Taxonomy of Australian Mammals

Corrigendum – Taxonomy of Australian Mammals

Page 22, Column 1, Lines 31–51. The entry for the Family Phalangeridae Thomas, 1888 sensu Flannery et al., 1987 [Cuscuses and Brush-tailed Possums] should read:

‘Family Phalangeridae Thomas, 1888 sensu Ruedas and Morales, 2005 [Cuscuses and Brush-tailed Possums]

Subfamily Phalangerinae Thomas, 1888

Phalanger mimicus Thomas, 1922 Southern Common Cuscus

Spilocuscus nudicaudatus (Gould, 1850) Australian Spotted Cuscus

Subfamily Trichosurinae Flynn, 1911

Trichosurus caninus (W. Ogilby, 1836) Short-eared Brush-tailed Possum

Trichosurus cunninghami Lindenmayer et al., 2002 Mountain Brush-tailed Possum

Trichosurus vulpecula (Kerr, 1792) Common Brush-tailed Possum

Wyulda squamicaudata Alexander, 1919 Scaly-tailed Possum’

Page 26, Column 1, Lines 14–27 and individual species entries on pages 238–241. The bats of the genus Pteropus are typically referred to as ‘Flying-fox’ rather than ‘Fruit-bat’, so these species should read:

Pteropus alecto Temminck, 1837 Black Flying-fox

† Pteropus brunneus Dobson, 1878 Percy Island Flying-fox

Pteropus conspicillatus Gould, 1850 Spectacled Flying-fox

Pteropus macrotis Peters, 1867 Large-eared Flying-fox

Pteropus natalis Thomas, 1887 Christmas Island Flying-fox

Pteropus poliocephalus Temminck, 1825 Grey-headed Flying-fox

Pteropus scapulatus Peters, 1862 Little Red Flying-fox

Page 123, Column 1. The heading for the Family Phalangeridae entry should include sensu Ruedas and Morales, 2005 not Flannery et al. 1987, so it should read:

‘Family Phalangeridae Thomas, 1888 sensu Ruedas and Morales, 2005

Family Phalangeridae Thomas, 1888a: xii, 126.’

Page 123, Column 2, Paragraph 1, Lines 9–18. The wording should read:

‘Flannery et al. (1987: 477, 503) recognised two subfamilies, the Ailuropinae (Flannery et al., 1987: 477, 503) that included the genus Ailurops Wagler, 1830: 26, and the Phalangerinae that included the tribes Phalangerini and Trichosurini. More recently Ruedas and Morales (2005: 362) proposed that the Family Phalangeridae is composed of the subfamilies Phalangerinae (including Phalanger and Spilocuscus), Ailuropinae (including Ailurops Wagler, 1830: 26 and Strigocuscus Gray, 1862a: 319) and Trichosurinae (including Trichosurus and Wyulda). This arrangement is followed here.’

Page 124, Column 1. The entry for the Subfamily Phalangerinae Thomas, 1888 should read:

‘Subfamily Phalangerinae Thomas, 1888

Subfamily Phalangerinae Thomas, 1888a: xii, 135.

Type genus: Phalanger Storr, 1780.

Comments: When originally proposed, this rank was placed in the Suborder Diprotodontia (Owen, 1877a) and included the subfamilies Tarsipedinae (Thomas, 1888a [ = Tarsipedidae (Gervais & Verreaux, 1842a)]), Phalangerinae (Thomas, 1888a) and Phascolarctinae (Thomas, 1888a [ = Phascolarctidae (Owen, 1839a)]). The inclusion of the possums, gliders and koala within the Family Phalangeridae was followed by Bensley (1903: 125) and Simpson (1945: 46), although the taxa representing the Families Phascolarctidae, Acrobatidae, Burramyidae, Petauridae, Pseudocheiridae and Tarsipedidae were subsequently removed (see individual entries).’

Page 124. Delete entry for Tribe Phalangerini Thomas, 1888 sensu Flannery et al., 1987.

Page 126, Column 2 and page 127, Column 1. The entry for the Tribe Trichosurini T. Flynn, 1911 should read:

‘Subfamily Trichosurinae T. Flynn, 1911

Family Trichosuridae T. Flynn, 1911: 120.

