicified wood occurred with the vertebrates, it might have been derived from the underlying Jurassic sediments. The silicified wood has been referred to the form-genus Mesembrioxylon, which is recorded from both Mesozoic and Tertiary sediments. Rigby (1995) also recorded a silicified Cocos nucifera fruit from the Pliocene sediments at Chinchilla, but Conran and Rozefelds (2003) queried this identification, noting that as the specimen had not been sectioned its affinities were unsubstantiated. They also similarly noted, as Sahni and Dunstan (1920) had much earlier, the likelihood that the silicified material was reworked from Jurassic or older sediments, which makes the identification of the fruit as a palm highly unlikely. Fragments of non-silicified, calcium carbonate-indurated wood occur with the vertebrate material and are possibly Pliocene in age (Andrew Rozefelds, personal observation 2013). The Chinchilla plants are probably irrelevant in understanding the palaeoecology of the Chinchilla Sand if the material is reworked from older sediments (Andrew Rozefelds, personal communication 2013).
Methods
We have attempted to identify every taxon that has been published from the Pliocene Chinchilla Sand, with the exception of those mentioned only in conference abstracts. We provide an abbreviated systematic palaeontology to the lowest taxonomic levels possible; a list of representative registered specimens for each taxon (where possible); the published authority of each taxon's record; illustrations of representative fossil material as assigned to each taxon (where possible); and brief but pertinent taxonomic, systematic and/or literary remarks for each taxon. Although every effort was made to locate every taxon published from Chinchilla, some specimens are missing from collections. This is noted. Provenance of specimens was judged on the basis of collection notes, museum registers, and the publications in which the specimens were described. Informal localities are listed in quotation marks. Most fossils recovered since the 1960s have been collected in the gully systems of the Chinchilla Rifle Range and surrounding areas.
Systematic palaeontology
Class Sarcopterygii Romer, 1955
Subclass Dipnoi Müller, 1845
Family Ceratodontidae Gill, 1873
Genus Metaceratodus Chapman, 1914
Type species: Metaceratodus wollastoni Chapman, 1914, Lightning Ridge, New South Wales, Early Cretaceous.
Metaceratodus palmeri (Krefft, 1874)
Fig. 3A.
Material.—QM F10537 (upper tooth plate) and QM F10540 (upper jaw) from unspecified locality near Chinchilla, Australia, Pliocene; QM F6564 (upper jaw) from the property adjacent to the Chinchilla Rifle Range, Australia, on the eastern fence in gully paralleling the Middle Gully, Pliocene.
Remarks.—The extant lungfish Neoceratodus forsteri was described by Krefft (1870). Later, Krefft (1874: 293) acknowledged receipt of the cast of an extinct species of lungfish found “in the alluvial deposits of the Darling Downs district of Queensland”, but without additional locality information. According to the note, Krefft had named the species Ceratodus palmeri, after the then Colonial Secretary of Queensland; however, according to de Vis (1884), Krefft did not publish a formal description of the taxon. Further, de Vis (1884) suggested that the range of variation exhibited by N. forsteri was sufficient to account for the differences between this species and Metaceratodus (then still Ceratodus) palmeri, and synonymised both taxa. He also provided information on the provenance of additional specimens attributable to M. palmeri, including Krefft's (1874) holotype, stating that they came from the “Chinchilla conglomerate” (= Chinchilla Sand). Metaceratodus palmeri was subsequently mentioned by Turner (1982) and Lees (1986), in their reviews on fossil fish held in the Queensland Museum, Brisbane, Australia (QM). Kemp and Molnar (1981), as part of a study on lungfish from Lightning Ridge, New South Wales, re-examined the synonymy of N. forsteri and M. palmeri, and concluded that they were, in fact, distinct species. That taxonomy was later followed in Kemp's (1991) subsequent review of Australian lungfish. Kemp (1997b) later reassigned the original Chinchilla fossil lungfish to the genus Metaceratodus, including referral of QM F10537 as a cast of the holotype of Metaceratodus palmeri (BMNH 45868). However QM F10537 (Fig. 3A) is not a cast; as such, the reason for its designation as a copy of the holotype is unclear.
Stratigraphic and geographic range.—Pliocene to Pleistocene; eastern mainland Australia.
Fig. 3.
Lungfish toothplates from unspecified locality near Chinchilla, Australia, Pliocene. A. Metaceratodus palmeri (Krefft, 1874), QM F10537. B. Neoceratodus forsteri (Krefft, 1870), QM F56224. Scale bars 10 mm.
Family Neoceratodontidae Miles, 1977
Genus Neoceratodus de Castelnau, 1876
Type species: Neoceratodus forsteri (Krefft, 1870), Burnett River, Queensland, Recent.
Neoceratodus forsteri (Krefft, 1870)
Fig. 3B.
Material.—QM F10539 (three toothplates) from Chinchilla, Australia, Pliocene; QM F56224 (toothplate) from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Two fossil lungfish specimens (FV O355 and FV O616) from Chinchilla were assigned to Neoceratodus forsteri by Kemp (1997a). Unfortunately, both specimens are from the private collection of Terry Poole, and thus not easily accessible. Three toothplates, registered together as QM F10539, (originally described as M. palmeri) are referable to N. forsteri.
Stratigraphic and geographic range.—Cretaceous to Recent; eastern mainland Australia.
Class Reptilia Laurenti, 1768
Order Testudines Linneaus, 1758
Family Trionychidae Fitzinger, 1826
Trionychidae gen. et sp. indet.
Fig. 4D.
Material.—QM F9037 (carapace fragment) from Fairymeadow near Chinchilla, Australia, probably Pliocene.
Remarks.—De Vis (1894b) described seven carapace fragments of a single individual (QM F1101) as Trionyx australiensis from the Darling Downs, possibly from the Chinchilla Sand. However, in the absence of locality data, the preservation led Alan Bartholomai (Gaffney and Bartholomai 1979) to suggest that they may have instead come from Tara Creek, Queensland. Gaffney and Bartholomai (1979) concluded that T. australiensis was only known from small carapace fragments and referred the taxon to Trionychidae indeterminate. They later described a specimen referred to Trionychidae sp. indet. from the “Southwest of Chinchilla”, and hence presumably from the Chinchilla Sand. That specimen was also listed in Gaffney's (1981) review of Australian turtles. Molnar (1982a) in his review of the reptile fossil material of Queensland lists only Emydura sp. as definitively from the Chinchilla Sand, although he also refers the Trionychidae sp. indet. as coming from Fairymeadow near Chinchilla.
Family Chelidae Gray, 1831
Genus Emydura Bonaparte, 1836
Type species: Emydura macquarrii (Gray, 1830), Maquarie River, New South Wales, Recent.
Emydura sp.
Fig. 4E.
Material.—QM F7035 (anterior half of carapace) and QM F7034 (anterior half of plastron) from Middle Gully of Chinchilla Rifle Range, Australia, Pliocene; QM F9038 (symphyseal region) and QM F9039 (left dentary) likely from Chinchilla, Australia, Pliocene.
Remarks.—Gaffney (1981) described QM F7034 from Middle Gully, near the type section of the Chinchilla Sand, QM F7035 from Chinchilla Sand, while QM F9038 and F9039 are also presumed to be from Chinchilla Sand based on preservation (Alan Bartholomai, personal communication to Gaffney 1981).
Order Crocodylia Owen, 1842
Family Crocodylidae Cuvier, 1807
Genus Pallimnarchus de Vis, 1886
Type species: Pallimnarchus pollens de Vis, 1886, type locality and age unknown.
Pallimnarchus pollens de Vis, 1886
Fig. 4F.
Material.—QM F11612 (symphyseal portion of dentary) from “Sand Scree locality, 4′6″ from top of bank of Condamine River”, Chinchilla, Australia, Pliocene; QM F1154 (premaxilla) and QM F1166 (quadrate) from Chinchilla, Australia, Pliocene; QM F30583 (femur) from north bank of the Condamine River, Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—De Vis (1886) described crocodilian material from the “Condamine drift” to which he assigned the name Pal limnarchus pollens. Molnar (1982b: 658) states that “De Vis did not designate a type specimen for Pallemnarchus [sic!] pollens, but described together material from four (possibly five) individuals of at least two, probably three, different taxa. De Vis' intention with regard to the type material cannot be deduced from the labels or the register entries, as the former treat all the specimens described by de Vis (1886), with the exception of the maxillary piece, as syntypes, while the register designates QM F1149 (two articulated dentaries) as the type. The maxillary piece figured by de Vis (1886: pl. 10) was not recognised in the collections.” Molnar (1982b) formalised the species Pallimnarchus pollens by designating a symphyseal portion of the dentary as the lectotype (QM F1149), noting that while the type locality was unknown, it was probably from the Darling Downs. He did refer several Chinchilla specimens (including QM F11612, F1154, and F1166) to this species. Willis and Molnar's (1999) revision of this genus does not change that. Subsequently, Mackness et al. (2010) added an additional pathological specimen (QM F30583) to the hypodigm (a femur showing fracture and puncture wounds).