Type genus: Trichosurus Lesson, 1828b.

Comments: When originally proposed, this rank was placed in the Marsupialia (Illiger, 1811) and included the genus Trichosurus Lesson, 1828b. Not recognised subsequently at family rank. Tribe Trichosurini recognised by Flannery et al. (1987: 477, 503), Marshall et al. (1990: 494), Norris (1994: 93), Kirsch et al. (1997: 245) and Groves (2005b: 49), but not by Strahan (1983: xxi; 1995: 7, 265) or Van Dyck and Strahan (2008: 10, 265). Synonymised within Phalangeridae by Marshall et al. (1990: 493) and McKenna and Bell (1997: 61). Subfamily Trichosurinae recognised by authors including Kirsch (1968a: 420), Marshall (1981: 28; 1984: 98), Kirsch and Wolman (2001: 23, 29) and Ruedas and Morales (2005: 362).’

Page 239, Column 1, Paragraphs 6 and 7. Homonym entries should read:

Pteropus nicobaricus Fitzinger, 1861, the Black-eared Flying-fox of the Class Mammalia (Order Chiroptera, Family Pteropodidae). Taxon is a nomen nudum and synonym of Pteropus melanotus Blyth, 1863 (see Simmons, 2005a: 341).

Pteropus nicobaricus Zelebor, 1869, the Blackeared Flying-fox of the Class Mammalia (Order Chiroptera, Family Pteropodidae). Taxon is a synonym of Pteropus melanotus Blyth, 1863 (see Simmons, 2005a: 341).

TAXONOMY OF

AUSTRALIAN MAMMALS

Stephen Jackson and Colin Groves

© Stephen Jackson and Colin Groves 2015

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Jackson, Stephen M., author.

Taxonomy of Australian mammals / Stephen Jackson and Colin Groves.

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Includes bibliographical references and index.

Mammals – Australia – Classification.

Groves, Colin P. (Colin Peter), author.

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Front cover (from top left): Koala (Phascolarctos cinereus), Torresian Striped Possum (Dactylopsila trivirgata) and Eastern Quoll (Dasyurus viverrinus). Photos by Stephen Jackson.

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Contents

Introduction

Acknowledgements

Definitions of nomenclatural and bibliographic terms

Abbreviations

Taxonomy of the mammals of Australia

References

Appendix

Index of common names

Index of scientific names

Introduction

The Australian continent, associated islands and territories oceans comprise 411 species (which includes one undescribed native species and 33 introduced species) of mammals that are recognised here. The scientific naming of the mammals of Australia began on 1 January 1758 when many of the cetaceans and animals that would subsequently be introduced into Australia were described by Carl Linnaeus in the 10th edition of the Systema Naturae, which is the starting point for all modern scientific names. The taxonomy of mammals from Australian soil began in 1785 when the Eastern Ring-tailed Possum was described, which was followed by the Eastern Grey Kangaroo in 1790 and the Yellow-bellied Glider in 1791.

After the colonisation of Australia by Europeans in 1788 there was a rapid increase in the number of native species described, with 165 new species recognised by 1860 and 230 species by 1900 (Fig. 1). Since this time each decade has resulted in the descriptions of many new species, with the most recent decades showing an increase in the rate of new species being discovered. These recent discoveries have included many native dasyurids, bats and rodents as might be expected because of their small size and more cryptic nature, but has also included five macropods. Incredibly, since 2000 a total of four new species of cetaceans have also been described from the waters off the Australian coast, with several others identified and awaiting formal description. With the development of new genetic technology and the reassessment of previously described populations, subspecies or synonyms that had not been adequately recognised (apparently unjustly) there has increasingly been a recognition of many of these as distinct species or subspecies. So it is likely that there are dozens of new species awaiting formal description or re-classification, which include not only the small cryptic species but also other larger species such as the cetaceans.

Fig. 1. The rate of descriptions of Australian mammals over time.

The rate at which species of the different orders of Australian mammals have been described by science has varied considerably over time (Fig. 2). Some groups such as the monotremes, bandicoots, seals, and exotic species were all described by 1900, while other groups such as the diprotodontians, chiropterans, rodents and dasyurids have continued to steadily increase in number in the last 30 years. Given the current knowledge of the taxonomy of Australian mammals it is clear that groups such as the dasyurids, chiropterans, rodents and the cetaceans will continue to reveal further species well into the future.

Fig. 2. Cumulative description of the different groups of Australian mammals.

Taxonomy is a continually evolving process where the work of each author is built upon as more species are described and as taxonomic studies of species, genera and larger groups reveal a clearer picture of their phylogenetic relationships. With this in mind this present research acknowledges the important and ground-breaking work of its predecessors, especially those by Waterhouse (1841a, 1846), Gould (1845–1863), Thomas (1888a), Lydekker (1894a), Iredale and Troughton (1934), Ride (1970),Walton (1988a), Strahan (1983, 1995), Van Dyck and Strahan (2008) and the enormous number of taxonomic revisions of specific groups that have been undertaken.

Despite the production of various excellent books on Australian mammals over the last few decades there is no current comprehensive checklist that provides a full list of scientific names and a list of agreed common names – indeed, there is no formal system for setting up an agreed list of common names. Therefore an effort has been made here to set and follow several rules in the formation of common names. With respect to higher taxonomic ranks, the absence of an updated checklist in Walton (1988a) has resulted in some authors such as Van Dyck and Strahan (2008) perforce retaining a somewhat outdated higher taxonomy, while Menkhorst and Knight (2011) included only the families within each order without any further division. Therefore an exhaustive effort has been made to include all higher ranks and their associated synonyms.

The taxonomy, common names and phylogenetic sequence used here typically follow Van Dyck and Strahan (2008) for accepted species names and ranks up to family, in order to maximise consistency with the most current treatment of all Australian mammals. Ranks above family typically follow Van Dyck and Strahan (2008), although there are some notable exceptions as a result of significant recent research. These include the acceptance of the subordinal division of Chiroptera into Yinpterochiroptera and Yangochiroptera in preference to the old division into Megachiroptera and Microchiroptera (the latter of which is now known to be non-monophyletic), and the placement of the cetaceans within the Artiodactyla. The synonymies of species typically follow Wilson and Reeder (2005), those of genera following Wilson and Reeder (2005) and McKenna and Bell (1997), and those above the generic level following McKenna and Bell (1997), but we have ventured to register our disagreement in many places due to the recognition of appropriate recent research, and we give reasons for this deviation.

Unlike previous taxonomic treatments of Australian mammals, a full list of all known homonyms is provided including their full citation and taxonomic hierarchy. This review is also unique in that it includes subspecies and associated synonyms that are extralimital in order to give a full account of the distribution of species that occur within Australia and its territories. To rationalise the taxa covered, and the volume of the final text, we do not include synonyms at any rank for those species that are introduced to Australia unless the taxa concerned have synonyms based on specimens collected within Australia.

A deliberate attempt has been made to undertake this revision from first principles by making every effort to personally view and obtain a copy of every citation used. This process has revealed many discrepancies within the published literature so it is hoped that this treatment has helped to rectify many of these inconsistencies while at the same time it is hoped to not create many new ones. We also hope that this review helps to identify priority areas for future research so that the cycle of taxonomic research and review can be expedited.

In order to assist in the accuracy of the dates of the references used, and in turn the date priority of names used, an effort has been made to give each reference an exact or approximate date of publication depending on the available information. To undertake this process the primary source or numerous secondary sources were used including the Biodiversity Heritage Library website, journal websites, and numerous secondary papers and books that have been published. Appendix 1 includes a table of references included in the text and sources that have been used to include their dates of publication. Some publications were released over multiple years, and have been referred to in the text by the year the particular information was published along with the range of years in which the book was published. For example, the dates of John Gould’s The Mammals of Australia spanned from 1845 to 1863 and it was released in 13 parts in three volumes. Taxa that were first described by Gould in that book have been given their specific year of publication using the publication dates proposed by Waterhouse (1885). References to this publication are cited with the year that the plate and associated text were published with [1845–1863] placed immediately afterwards so there is no confusion within the reference list.