Stratigraphic and geographic range.—Pliocene to Pleistocene; central and eastern mainland Australia.
Fig. 4.
Reptiles from Chinchilla Sand, Australia, Pliocene. A. ?Reptilia, QM F56197, Chinchilla Rifle Range, unidentified fragments (A1–A3), showing the WPC registration numbers of Agamidae sp. indet. cited by Hutchinson and Mackness (2002). B. Varanus sp. QM F56189, “Wilkinson's Quarry” Chinchilla Rifle Range, dorsal vertebra. C. Quinkana sp., QM F10204, Chinchilla Rifle Range, isolated tooth. D. Trionychidae sp. indet. QM F9037, Fairymeadow, carapace fragment. E. Emydura sp., QM F7035, Middle Gully, Chinchilla Rifle Range, anterior half of carapace. F. Pallimnarchus pollens de Vis, 1886, QM F11612, “Sand Scree locality, 4′6″ from top of bank of Condamine River”, symphyseal portion of dentary. G. Varanus komodoensis Ouwens, 1912, QM F874, unspecified locality near Chinchilla, right maxilla. Scale bars 20 mm.
Genus Quinkana Molnar, 1981
Type species: Quinkana fortirostrum Molnar, 1981, Tea Tree Cave near Chillagoe, Queensland, Pleistocene.
Quinkana sp.
Fig. 4B.
Material.—QM F10204 (tooth) and QM F10205 (tooth) from Chinchilla Rifle Range, Australia, Pliocene; QM F1152 (jugal) from Chinchilla, Australia, Pliocene.
Remarks.—Molnar (1981) refers to two isolated teeth and a jugal from a ziphodont crocodile from the Chinchilla Sand. He considered them to be referrable to Quinkana.
Order Squamata Oppel, 1811
Family Agamidae Hardwicke and Gray, 1827
Agamidae gen. et sp. indet.
Fig. 4A.
Material.—QM F56197, unidentified fragments from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Specimens supposed to represent agamids by Hutchinson and Mackness (2002) previously held in a private collection (WPC) were accessioned into the collections of the QM (see SOM: Table 1, Supplementary Online Material available at http://app.pan.pl/SOM/app60-Louys_Price_SOM.pdf). However the specimen referred to the Agamidae by Hutchinson and Mackness (2002; WPC1354) does not correspond with the specimen of that number donated to the QM by the collectors (Ces and Doris Wilkinson, WPC). The specimen in question (now registered as QM F56197; Fig. 4A) is a damaged and poorly preserved fossil in three pieces which can at best be referred to indeterminate ?Reptilia. A similar error most likely explains the incorrect assignment of several specimens referred to as ?Megalania by Hutchinson and Mackness (2002), which are clearly not squamates (see below).
Family Gekkonidae Gray, 1825
Genus Diplodactylus Gray, 1832
Type species: Diplodactylus vittatus Gray, 1832, Australia, Recent.
Diplodactylus sp. cf. Diplodactylus steindachneri Boulenger, 1885
Material.—QM F30573 (right maxilla), QM F30574 (left parietal), and QM F30572 (partial right mandible) from “Wilkinson's Quarry”, Chinchilla Rifle Range, Australia, Pliocene (Hutchinson and Mackness 2002).
Remarks.—Specimens were recovered from synonymous locations “Wilkinson's Quarry” or “Quarry Site” (Hutchinson and Mackness 2002). Those authors suggest that the specimens are the remains from a single individual.
Family Scincidae Gray, 1825
Genus Cyclodomorphus Fitzinger, 1843
Type species: Cyclodomorphus casuarinae (Duméril and Bibron, 1839), Australia, Recent.
cf. Cyclodomorphus sp.
Material.—QM F30568 (partial dentary) from “Wilkinson's Quarry”, Chinchilla Rifle Range, Australia, Pliocene (Hutchinson and Mackness 2002).
Genus Tiliqua Gray, 1825
Type species: Tiliqua scincoides (White, 1790), Australia, Recent.
Tiliqua wilkinsonorum Hutchinson and Mackness, 2002
Material.—QM F30567 (holotype, right dentary) from “Wilkinson's Quarry”, Chinchilla Rifle Range, Australia, Pliocene (Hutchinson and Mackness 2002).
Remarks.—While having been assigned a QM specimen number, the specimen has never been received by the museum.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Family Varanidae Merrem, 1820
Genus Varanus Merrem, 1820
Type species: Varanus varius (White, 1790), Australia, Recent.
Varanus komodoensis Ouwens, 1912
Fig. 4G.
Material.—QM F874 (right maxilla), QM F42105 (partial right maxilla), QM F870 (partial left dentary), QM F871 (partial left dentary), QM F42156 (quadrate), QM F866 (scapulacoracoid), QM F53954 (partial left humerus), and QM F53955 (partial right humerus) from Chinchilla, Australia, Pliocene; QM F25392 supraorbital) from “Wilkinson's Dig, near Graham's Gully”, Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—De Vis (1889b) erected the species Varanus dirus on the basis of a single tooth from King Creek, Darling Downs. He later assigned the first lizard material described from Chinchilla, a partial maxilla (QM F874) to that species (de Vis 1900), a referral that was later confirmed by Feférváry (1918). Hecht (1975), in his review of Megalania, subsumed Varanus dirus into Megalania prisca; however, he could not confidently refer de Vis's maxillary fragment to either Megalania or Varanus komodoensis. He also referred four vertebrae from Chinchilla (QM C20 and C106) to Megalania sp. (Molnar 1982a; Lees 1986). Several additional varanid specimens were assigned to ?Megalania sp. and Varanus sp. by Hutchinson and Mackness (2002). Those specimens were then part of a private collection (WPC); however, ten of the twelve specimens were later donated to the QM and have been assigned registration numbers (SOM: Table 1). Of those, four are clearly not varanid, (SOM: Table 1) and this is probably due to either numbers being incorrectly quoted or the result of typographical errors (see comments for Agamidae gen. et sp. indet.). The six specimens that are varanid (QM F56191–F56196) are comparable with Varanus komodoensis and are here assigned to that species. Hocknull et al. (2009) referred several QM specimens from Chinchilla, including de Vis's (1900) maxillary fragment and several previously unidentified specimens, to Varanus komodoensis.
Stratigraphic and geographic range.—Pliocene to Recent; eastern Australia (fossil only) and Indonesia.
Varanus sp.
Fig. 4D.
Material.—QM F56189 (dorsal vertebra) and QM F56190 (dorsal vertebra) from “Wilkinson's Quarry”, Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Both specimens were referred to Varanus sp. by Hutchinson and Mackness (2002) under WPC numbers (see SOM: Table 1).
Family Madtsoiidae Hoffstetter, 1961
Genus Yurlunggur Scanlon, 1992
Type species: Yurlunggur camfieldensis Scanlon, 1992, north central Northern Territory, middle Miocene.
Yurlunggur sp.
Material.—QM F30560 (cast of a vertebra) from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Mackness and Scanlon (1999) referred the first snake fossil described from the Chinchilla Sand to Yurlunggur sp. While assigned a QM number, the specimen has never been received by the museum.
Class Aves Linneaus, 1758
Order Casuariiformes Sclater, 1880
Family Casuariidae Kaup, 1847
Genus Dromaius Vieillot, 1816
Type species: Dromaius novaehollandiae (Latham, 1790), Syndey, New South Wales, Recent.
Dromaius novaehollandiae (Latham, 1790)
Fig. 5K.
Material.—QM F56203 (third trochlea of the tarsometatarsus) likely from Chinchilla, Australia, Pliocene.
Remarks.—De Vis (1892: 446) described several specimens (QM F1121, QM F56202 [originally QM F1143, in part], QM F56204 [originally QM F1143, in part] and a “calcaneal region of another metatarse” [specimen missing] which he referred to Dromaius patricius. De Vis (1892) also erected a new species of emu, D. gracileps (QM F1142) and kiwi, Metapteryx bifrons (QM F1135). All three species were synonymised with the modern emu D. novaehollandiae by Patterson and Rich (1987). These authors described an additional specimen, also originally numbered QM F1143—a third trochlea of the tarsometatarsus (QM F56203)—and assigned it to D. novaehollandiae. Confusingly, three D. novaehollandiae specimens were registered, in addition to a specimen referred to Leipoa (Progura) gallinacea, as QM F1143. In order to avoid further confusion, all the D. novaehollandiae specimens have now been assigned new, individual QM F numbers (QM F56202–F56204).