The most recent complete review of Australian mammal taxonomy by Walton (1988a) is now well out of date by the description of new species and the major rearrangements of genera, families and even orders that has occurred as a result of subsequent research. Therefore the aims of this publication are to:

• Present an up to date taxonomic list of all Australia mammals and every rank from species (and subspecies and synonyms) up to the Class Mammalia.

• Provide a history of the major taxonomic changes that have occurred during the life of each taxon’s name from the time of its initial description, and therefore provide a justification of the taxonomy used in this review.

• Provide an indication of the current stability of names by showing how names have changed over time, and show whether each name has stabilised over time or still shows fluctuation.

• Make taxonomic decisions where appropriate to assist in the stabilisation of names used.

• Include all taxa that occur outside Australia, but which have representative native species within Australia, in order to give a greater context to the Australian taxa.

• Include all species that occur within island territories of Australia including Christmas Island, Torres Strait islands, Lord Howe Island, Norfolk Island, Macquarie Island, Heard Island and McDonald Islands, and also the Australian Antarctic Territory, for all that this is not formally recognised internationally.

• Include all species occurring within the territorial waters of Australia and the island territories listed above, and the Australian Antarctic Territory.

• Provide details of all homonyms including non-mammalian names.

• Identify areas where future taxonomic research should be focussed.

Taxonomic decisions

The aim of this work is to provide a taxonomy that reflects the current state of knowledge and to document the enormous literature that has been published on studies focussed at all ranks within the Class Mammalia down to subspecies, and the synonyms for each rank. For each taxonomic rank an effort has been made to review the primary literature to either support its rank or synonymise it within other taxa. As part of this process the major taxonomic decisions have been identified from the primary literature. The construction of synonyms has endeavoured to follow that developed by Gardner and Hayssen (2004).

Common names

The first attempt to establish a recommended list of common names, or vernacular names, was published by Strahan (1980a) on behalf of a committee of the Australian Mammal Society. The general principles of the names proposed by the committee were to (where possible) be descriptive, indicate relationships, be memorable, be euphonious and be as short as possible. They also recognised the value of Aboriginal names such as Kultarr, Mulgara, Quoll, Kowari, Dunnart, Numbat, Bilby, Bettong, Pademelon, Potoroo and Quokka.

Common names used here are generally derived from those of Strahan (1980a) and more recently by Van Dyck and Strahan (2008) but with several differences. The names used are typically well known and commonly referred to, ideally descriptive, short and pronounceable. The noun-part of the name should correspond with the genus and the adjectival part should, if possible, refer to a diagnostic external character but may refer to the distribution. Whenever possible, names that are parochial and convey little or no information, for example ‘Mitchell’s Hopping-mouse’, should be avoided. Names such as Monjon for Petrogale burbidgei and Nabarlek for Petrogale concinna are also not encouraged as they should include ‘Rock-wallaby’ in the name. With this in mind the names Kakarratul for Notoryctes caurinus and Itjaritjari for Notoryctes typhlops have been replaced by the Northern Marsupial Mole and Southern Marsupial Mole respectively following Groves (2005a). Familiarity may make terms such as Ringtail and Brushtail, as used by Groves (2005b), acceptable but for clarity are improved by the inclusion of ‘Possum’ in the group name.

Though common names can cause confusion on occasion, a perhaps unexpected advantage is that they typically remain constant when scientific names are changed. Importantly, common names and scientific names perform different functions, with common names reflecting the phenotype while the scientific names reflect the genotype (Andrew 2008). An example of this is the term Marsupial Mole that is used for the marsupial Order Notoryctemorphia as opposed to the true placental golden moles of the Order Afrosoricida. Common names have generally been restricted to the species rank, although they have been allocated to subspecies in some cases where these have been recognised in the modern literature – usually because the subspecies have extralimital distributions with recognised regional names or have been proposed to potentially warrant being elevated to species rank.