De Vis (1892) did not give a location for any of these specimens, but it is likely that all his specimens came from Chinchilla, following Olson's (1975) inference regarding the origin of the specimens described by de Vis (1888c) (see Remarks for Gallinula morterii). It is unclear why Patterson and Rich (1987) only identified one of the original QM F1143 specimens (the third trochlea of the tarsometatarsus, QM F56203) as coming from Chinchilla, the other two being ascribed to the eastern Darling Downs. That designation is reflected on both specimen labels, and yet the QM registry book records all four specimens originally registered as QM F1143 as coming from the “Darling Downs”. We accept a “Darling Downs” provenance (inclusive of both the Pleistocene eastern Darling Downs and the Pliocene western Darling Downs), although we consider it likely that all four specimens (three D. novaehollandiae and one L. gallinacea) originally registered as QM F1143, are from Chinchilla.
Finally, it does not appear that de Vis (1892) mentioned the third trochlea of the tarsometatarsus (QM F56203) described by Patterson and Rich (1987); however, de Vis probably recorded this specimen as a “calcaneal region of another metatarse” (de Vis 1892: 446). If correct, this would: (i) explain why all three QM F1143 emu specimens were given the same registration number; (ii) account for the missing “calcaneal region” specimen; and (iii) resolve the disparity in locations recorded for the different specimens.
Stratigraphic and geographic range.—Pliocene to Pleistocene; Australia.
Order Galliformes Temmink, 1820
Family Megapodiidae Lesson, 1831
Genus Leipoa Gould, 1840
Type species: Leipoa ocellata Gould, 1840, Australia, Recent.
Leipoa gallinacea (de Vis, 1888)
Fig. 5B.
Material.—QM F1132 (partial carpometacarpus) and QM F5558 (partial scapula) from Chinchilla, Australia, Pliocene.
Remarks.—These two Chinchilla specimens, one assigned to Chosornis praeteritus by de Vis (1889a) and the other to Otididae sp. indet. by de Vis (1888c) were subsequently referred to Progura gallinancea by van Tets (1974). Progura was subsumed into Leipoa by Boles (2008).
Stratigraphic and geographic range.—Pliocene to Pleistocene; eastern Australia.
Order Anseriformes (Wagler, 1831)
Family Anatidae (Leach, 1819)
Genus Biziura Stephens, 1824
Type species: Biziura lobata Shaw, 1796, Australia, Recent.
Biziura lobata Shaw, 1796
Fig. 5C.
Material.—QM F1125 (partial left humerus), QM F1133 (left tarsometatarsus), and QM F7057 (left femur) from Chinchilla, Australia, Pliocene.
Remarks.—Olson (1977) referred species formerly assigned to Dendrocygna validipinnis and Biziura exhumata by de Vis (1888c) to the extant Musk duck, Biziura lobata. Olson (1977) noted that one specimen (QM F5551), supposedly from Chinchilla, had a different preservation, and might come from the late Pleistocene Darling Downs deposits.
Stratigraphic and geographic range.—Pliocene to Recent; Australia.
Genus Anas Linneaus, 1758
Type species: Anas platyrhynchos Linneaus, 1758, Europe, Recent.
Anas superciliosa Gmelin, 1789
Fig. 5D.
Material.—QM F1122 (partial right humerus) and QM F5550 (left coracoid) from Chinchilla, Australia, Pliocene.
Remarks.—De Vis (1888c) described a new species of duck, Nyroca robusta, from Chinchilla. This species was considered by Olson (1977) to be indistinguishable from the living Pacific black duck, Anas superciliosa.
Stratigraphic and geographic range.—Pliocene to Recent, Australia, New Guinea, New Zealand, Indonesia, Pacific.
Genus Aythya Boie, 1822
Type species: Aythya marila (Linneaus, 1761), Europe, Recent.
Aythya australis Eyton, 1838
Fig. 5A.
Material.—QM F1123 (left coracoid) and QM F1124 (left tibiotarsus) from Chinchilla, Australia, Pliocene.
Remarks.—Olson (1977), continuing his review of de Vis's (1888c) ducks, referred two other Chinchilla species (Nyroca recluse and Anas elapsa) to the extant White-eyed duck Aythya australis.
Stratigraphic and geographic range.—Pliocene to Recent; Australia.
Order Phalacrocoraciformes Christidis and Boles, 2008
Family Phalacrocoracidae Reichenbach, 1850
Genus Microcarbo Bonaparte, 1856
Type species: Microcarbo pygmaeus (Pallas, 1773), Europe, Recent.
Microcarbo melanoleucos Vieillot, 1817
Fig. 5I.
Material.—QM F1130 (right humerus) from Chinchilla, Australia, Pliocene.
Remarks.—De Vis (1888c) described a species of Anhingidae on the basis of a specimen from Chinchilla which he called Plotus parvus. Miller (1966a) revised that diagnosis and assigned the specimen to Halietor (now Microcarbo) melanoleucos, synonymising Plotus parvus with the extant species.
Stratigraphic and geographic range.—Pliocene to Recent; Australia, Indonesia, New Guinea, New Zealand, Pacific.
Order Pelecaniformes Sharpe, 1891
Family Pelecanidae Rafinesque, 1815
Genus Pelecanus Linneaus, 1758
Type species: Pelecanus onocrotalus Linneaus, 1758, Europe, Recent.
Pelecanus proavus de Vis, 1892
Material.—Unknown missing specimen (described as Pelicanus [sic!] proavus and illustrated in de Vis (1892: pl. 24: 6a, b), probably from Chinchilla, Australia, Pliocene.
Remarks.—De Vis (1892) described a new species of pelican on the basis of two specimens possibly from Chinchilla (see remarks in Gallinula morterii). Miller (1966b) reviewed the one remaining specimen available for study (QM F1141), and concluded that it was not referable to Pelecanidae. He compared de Vis's (1892) illustration of the other specimen, the type of Pelecanus proavus, with the modern pelican and could find no characters to distinguish them. However, Rich and van Tets (1981) disagreed with that assessment due to the quality of de Vis's (1892) illustrations, and reserved judgement on P. proavus until the original specimen is located, a view we support.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Order Ciconiiformes Bonaparte, 1854
Family Ciconiidae Sundevall, 1836
Genus Ciconia Brisson, 1760
Type species: Ciconia ciconia (Linneaus, 1758), Europe, Recent.
Ciconia nana (de Vis, 1888)
Fig. 5J.
Material.—QM F1131 (lectotype, distal right tibiotarsus) and QM F5514 (paralectotype, proximal right ulna) from the north bank of the Condamine River, 5 km from the town of Chinchilla, Australia, Pliocene.
Remarks.—De Vis's (1888c) description of a stork from Chinchilla (Xenorhynchus nanus) was reviewed by Boles (2005), who agreed with the specific diagnosis, but assigned the species to the extant genus Ciconia. Boles (2005) erected the appropriate lectotypes on the basis of Chinchilla material.
Stratigraphic and geographic range.—Pliocene to Pleistocene; central and eastern Australia.
Order Falconiformes Sharpe, 1874
Family Accipitridae Vigors, 1824
Genus Necrastur de Vis, 1892
Type species: Necrastur alacer de Vis, 1892, type locality unknown but probably Chinchilla, Pliocene.
Necrastur alacer de Vis, 1892
Fig. 5H.
Material.—QM F1136 (proximal right humerus) and QM F5552 (distal right ulna), probably from Chinchilla, Australia, Pliocene.
Remarks.—The species was described de Vis (1892), and is therefore of possible Chinchilla provenance (see remarks in Gallinula morterii). These two specimens were listed by Rich and van Tets (1982), but as ?Harpyopsis and Butonidae. They stated that a revision of the taxon was in preparation at the time, presumably by them. We were unable to locate the material, and until a revision is published, we follow the taxonomy as currently published.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Fig. 5.
Birds from unspecified localities near Chinchilla, Australia, Pliocene. A. Aythya australis Eyton, 1838, QM F1124, left tibiotarsus. B. Leipoa gallinacean (de Vis, 1888), QM F1132, partial carpometacarpus. C. Biziura lobata Shaw, 1796, QM F1125, partial left humerus. D. Anas superciliosa Gmelin, 1789, QM F5550, left coracoid. E. Fulica atra Linneaus, 1758, QM F1129, proximal right humerus. F. Gallinula morterii du Bus, 1840, QM F1138, distal right humerus. G. Charadriiformes gen. et sp. indet., QM F5543, proximal left femur. H. Necrastur alecer de Vis, 1892, QM F1136, proximal right humerus. I. Microcarbo melanoleucos Vieillot, 1817, QM F1130, right humerus. J. Ciconia nana (de Vis, 1888), QM F1131, distal right tibiotarsus. K. Dromaius novaehollandiae (Latham, 1790), QM F56203, third trochlea of the tarsometatarsus. Scale bars 10 mm.
Order Gruiformes Bonaparte, 1854
Family Rallidae Rafinesque, 1815
Genus Fulica Linneaus, 1758
Type species: Fulica atra Linneaus, 1758, Europe, Recent.
Fulica atra Linneaus, 1758
Fig. 5E.
Material.—QM F1129 (proximal right humerus) from Chinchilla, Australia, Pliocene.
Remarks.—Olson (1975) synonymised Fulica prior, originally described by de Vis (1888c), with Fulica atra.
Stratigraphic and geographic range.—Pliocene to Recent; Europe, Asia, Australia, Africa.
Genus Gallinula Brisson, 1760
Type species: Gallinula chloropus (Linnaeus, 1758), Europe, Recent.
Gallinula mortierii du Bus, 1840
Fig. 5F.
Material.—QM F1138 (distal right humerus), QM F1144 (right humerus), QM F1128 (left tarsometatarsus), QM F5554 (tibia), QM F5555 (tibia), QM F1126 (distal right tarsometatarsus), QM F7008 (femur), QM F7007 (femur), QM F7009 (tibia), QM F7029 (proximal tibiotarsus), QM F7030 (distal tibiotarsus), and QM F7058 (partial humerus), all probably from Chinchilla, Australia, Pliocene.
Remarks.—Olson (1975) synonymised four species described by de Vis (1888c), namely Gallinula strenuipes, Gallinula peralata, Tribonyx effluxus, and Porphyrio mackintoshi into Gallinula (Tribonyx) mortierii. Olson (1975) also drew attention to the fact that de Vis's (1892) account of fossil birds lacked an introduction, and inferred that it was meant as a continuation of de Vis (1888c). In the introduction, de Vis (1888c: 1277) refers to specimens “yielded by the Darling Downs in the immediate neighbourhood of Chinchilla”. It is therefore probable that all the specimens mentioned by de Vis (1892) came from the Chinchilla Sand. QM F7029 and F7030 represent two parts (proximal and distal tibiotarsus, respectively) of the same specimen.
Stratigraphic and geographic range.—Pliocene to Recent; Australia.
Order Charadriiformes Huxley, 1867
Charadriiformes gen. et sp. indet.
Fig. 5G.
Material.—QM F5543 (proximal left femur) from Chinchilla, Australia, Pliocene.
Remarks.—The specimen referred to Anas elapsa by de Vis (1888c) was identified by Olson (1977) as Charadriiformes indeterminate.
Family Charadriidae Leach, 1820
Genus Vanellus Brisson, 1760
Type species: Vanellus vanellus (Linnaeus, 1758), Europe, Recent.
Vanellus sp.
Material.—None (destroyed).
Remarks.—De Vis (1892) described a species of lapwing (Lobivanellus sp., now Vanellus sp.), but stated in his description that the specimen was destroyed before a full taxonomic assignment could be made. As this specimen was published in de Vis's 1892 treatise, it is possible that it was derived from the Chinchilla Sand (see remarks in Gallinula morterii).
Class Mammalia Linneaus, 1758
Infraclass Marsupialia Illiger, 1811
Order indeterminate
Genus Archizonurus de Vis, 1889
Type species: Archizonurus securus de Vis, 1889, Chinchilla, Pliocene.
Archizonurus securus de Vis, 1889
Fig. 6C.
Material.—QM F682 (holotype, proximal left scapula fragment) from Chinchilla, Australia, Pliocene.
Remarks.—Meston (1895) suggested Archizonurus securus could be a possum, whereas Mahoney and Ride (1975) regard this species as Marsupialia incertae sedis. The specimen is clearly from a taxon much larger than a possum (e.g., large-bodied macropodid or wombat-sized individual). We follow the latest taxonomic authority on the species, but nevertheless consider Archizonurus securus as a species inquirenda.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Order Dasyuromorphia Gill, 1872
Family Dasyuridae Goldfuss, 1820
Genus Dasyurus Geoffroy Sainte-Hilaire, 1796
Type species: Dasyurus maculatus (Kerr, 1792), type locality not given, Recent.
Dasyurus dunmalli Bartholomai, 1971
Fig. 7A.
Material.—QM F6579 (holotype, left mandibular ramus), QM F6580 (left mandibular ramus), and QM F12682 (dentary fragment) from Chinchilla Rifle Range, Australia, Pliocene; QM F30477 (dentary fragment) from Chinchilla Rifle Range, Pliocene; QM F57539 (= “W3357″ and “QM F3357″ in Wroe and Mackness 1998, and Wroe and Mackness 2000a, respectively; dentary fragment) from north bank of the Condamine River, Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—A species of extinct quoll from Chinchilla, Dasyurus dunmalli, was described by Bartholomai (1971). An additional specimen (lower right mandibular fragment) from Chinchilla was later referred to D. dunmalli by Wroe and Mackness (2000a), who listed the specimen as QM F3357. However, QM F3357 is actually a specimen of the diprotodontid, Euryzygoma dunense, as registered in the QM catalogue. Confusingly, Wroe and Mackness (1998) had earlier noted the same specimen as “W3357″, so part of the Wilkinson Private Collection (i.e., WPC 3357), not the Queensland Museum (i.e., QM F), thus providing an explanation for the inconsistency in the QM registration book. Even more confusingly, the WPC registration book lists the number “3357″ as a vertebra, not a jaw, and certainly not D. dunmalli. However, the transfer of the number “3357″ to the D. dunmalli specimen is an error, for the WPC registration book lists specimen “3356″ as a jaw fragment of D. dunmalli. The description of WPC 3356 in the registration book appears to be consistent with the figured specimen (i.e., “QM F3357″) featured in Wroe and Mackness (2000a). To avoid confusion, the specimen is renumbered as QM F57539.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Genus Archerium Wroe and Mackness, 2000
Type species: Archerium chinchillaensis Wroe and Mackness, 2000, Chinchilla, Pliocene.
Archerium chinchillaensis Wroe and Mackness, 2000
Fig. 7E.
Material.—QM F39847 (holotype, left maxillary fragment) from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—A new genus and species of dasyurid, Archerium chinchillaensis, was erected by Wroe and Mackness (2000b) on the basis of Chinchilla material.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Family Thylacinidae Bonaparte, 1838
Genus Thylacinus Temmink, 1824
Type species: Thylacinus cynocephalus (Harris, 1808), Tasmania, Recent.
Thylacinus cynocephalus (Harris, 1808)
Fig. 7K.
Material.—QM F3741 (right maxillary fragment) and QM F9476 (right mandibular fragment) from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—De Vis (1894a) described two specimens of thylacine from Chinchilla (namely Cat. No. 12,029 = QM F726 in part, and Cat. No. 12,031 = QM F3744) which he attributed to a new species, Thylacinus rostralis. Ride (1964) referred an additional specimen (QM F2159) to that species. Thylacinus rostralis was later considered a junior synonym of the modern species T. cynocephalus by Dawson (1982). Mackness et al. (2002) reviewed the history and occurrence of Thylacinus at Chinchilla, and concluded that QM F726 and Cat. No. 12,026 were not from Chinchilla, but rather from the Darling Downs, on the basis of label information and preservation. However, they did not realise that Cat. No. 12,026 was also registered as QM F726, and that Cat. No. 12,026 and 12,029 are very likely part of the same specimen. It is worth citing the complete label information attached with QM F726: “Only the anterior portion of this specimen was listed by de Vis (1894[a]) as being in the Museum collections. He claimed that it came from the ?Pliocene Chinchilla Sand at Chinchilla, N.W. Darling Downs. It bears the de Vis number 12029. Preservation indicates it has been derived from the Pleistocene fluviatile deposits [of the eastern Darling Downs]. The posterior portion of the ramus was associated during the early period of Longman's time and I think there will be no hesitation about accepting the association. This portion bears the de Vis number 12026 but was not listed in de Vis' (1894[a]) paper. The numbering indicates that he most certainly had the specimen. Longman noted that the posterior portion was from Pilton, S.E. Darling Downs and registered the specimen as such. Remains of the purchase number are still visible on the posterior portion, and this locality represents a much more likely source for the ramus” (Alan Bartholomai, undated). Mackness et al. (2002) did not discuss the other specimen described by de Vis (QM F3744); its label information indicates that it was recovered from the ?Pliocene Chinchilla Sand. They also suggest that the specimen described by Ride (1964) was not derived from the Chinchilla Sand on the basis of preservation. Mackness et al. (2002) described material from Chinchilla (registered WPC), but suggested that it was only attributable to Thylacinus sp. However, two additional specimens in the QM collection, recovered in 1962 and 1977, and identified as T. rostralis (= T. cynocephalus) are definitely from Chinchilla.