The protocols for the construction, hyphenation and capitalisation of common names used here draws on Duckworth and Pine (2003), Armstrong and Reardon (2006) and Andrew (2008) in an endeavour to give the names a consistent construction. For example, the free-tail bats have been spelt Freetail Bat by Menkhorst and Knight (2011), Free-tailed Bat by Simmons (2005a) and Van Dyck and Strahan (2008), and free-tailed bat by Armstrong and Reardon (2006).

The protocol of compounding group names is well established in some Australian taxa, with names including Rock-wallaby and Tree-kangaroo in common usage. Elsewhere, retaining the adjectival form, where possible, keeps groups large for unfamiliar taxa, for example Bent-winged Bat (rather than Bentwing-bat, but Horseshoe-bat because the adjectival form is too clumsy as in Horse-shoe-marked Bat). Thus the generalised rule adopted by Andrew (2008) and followed here is to compound group names comprising two nouns, but to retain two words when the first part of a group name is adjectival. Thus the group of possums with ‘ringtails’ (the ‘ringtail possum’ of Strahan, 1980a) should be compounded as Ringtail-possum or become adjectival as in Ring-tailed Possum. The latter retains larger groups and is easier to find in an index for those less familiar with, for example, the Pseudocheiridae. This has the added advantage of being grammatically correct but we recognise there are still challenges in Cetacea.

Species concepts

In the light of the discussions of species concepts over the last 20 years or so, the failure of earlier checklists to specify what precisely they mean by species and subspecies seems a fault; but, given that the Biological Species Concept of Dobzhansky (1937) and Mayr (1942) held sway almost unchallenged for some 50 years since the period of the Evolutionary Synthesis in the 1930s, it was understandable. Today, however, it is mandatory for taxonomists to be clear about what species concept they adopt, and whether or not they recognise the rank of subspecies.

The Dobzhansky/Mayr concept defined species as not interbreeding with each other in nature. It has been pointed out many times that this gives no guidance for allopatric populations: under what circumstances should they be classified as distinct species, and when should they be combined into one species? For this reason alone, taxonomists should be aware that a simple claim to be working with the Biological Species Concept cannot be accurate: they are very evidently using some extra, unspoken criterion or multiple criteria.

We now know, however, that what we have always classed as distinct species, which remain discrete in sympatry, may actually be interbreeding on the sly – introgression, unbeknownst to the taxonomist and even usually to the field worker, may well be occurring or have occurred, its only evidence being the presence of the ‘Wrong’ mitochondrial or other genetic components in a given population. It would clearly be nonsense to unite two species on such grounds. The Biological Species Concept, defining species as not interbreeding, actually does not work.

Molecular data have been used to create what has become known as the Genetic Species Concept, based on the idea that the amount of genetic difference necessary for reproductive isolation can be measured (Mayden, 1997). Bradley and Baker (2001) tested this using Cytochrome-b sequences for several bat and rodent genera, and found that genetic distances do indeed tend to increase in the expected sequence from intrapopulational via intrasubspecific and intraspecific to intrageneric; moreover, distances between presumed sister species were less than between congeneric species in general. In a later paper the same authors amplified this, and offered a definition of the genetic species as ‘a group of genetically compatible interbreeding natural populations that is genetically isolated from other such groups. Under our definition of the Genetic Species Concept, speciation is the accumulation of genetic changes in 2 lineages’ (Baker & Bradley, 2006). In their accompanying table, they compared the Genetic Species Concept with other concepts, showing that most of the criteria for specific status under the concept were very close to those under the Phylogenetic Species Concept (see below), with this important difference: there must be a certain amount of difference in certain systems (nowadays, this would equate to DNA sequences) between two taxa in order to qualify, which, they argue, would constitute proof that they have entered separate evolutionary trajectories. This amount would have, of necessity, to vary across taxonomic groups, unless it is to be set arbitrarily.

Many authors working with DNA data have either explicitly or (more usually) tacitly used ‘amount of difference’ in making their taxonomic judgments. This raises the question of precisely what amount of difference would suffice, and, importantly, why. Baker and Bradley (2006) faced this problem and noted that it is a matter of probabilities: there is simply a greater probability that speciation has occurred with genetic distances of >5% than with distances of <2%. When faced with allopatric populations, the response of the Genetic Species Concept is similar to that of the Biological Species Concept: compare the amount of difference between two allopatric populations with that between two acknowledged congenerics (an argument which risks being circular!). There is also the question of how a difference in one DNA sequence (such as the Control Region) would equate to a difference in another (such as 12S or, even more problematic, a nuclear sequence such as an intron or pseudogene). Then there is of course the problem of species for which no genetic data are available, including fossils for which – except for rather recent specimens, preserved under appropriate conditions of fossilisation – no DNA sequencing will ever be possible. Given the arbitrariness of the ‘amount of difference’ criterion, we cannot follow the Genetic Species Concept, while recognising the enormous contributions that its adherents have made to the field. In effect, the Genetic Species Concept is less a species concept as such than a recipe for uncovering the existence of cryptic species; Baker and Bradley (2006), indeed, emphasise this role of the concept, and indeed we acknowledge that its considerable merit is its past history of, and future potential for, recognising cryptic evolutionary lineages within what has hitherto been considered a single species.

Our criterion for species status in this work is what has been called the Phylogenetic Species Concept: ‘species are populations (or aggregations of populations) that are diagnosably distinct’. In this concept, we often have to assume that our species are populations (although this is often amenable to testing); apart from that, we need make no assumptions, noting only that on the evidence before us the ranges of variation of two species are separate. To put it another way, species have fixed heritable differences between them (Groves and Grubb, 2011). Such differences may be pelage characters, morphometric characters (either single measurements, or separation under Discriminant Analysis), or consistent differences in DNA sequences. It is in this latter area that genetic data are valuable for the delimitation of species.

Above all, the Phylogenetic Species Concept is testable, as a scientific proposition should be: ‘the criterion of the scientific status of a theory is its falsifiability, or refutability, or testability’ (Popper, 1963). Diagnosability (whether the differences are fixed or not) is testable; the question of whether the candidates for species status are populations (or metapopulations) is in principle testable; and the heritability of the differences is likewise in principle testable, although we must acknowledge that the heritability of morphological differences may well be unproven on the available evidence, but it is clearly best to act on the working hypothesis that they are heritable until breeding or rearing experiments prove otherwise.

In a significant clarification, De Queiroz (2007) proposed what he calls the Unified Species Concept – species are ‘separately evolving metapopulation lineages (or, more properly, segments thereof)’ (De Queiroz, 2007); there are particular properties, or lines of evidence, by which species may be delimited, but these are not themselves ‘species concepts’. This has the effect of transferring the disputes from the question of which ‘species concepts’ are appropriate to the question of what evidence is appropriate for delimiting species, and it is relevant that De Queiroz goes on to argue that one consequence of this view is that ‘any evidence of lineage separation is sufficient to infer the existence of separate species . . . Although presence of a single property provides evidence for lineage separation, a highly corroborated hypothesis of lineage separation . . . requires multiple lines of evidence’ (De Queiroz, 2007: 884). That is to say, species delimitation relies on what has hitherto been called the Phylogenetic Species Concept.

The De Queiroz model significantly clarifies the discussion: there is only one way of defining species, and that is as evolutionary lineages. The ‘phylogenetic species’ is the minimal delimitation of species; while not necessarily ‘highly corroborated’, it offers clear evidence, falsifiable when fresh evidence comes to hand, that lineage separation has occurred.

In this way, the decision of whether two populations are specifically distinct or not is objective; it depends entirely on the evidence to hand, not on an arbitrary ‘enough difference’ nor on the basis of whether they would interbreed or not if given the chance.

Subspecies

Subspecies are geographic divisions within a species, which differ to a certain extent from each other in heritable features. The usual criterion is that 75% of individuals of a subspecies should be distinguishable from all individuals of other subspecies within that species, but this is a somewhat arbitrary criterion (Amadon, 1949). Essentially, a subspecies may be regarded as the point on a continuum at which it becomes convenient to dignify a population with a separate name (see, for example, Groves, 2001): the concept is, accordingly, a ‘convenience category’, and subspecies should on no account be reified. Nonetheless, as argued by Parkes (1982), subspecies serve as a conservative buffer for taxonomic uncertainty; this is especially true for cases where future taxonomic inquiry may demonstrate that currently recognised subspecies should be elevated to specific rank (e.g. Braby et al., 2012). In contrast to the more recent support of subspecies, several authors have suggested that they be abandoned because they are often poorly diagnosed, their delimitation is difficult to determine (especially for parapatric populations), and taxonomic decisions made for a particular set of populations are often arbitrary, subjective, and based on too few characters (Wilson and Brown, 1953; Gillham, 1956).