Stratigraphic and geographic range.— Pliocene-Holocene; mainland Australia and New Guinea (fossil only), and Tasmania.
Fig. 6.
Mammal species of uncertain taxonomic identity from unspecified localities near Chinchilla, Australia, Pliocene. A. Koalemus ingens de Vis, 1889, QM F683, partial right fibula. B. Synaptodon aevorum de Vis, 1888, QM F811, right dentary fragment. C. Archizonurus securus de Vis, 1889, QM F682, proximal left scapula. D. Phalanger procuscus (de Vis, 1889), QM F687, right scapula. E. Chronozoon australe de Vis, 1883, QM F610, calvarium. F. Brachalletes palmeri de Vis, 1883, QM F3308, right femur. Scale bars 10 mm.
Order Peramelemorphia Kirsch, 1968
Family Peramelidae Gray, 1825
Genus Perameles Geoffroy, 1804
Type species: Perameles nasuta Geoffroy, 1804, New South Wales, Recent.
Perameles bowensis Murihead, Dawson, and Archer, 1997
Material.—QM F30580 (left M1) and QM F30581 (left dentary), both from “Wilkinson's Quarry”, Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Mackness et al. (2000b) described the only known bandicoot from Chinchilla as Perameles bowensis, and referred two specimens to the species, QM F30580 and QM F30581. QM F30580 could not be located, and QM F30581 is a specimen of a snake, Liasis. Both registration numbers have also inadvertently been assigned to two fossil bat specimens, Icarops sp. cf. I. paradox and I. paradox, respectively, from Miocene deposits of Riversleigh (Hand et al. 2005). An erratum is currently being prepared by Sue Hand (personal communication 2015) to correct the registration discrepancy.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Order Diprotodontia Owen, 1866
Suborder Vombatiformes Woodburne, 1984
Family indeterminate
Genus Koalemus de Vis, 1889
Type species: Koalemus ingens de Vis, 1889, unknown location, but likely Chinchilla, Pliocene.
Koalemus ingens de Vis, 1889
Fig. 6A.
Material.—QM F683 (partial right fibula), locality uncertain, ?Pliocene.
Remarks.—Bartholomai (1968) considered the specimen to be from Chinchilla on the basis of its preservation. Although de Vis (1889c) suggested affinities of the species with the modern koala, Bartholomai (1968) suggested it be referred to Diprotodontidae incertae sedis. We consider Koalemus ingens a as species inquirenda.
Stratigraphic and geographic range.—Uncertain, possibly Pliocene, eastern Australia.
Family Phascolarctidae Owen, 1839
Phascolarctidae gen. et sp. indet.
Fig. 7C.
Material.—QM F52287 (left dentary fragment) from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—This large dentary fragment appears to be morphologically distinct from all other known koala dentaries (Price et al. 2009). However the lack of diagnostic characters on the specimen means it can only be referred to Phascolarctidae gen. et sp. indet.
Genus Koobor Archer, 1977
Type species: Koobor notabilis (de Vis, 1889), unknown location, but likely Chinchilla, Pliocene.
Koobor notabilis (de Vis, 1889)
Fig. 7B.
Material.—QM F691 (holotype, left maxillary fragment) probably from Chinchilla, Australia, Pliocene; QM F8976 (P3) from between “Col's Knob” and “Liz's Hill”, Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—De Vis (1889c) described a maxillary fragment, probably from Chinchilla (see discussion in Archer 1977), as Pseudochirus ?notabilis. Archer (1977) re-assigned it to Koobor, and referred another specimen (QM F8976) of definite Chinchilla provenance to the species. Koobor was assigned to Phascolarctidae by Black et al. (2012a).
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Genus Phascolarctos Blainville, 1816
Type species: Phascolarctos cinereus Blainville, 1816, New South Wales, Recent.
Phascolarctos ?stirtoni Bartholomai, 1968
Fig. 7D.
Material.—QM F52289 (isolated RM1, RM2 or RM3 fragment) from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Assigned to Phascolarctos ?stirtoni by Price (2008a) and Price et al. (2009).
Family Diprotodontidae Gill, 1872
Genus Euowenia de Vis, 1891
Type species: Euowenia grata (de Vis, 1887), Chinchilla, Pliocene.
Euowenia grata (de Vis, 1887)
Fig. 7J.
Material.—QM F519 (holotype, mandible) from north bank of Condamine River, Chinchilla, Australia, Pliocene.
Remarks.—Euowenia was first named Owenia, in honour of Sir Richard Owen, by de Vis (1887, 1888a), but the name was preoccupied, and de Vis (1891a) changed it to Euowenia.
Stratigraphic and geographic range.—Pliocene-?Pleistocene; central and eastern Australia.
Fig. 7.
Non-macropodid marsupials from Chinchilla Sand, Australia, Pliocene. A. Dasyurus dunmalli Bartholomai, 1971, QM F6579, Chinchilla Rifle Range, left mandibular ramus. B. Koobor notabilis (de Vis, 1889), QM F691, unspecified locality near Chinchilla, left maxillary fragment. C. Phascolarctidae gen. et sp. indet., QM F52287, Chinchilla Rifle Range, left dentary fragment. D. Phascolarctos ?stirtoni Bartholomai, 1968, QM F52289, Chinchilla Rifle Range, isolated RM1, RM2 or RM3 fragment. E. Archerium chinchillaensis Wroe and Mackness, 2000, QM F39847, Chinchilla Rifle Range, left maxillary fragment. F. Thylacoleo crassidentatus Bartholomai, 1962, QM F3565, right mandibular ramus. G. Euryzygoma dunense (de Vis, 1888), QM F376, unspecified locality near Chinchilla, left mandible. H. Vombatus ursinus (Shaw, 1800), QM F743, unspecified locality near Chinchilla, proximal right tibia. I. Palorchestes parvus de Vis, 1895, QM F783, unspecified locality near Chinchilla, left mandibular fragment. J. Euowenia grata (de Vis, 1887), QM F519, north bank of Condamine River, mandible. K. Thylacinus cynocephalus (Harris, 1808), QM F9476, Chinchilla Rifle Range, right mandibular fragment.
Genus Euryzygoma Longman, 1921
Type species: Euryzygoma dunense (de Vis, 1888), Chinchilla, Pliocene
Euryzygoma dunense (de Vis, 1888)
Fig. 7G.
Material.—QM F376 (holotype, left mandible) from Chinchilla, Australia, Pliocene.
Remarks.—De Vis (1888b) described Euryzygoma dunense as a species of Nototherium. Longman (1921), who had access to more complete cranial material, assigned the species to its own genus, Euryzygoma.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Genus Zygomaturus Macleay, 1857
Type species: Zygomaturus trilobus Macleay, 1858, King Creek, Darling Downs, Queensland, Pliestocene.
Zygomaturus sp.
Material.—QM F3829 (upper premolar) from Chinchilla, Australia, Pliocene.
Remarks—Archer and Wade (1976) discussed a Zygomaturus specimen from the Chinchilla Sand, drawing similarities between it and an isolated premolar from the early Pliocene Allingham Formation. The specimen could not be located in the QM collection.
Family Palorchestidae Tate, 1948
Genus Palorchestes Owen, 1873
Type species: Palorchestes azael Owen, 1873, Victoria, Pleistocene.
Palorchestes parvus de Vis, 1895
Fig. 7I.
Material.—QM F783 (lectotype, left mandibular fragment) from Chinchilla, Australia, Pliocene.
Remarks.—Palorchestes parvus was named by de Vis (1895) on the basis of material from the “Darling Downs”. In his review of Palorchestes, Woods (1958) suggested that the species was derived solely from specimens from Chinchilla and designated a lectotype for the species. However, it is noted that small-bodied species of Palorchestes potentially referable to P. parvus do occur in younger, Pleistocene deposits of the Darling Downs (Price and Hocknull 2005). Rich et al. (1991) also listed cf. Palorchestes sp. from Chinchilla, but we could find no further reference to that taxon.
Stratigraphic and geographic range.—Pliocene to ?Pleistocene; eastern Australia.
Family Vombatidae Burnett, 1830
Genus Vombatus Geoffroy, 1803
Type species: Vombatus ursinus (Shaw, 1800), Tasmania, Recent.
Vombatus ursinus (Shaw, 1800)
Fig. 7H.
Material.—QM F743 (proximal right tibia) from Chinchilla, Australia, Pliocene.
Remarks.—De Vis (1883c) erected Sarcophilus prior (marsupial devil) on the basis of a tibia fragment. Bartholomai and Marshall (1973) referred the specimen to ?Vombatus prior (wombat) noting marked similarities with the common wombat, V. ursinus, although they did not assign it to that species due to the paucity of fossil Vombatus material. Murray (1998) regarded ?Vombatus prior as a nomen dubium, and listed it as a junior synonym of V. ursinus. The Vombatus material from Chinchilla has since been revised (Louys 2015; see note added in proofs below).
Stratigraphic and geographic range.—Pliocene to Holocene; eastern and southeastern mainland Australia, and Tasmania (extant).
Family Thylacoleonidae Gill, 1872
Genus Thylacoleo Gervais, 1852
Type species: Thylacoleo carnifex Owen, 1858, Lake Colongulac, Victoria, Pleistocene.
Thylacoleo crassidentatus Bartholomai, 1962
Fig. 7F.
Material.—QM F3565 (holotype, right mandibular ramus) from Chinchilla Rifle Range, Australia, Pliocene; QM F2957 (mandibular ramus), QM F2961 (mandibular ramus), QM F2962 (mandibular ramus), QM F2960 (mandibular ramus), QM F2964 (mandibular ramus), QM F2963 (p3), QM F2495 (mandibular ramus), QM F2941 (partial maxilla), QM F2954 (partial maxilla), and QM F2955 (partial maxilla) from Chinchilla, Australia, Pliocene.
Remarks.—In his revision of Thylacoleo in the QM collections, Woods (1956) suggested that specimens from Chinchilla warranted specific designation. This was supported by Bartholomai (1962), who erected T. crassidentatus for the Chinchilla material.
Stratigraphic and geographic range.— Pliocene; eastern Australia.
Suborder Phalangerida Aplin and Archer, 1987
Family Phalangeridae Thomas, 1888
Genus Phalanger Storr, 1780
Type species: Phalanger orientalis (Pallas, 1766), Ambon Island, Indonesia, Recent.
Phalanger procuscus (de Vis, 1889)
Fig. 6D.
Material.—QM F687 (holotype, right scapula) from Chinchilla, Australia, Pliocene.
Remarks.—De Vis (1889c) originally named the specimen Cuscus procuscus. Referral to Phalanger here is based on the priority of the name. Mahoney and Ride (1975) referred this species to Marsupialia incertae sedis. Similar to Archizonurus securus, the specimen is clearly from a large-bodied taxon, much larger than any known phalangerid. We consider Phalanger procuscus as a species inquirenda.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Family Macropodidae Gray, 1821
Genus Brachalletes de Vis, 1883
Type species: Brachalletes palmeri de Vis, 1883, Chinchilla, Pliocene.
Brachalletes palmeri de Vis, 1883
Fig. 6F.
Material.—QM F3308 (holotype, right femur) from Chinchilla, Australia, Pliocene.
Remarks.—Described by de Vis (1883b), the taxon was included in the family Macropodidae by Mahoney and Ride (1975), although they also suggested that it might be better placed in the Diprotodontidae. Dawson and Flannery (1985) suggested it possessed no characters diagnostic of the Macropodinae (long-faced kangaroos), and its placement remains uncertain, if indeed it is a valid taxon. We consider Brachalletes palmeri as a species inquirenda.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Genus Synaptodon de Vis, 1888
Type species: Synaptodon aevorum de Vis, 1888, Chinchilla, Pliocene.
Synaptodon aevorum de Vis, 1888
Fig. 6B.
Material.—QM F811 (holotype, right dentary fragment) from Chinchilla, Australia, Pliocene.
Remarks.—Both Bartholomai (1975) and Dawson and Flannery (1985) argued that the taxon is based on inadequate, poorly preserved and undiagnostic material. Dawson and Flannery (1985) suggested that it bears no characters that allow it to be referred to Macropus. We consider Synaptodon aevorum as a nomen dubium.
Subfamily Lagostrophinae Prideaux and Warburton, 2010
Genus Troposodon Bartholomai, 1967
Type species: Troposodon minor (Owen, 1877), Talbragar country, New South Wales, ?Pleistocene.
Fig. 8.
Macropodids from Chinchilla Sand, Australia, Pliocene. A. Simosthenurus antiquus (Bartholomai, 1963), QM F2975, partial left maxilla of probable Chinchilla provenance. B. Sthenurus andersoni Marcus, 1962, QM F814, unspecified locality near Chinchilla, left juvenile dentary. C. Macropus agilis siva (de Vis, 1895), QM F4733, unspecified locality near Chinchilla, right mandibular fragment. D. Sthenurus notabilis Bartholomai, 1963, QM F3817, Chinchilla Rifle Range, right mandibular ramus. E. Troposodon gurar Flannery and Archer, 1983, QM F4609, unspecified locality near Chinchilla, right dentary of probable Chinchilla provenance. F. Protemnodon devisi Bartholomai, 1973, QM F4710, unspecified locality near Chinchilla, partial left mandibular ramus. G. Macropus dryas de Vis, 1895, QM F3582, partial right maxilla of probable Chinchilla provenance. H. Protemnodon chinchillaensis Bartholomai, 1973, QM F5246, unspecified locality near Chinchilla, partial right mandibular ramus. I. Bohra wilkinsonorum Dawson, 2004, QM F43277, Chinchilla Rifle Range, right maxillary fragment. J. Wallabia indra (de Vis, 1895), QM F3595, unspecified locality near Chinchilla, left mandibular ramus. K. Troposodon minor (Owen, 1877), QM F4389, Condamine River, “50 yards east of Chinchilla Rifle Range”, right maxillary fragment. L. Prionotemnus palankarinnicus Stirton, 1955, QM F3589, partial right mandibular ramus of probable Chinchilla provenance. M. Silvaroo bila Dawson, 2004, QM F43276, Chinchilla Rifle Range, left maxillary fragment. N. Silvaroo sp., QM F43281, Chinchilla Rifle Range, right mandibular ramus. O. Macropus woodsi Bartholomai, 1975, QM F5465, Chinchilla Rifle Range, partial left maxilla. P. Macropus pan de Vis, 1895, QM F2925, partial right maxilla of probable Chinchilla provenance. Scale bars 10 mm.
Troposodon minor (Owen, 1877)
Fig. 8K.
Material.—QM F4389 (right maxillary fragment) from Condamine River, “50 yards east of Chinchilla Rifle Range, 30′ above low water”, Australia, Pliocene.
Remarks.—Bartholomai (1967) erected the genus Troposodon and assigned many Chinchilla specimens to the Troposodon minor.
Stratigraphic and geographic range.—Pliocene to Pleistocene; eastern Australia.
Troposodon gurar Flannery and Archer, 1983
Fig. 8E.
Material.—QM F4609 (holotype, right dentary) most likely from Chinchilla, Australia, Pliocene.
Remarks.—Flannery and Archer (1983) described this new species on the basis of material of probable Chinchilla provenance.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Subfamily Sthenurinae (Glauert, 1926)
Genus Sthenurus Owen, 1873
Type species: Sthenurus atlas (Owen, 1838), Wellington Valley, late Pleistocene.
Sthenurus notabilis Bartholomai, 1963
Fig. 8D.
Material.—QM F3817 (holotype, right mandibular ramus) from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Bartholomai (1963) erected this species on the basis of Chinchilla material.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Sthenurus andersoni Marcus, 1962
Fig. 8B.
Material.—QM F813 (right adult dentary), QM F814 (left juvenile dentary), UCR 23029 (right lower molar), and UCR 23033 (left incisor) from Chinchilla, Australia, Pliocene.
Remarks.—Prideaux (2004) assigned several specimens from Chinchilla to this species.
Stratigraphic and geographic range.—Pliocene to Pleistocene; central to southeastern mainland Australia.
Genus Simosthenurus (Tedford, 1966)
Type species: Simosthenurus occidentalus (Glauret, 1910), Mammoth Cave, Western Australia, late Pleistocene.
Simosthenurus antiquus (Bartholomai, 1963)
Fig. 8A.
Material.—QM F2975 (holotype, partial left maxilla) probably from Chinchilla, Australia, Pliocene; QM F3818 (right p3) from Chinchilla Rifle Range, Australia, Pliocene; QM F3816 (partial left M2) from Middle Gully of Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Bartholomai (1963) erected this species with a holotype (QM F2975) probably from Chinchilla. The species was subsequently assigned to the new genus Simosthenurus by Pledge (1980) and later “Simosthenurus” by Prideaux (2004). Other specimens (e.g., those listed above) are of definite Chinchilla provenance.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Subfamily Macropodinae Gray, 1821
Genus Wallabia Trouessart, 1905
Type species: Wallabia bicolor (Desmarest, 1804), type locality unknown, Recent.
Wallabia indra (de Vis, 1895)
ig. 8J.
Material.—QM F3595 (holotype, left mandibular ramus) from Chinchilla, Australia, Pliocene.
Remarks.—Bartholomai (1976) reassigned material previously attributed to Halmaturus indra by de Vis (1895) to Wallabia indra.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Genus Macropus Shaw, 1790
Type species: Macropus giganteus Shaw, 1790, Cooktown, Queensland, Recent.
Macropus pan de Vis, 1895
Fig. 8P.
Material.—QM F2925 (holotype, partial right maxilla) probably from Chinchilla, Australia, Pliocene.
Remarks.—Bartholomai (1975) referred several specimens from Chinchilla to M. pan. Although the provenance of the material had not been noted before, Bartholomai (1975) suggested that the type specimen exhibits preservation consistent with other undoubted Chinchilla specimens.
Stratigraphic and geographic range.—Pliocene; eastern and northwestern Australia.
Macropus agilis siva (de Vis, 1895)
Fig. 8C.
Material.—QM F4733 (right mandibular fragment) and QM F4735 (left mandibular fragment) from Chinchilla, Australia, Pliocene.
Remarks.—Bartholomai (1975) examined material from Chinchilla and attributed it to M. agilis siva, noting that the variation in the Chinchilla material was insufficient to warrant specific distinction.
Stratigraphic and geographic range.—Pliocene to Pleistocene; central, eastern and southeastern Australia.
Macropus dryas de Vis, 1895
Fig. 8G.
Material.—QM F3582 (lectotype, partial right maxilla) from Chinchilla, Australia, Pliocene.
Remarks.—Bartholomai (1966) selected this lectotype on the basis of material of probable Chinchilla provenance, and subsequently (Bartholomai 1975) assigned several additional Chinchilla specimens to the species.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Macropus woodsi Bartholomai, 1975
Fig. 8O.
Material.—QM F3920 (holotype, partial right mandibular ramus) from Middle Gully of Chinchilla Rifle Range, Aus tralia, Pliocene; QM F5465 (partial left maxilla) from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Bartholomai (1975) erected this species on the basis of Chinchilla material, and referred several Chinchilla specimens to the species.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Genus Prionotemnus Stirton, 1955
Type species: Prionotemnus palankarinnicus Stirton, 1955, Lake Palankarinna, South Australia, early Pliocene.
Prionotemnus palankarinnicus Stirton, 1955
Fig. 8L.
Material.—QM F3589 (partial right mandibular ramus) most likely from Chinchilla, Australia, Pliocene.
Remarks.—Bartholomai (1975) assigned two specimens to Macropus (Prionotemnus) palankarinnicus, but only one (QM F6989) is of definite Chinchilla provenance. Bartholomai (1975: 232) incorrectly identified this specimen as QM F6869, but the correct number is provided later (Bartholomai 1975: 233). Bartholomai (1975) strongly suggested that the other specimen (QM F3589; Fig. 8L) was also derived from the Chinchilla Sand.
Stratigraphic and geographic range.—Pliocene; central and eastern Australia.
Genus Protemnodon Owen, 1874
Type species: Protemnodon anak Owen, 1874, eastern Darling Downs, Pleistocene.
Protemnodon devisi Bartholomai, 1973
Fig. 8F.
Material.—QM F4710 (holotype, partial left mandibular ramus) from Chinchilla, Australia, Pliocene.
Remarks.—Bartholomai (1973) revised the genus Protemnodon, and erected two new species on the basis of Chinchilla material.
Stratigraphic and geographic range.— Pliocene; central and eastern Australia.
Protemnodon chinchillaensis Bartholomai, 1973
Fig. 8H.
Material.—QM F5246 (holotype, partial right mandibular ramus) from Chinchilla, Australia, Pliocene.
Remarks.—See above.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Genus Bohra Flannery and Szalay, 1982
Type species: Bohra paulae Flannery and Szalay, 1982, Cathedral Cave, New South Wales, Pleistocene.
Bohra wilkinsonorum Dawson, 2004
Fig. 8I.
Material.—QM F43277 (holotype, right maxillary fragment) from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Dawson (2004b) described the first tree kangaroo from Chinchilla. Hocknull (2005) later confirmed the presence of Bohra sp. in the Chinchilla Local Fauna on the basis of an isolated calcaneum (QM F49453). Its relationship to B. wilkinsonorum is unclear.
Stratigraphic and geographic range.—Pliocene; Chinchilla.
Genus Silvaroo Dawson, 2004
Type species: Silvaroo bila Dawson, 2004, Chinchilla, Pliocene.
Silvaroo bila Dawson, 2004
Fig. 8M.
Material.—QM F43280 (holotype, left mandibular ramus), QM F43276 (paratype, left maxillary fragment), and QM F43292 (paratype, incisor) all from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Dawson (2004a) described the above material as a new genus and species of wallaby.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Silvaroo sp.
Fig. 8N.
Material.—QM F43281 (right mandibular ramus) from Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Dawson (2004a) described a right mandibular ramus from Chinchilla, which she suggested might represent a second, smaller species of Silvaroo.
Infraclass Eutheria Huxley, 1880
Order indeterminate
Genus Chronozoon de Vis, 1883
Type species: Chronozoon australe de Vis, 1883, Chinchilla, Pliocene.
Chronozoon australe de Vis, 1883
Fig. 6E.
Material.—QM F610 (holotype, calvarium) from Chinchilla, Australia, Pliocene.
Remarks.—De Vis (1883a) originally suggested the species might share affinities with sirenians. Mahoney and Ride (1975) suggested that it might be part of the skull of a giant wombat, whereas Reinhart (1976) interpreted it as a likely sirenian of unknown affinity. We consider Chronozoon australe as species inquirenda.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Order Rodentia Bowdich, 1821
Family Muridae Illiger, 1811
Genus Pseudomys Gray, 1832
Type species: Pseudomys australis Gray, 1832, Liverpool Plains, eastern New South Wales, Recent.
Pseudomys vandycki Godthelp, 1990
Material.—QM F16834 (holotype, right maxillary fragment) from conglomerate, main gully of Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Godthelp (1990) described the only species of rodent from Chinchilla. This specimen could not be located in the QM collection. Rich et al. (1991) suggested that there might be other Pseudomys species in the Chinchilla deposits.
Stratigraphic and geographic range.—Pliocene; eastern Australia.
Order Chiroptera Blumenbach, 1779
Family Molossidae Gray, 1821
Genus Mormopterus Peters, 1865
Type species: Mormopterus jugularis Peters, 1865, Madagascar, Recent.
Mormopterus sp.
Material.—QM F30575 (left upper canine) from “Wilkinson's Quarry”, Chinchilla Rifle Range, Australia, Pliocene.
Remarks.—Hand et al. (1999) described the first bat from Chinchilla, a species of Mormopterus, and referred the specimen, QM F30575 (canine), to the genus. The registration number QM F30575 has also inadvertently been assigned to two different fossil bats: a maxilla of Hipposideros winsburyorum (Hand and Godthelp 1999) and an isolated molar of Icarops aenae (Hand et al. 2005); both fossils are from Pleistocene and Miocene sites, respectively, of Riversleigh. An erratum is currently being prepared by Sue Hand (personal communication 2015) to correct the registration discrepancy.
Discussion
Sixty-three taxa have been reported in the literature from the Chinchilla Sand (SOM: Tables 2, 3). There are two families of fishes (Ceratodontidae, 1; Neoceratodontidae, 1), although a conference abstract by Mackness et al. (1999) also mentioned the occurrence of catfish (Plotosidae). There are 11 reptile taxa, comprising two families of Testudines (Trionychidae, 1; Chelidae 1), one family of Crocodylia (Crocodylidae, 2), and five families of Squamata (Agamidae, 1; Gekkonidae, 1; Scincidae, 2; Varanidae, 2; Madtsoiidae, 1). Thirteen bird taxa have been described from Chinchilla, comprising one family each of Anseriformes (Anatidae, 3), Ciconiiformes (Ciconiidae, 1), Galliformes (Megapodiidae, 1), Gruiformes (Rallidae, 2), Charadriformes (Charadriidae, 1) and one Charadriformes indeterminate, Casuariiformes (Casuariidae, 1), Pelecaniformes (Pelecanidae, 1), Phalacrocoraciformes (Phalacrocoracidae, 1), and Falconiformes (Accipitridae, 1). The fauna includes 39 mammal taxa, including one order indeterminate; two families of Dasyuromorphia (Dasyuridae, 2; Thylacinidae, 1), seven families of Diprotodontia (Phascolarctidae, 3; Diprotodontidae, 3; Palorchestidae, 1; Vombatidae, 1 (however see note added in proofs below); Thylacoleonidae, 1; Phalangeridae, 1; Macropodidae, 18; Phalangeridae, 1), and one Diprotodontia indeterminate, and one family each of Peramelemorphia (Peramelidae, 1), Rodentia (Muridae, 1), and Chiroptera (Molossidae, 1).
Nineteen species have been erected on the basis of specimens undoubtedly from the Chinchilla Sand: Metaceratodus palmeri, Tiliqua wilkinsonorum, Dasyurus dunmalli, Archerium chinchillaensis, Euowenia grata, Thylacoleo crassidentatus, Sthenurus notabilis, Wallabia indra, Macropus woodsi, Protemnodon devisi, Protemnodon chinchillaensis, Bohra wilkinsonorum, Silvaroo bila, Pseudomys vandycki, Archizonurus securus, Phalanger procuscus, Brachalletes palmeri, Synaptodon aevorum, and Chronozoon australe. Taxonomic revision of some of those species, in particular the last five, is urgently needed and will probably reduce this number. Type specimens of ten additional species are most likely derived from the Chinchilla area: Ciconia nana, Pelecanus proavus, Necrastur alacer, Koalemus ingens, Koobor notabilis, Euryzygoma dunense, Troposodon gurar, Simosthenurus antiquus, Macropus pan, and Macropus dryas.
Uncertainties exist regarding the provenance of certain species supposedly from Chinchilla (SOM: Table 3), and in almost all cases, of provenance within the Chinchilla area itself (Price 2012). Most early collecting did not differentiate between Chinchilla and the eastern Darling Downs, so confusing Pliocene and Pleistocene localities respectively. Bartholomai (1963, 1966, 1967, 1968, 1971, 1973, 1975, 1976), Olsen (1975, 1977), Boles (2005), van Tets (1974), and others have provided some clarification on provenance. Later collectors differentiated between Chinchilla and the eastern Darling Downs, but precise geographic or stratigraphic information for most specimens in the QM is lacking. Most are listed as coming from the Chinchilla Rifle Range, and derived from the Chinchilla Sand Formation (sensu Woods 1960); exact horizons within the Pliocene stratigraphic sequence is rarely documented. Therefore, the Chinchilla Local Fauna may represent an assemblage with considerable temporal and spatial averaging (Price 2012). These problems have arisen because of the lack of stratigraphically controlled, systematic excavation. Difficulties of dating the site add to the problems, with age differences of up to ca. 500 ka between biocorrelated Local Faunas (Tedford et al. 1992; Mackness et al. 2000a).
Several other taxa are not discussed above, but have been listed by others as from Chinchilla. Three species of birds are mentioned by Rich et al. (1991) in their species list for Chinchilla fossils: a magpie goose (cf. Anseranas semipalmata), a probable Cape Barren goose (cf. Cereopsis), and a black swan (Cygnus sp. cf. C. atratus), none of which are represented by fossil material in the Queensland Museum. Although de Vis (1905) was responsible for erecting those three taxa, he based his descriptions on specimens from the late Pleistocene Lake Eyre Basin of central Australia, and we can find no evidence that these three birds are part of the Chinchilla fauna.
De Vis (1892) described a number of fossil bird species, but with no introductory remarks, so the provenance of the fossils described therein is uncertain. Olson (1975) noted the similarity between the 1892 paper and de Vis's (1888c) previous treatise on fossil birds, and inferred that the 1892 paper was a continuation of the 1888 paper. De Vis (1888c: 1277) referred to specimens “yielded by the Darling Downs in the immediate neighbourhood of Chinchilla”, and we follow Olson (1975) in assigning all fossils described in de Vis (1888c) and de Vis (1892) to the Chinchilla Sand.
Dawson and Flannery (1985) mistakenly listed Macropus thor as being restricted to the Chinchilla Sand, incorrectly citing Bartholomai (1975), who in fact stated that the species is restricted to Pleistocene deposits. Bartholomai (1975) referred a juvenile mandibular ramus (QM F4230) supposedly from Chinchilla to Macropus titan (= M. giganteus titan; Dawson and Flannery 1985), but suggested that on basis of its preservation, the specimen was more likely from Pleistocene deposits.
Flannery and Archer (1983) recognised Troposodon kenti among the Chinchilla fossil material in the QM, but did not refer any specimens to the species. A search of the collections did not locate any specimens of definitive Chinchilla provenance.
Tedford et al. (1992) suggested that two specimens (QM F5580 and F10293) attributable to Diprotodon sp. cf. Diprotodon optatum are from Chinchilla. Mackness and Godthelp (2001) later concluded that those specimens were more likely from the Pleistocene site of Warra, and hence that Diprotodon is exclusively Pleistocene (Price 2008b).
Both Bartholomai and Marshall (1973) and Archer and Bartholomai (1978) noted Phascolonus from Chinchilla. Ride (1964) suggested that the “Darling Downs” term used by de Vis (1891b) in his description of Queensland wombats might include the Chinchilla Sand. Fossil material attributable to large-bodied wombats that are of definitive Chinchilla provenance are currently under study.
Gaffney (1981) reported a specimen of giant horned turtle (meiolanid) supposedly from the Chinchilla Sand, but coming from Armour Station, near Macalister, over 50 km southeast of the known exposures of the Chinchilla Sand. Molnar and Kurz (1997) also recorded Diprotodon from Armour Station, which on the basis of biostratigraphy, suggests that the meiolanid is actually Pleistocene, not Pliocene, and certainly not from the Chinchilla Sand.
Tedford et al. (1992) reported Zygomaturus sp. cf. Zygomaturus trilobus in the Chinchilla deposits. It is unclear whether this is the same specimen examined by Archer and Wade (1976) who assigned it to Zygomaturus sp. (listed above).
Finally, Rich et al. (1991) and Archer and Hand (1987) both mention unidentified phalangerids from Chinchilla; however, the only potential phalangerids from Chinchilla are the equally problematic Phalanger procuscus and Archizonurus securus. Undoubted phalangerid material from Chinchilla is in the QM collection, but is currently unregistered and awaits detailed study.
Despite these limitations and uncertainties, Chinchilla remains one of, if not the most important Pliocene sites in Australia. Its faunal diversity is unparalleled amongst current Australian Pliocene vertebrate records, and the preservation of its fossil material is largely excellent. Moreover, due to its richness and long history of collection, abundant specimens are available for study (see Note added in proofs below).
Note added in proofs
Since the online publication of this article, the wombat material from Chinchilla has been described and published (Louys 2015). Five species referred to the Vombatidae are described from the Chinchilla Sand, bringing the total number of vertebrate taxa recovered from this formation to sixty-seven.
Acknowledgements
We thank Ces and Doris Wilkinson for their generous donations of Chinchilla specimens to the Queensland Museum (Brisbane, Australia), and their continued support of our research. We thank the staff at the QM, in particular Jo Wilkinson, Kristen Spring, Andrew Rozefelds, Pam Wilson, and Scott Hocknull, who provided access to specimens and checking of museum registers. Grant Gully and Gavin Prideaux (Flinders University, Bedford Park, Australia) kindly provided photographs of Macropus woodsi, Prionotemnus palankarinnicus, and Silvaroo sp. specimens; and Robin Beck (University of New South Wales, Sydney, Australia) photographs of Archerium chinchillaensis. Mel Wilkinson and Gregg Webb (University of Queensland, St Lucia, Australia) provided helpful discussions regarding the geology of the site. GJP acknowledges the support of an ARC Discovery Early Career Researcher Award (DE120101533). JL was supported by a UQ Postdoctoral Fellowship and a UQ Early Career Research Grant (2012003222). We thank Robin Beck (UNSW), Karen Black (UNSW), Andy Herries (La Trobe University, Melbourne, Australia), Kenny Travouillon (UQ), Gavin Prideaux (Flinders University), Andrew Rozefelds (QM), Trevor Worthy (Flinders University) and Editor Mike J. Benton (University of Bristol, UK) for their comments and suggestions that significantly improved this manuscript.