Subspecies are not the only infraspecific category. Evolutionarily Significant Units (‘ESU’) (Moritz, 1994), are discrete populations of notably divergent evolutionary history and (consequently) genetic composition; very often, they are found to be equivalent to phylogenetic species.

Genera

A genus is a monophyletic group of species, but where to draw the line is at present arbitrary. Groves (2001) (see also Groves and Grubb, 2011) proposed that, to qualify as genera, two monophyletic groups should have been separated at least since the Miocene–Pliocene boundary. There seems little problem if molecular clock data (in the absence of the fossil record) indicate separation times, between candidates for generic status, of greatly above or below this level, but very often the inferred dates fall quite close to this boundary, and in such a case we would require much firmer data – more loci, better calculations of evolutionary rates, and so on – before proposing to change ‘traditional’ generic rankings. But it is mandatory that genera, and indeed other higher categories, be monophyletic, and if the evidence indicates otherwise, we have in this research ventured to alter the ‘traditional’ classification.

Families

A family is a monophyletic group of genera, but where to draw the line is at present arbitrary, as in the case of genera. Groves (2001, and elsewhere) proposed that, to qualify as families, two monophyletic groups should have been separated since at least the Oligocene–Miocene boundary. As in the case of genera, we have been conservative unless the evidence is strongly in favour of changing a ‘traditional’ arrangement.

Subgenera, subfamilies

When a genus or a family is too large and unwieldy in its contents, it may be convenient to divide it into two or more monophyletic groups called subgenera and subfamilies respectively. Their use is optional; for example, if a family contains just two known genera, however well differentiated, there is no need for subfamilies. Frequently the use of subfamilies is not sufficient, and tribes, and below that subtribes, may be used as subordinate (but always monophyletic) divisions. Subgenera, on the other hand, have been all too often used simply as a way of dividing a large and speciose genus without asking whether they are actually monophyletic. The informal term ‘species group’, which carries no implication other than sheer convenience, performs this function much better, and it is perhaps appropriate that the subgenus category seems to be falling out of use. Given the issues associated with subgenera they are not recognised here, so proposed subgenera including Notamacropus and Osphranter have been separated from Macropus, while Micronomus, Ozimops and Setirostris have been removed from Mormopterus as distinct genera.

Ranks above genus

In order to better understand the phylogeny of the different groups of Australian mammals, an effort has been made to include all ranks above genera and their associated synonyms. To better understand the original scope of each individual rank, the ranks above and below have been included. For family group names this includes all genera.

Given that the composition of most higher taxa has changed considerably since their original description, the current usage of the rank is also attributed to the author(s) from which this revision follows.

Higher non-Linnean ranks

The increasing research of the higher-order relationships of the different groups of mammals has resulted in a proliferation of names for ranks or clades above family group; as these names are not subject to the rules of priority that are required by the Code of Zoological Nomenclature there has been a degree of confusion as to which ones should be used. In an effort to provide stability to these names the principles of priority and stability as outlined by Simpson (1945), and followed by Brothers (1983) and enhanced by Asher and Helgen (2010), are followed. One of the main principles used is that the older name is retained if the proposed new name retains at least its original families. The senior and junior names of higher ranks of placental mammals advocated by Asher and Helgen (2010) are specifically followed.

Book outline

Each currently recognised taxon is identified with the relevant author and year of publication under which is the common name for species (and some subspecies) ranks. This is followed by a complete synonymy including the senior synonym and all known junior synonyms using the spelling and formation as described with the relevant author, year of publication and page number on which the name occurs. Sometimes two page numbers are included and these comprise the first listing of the name in the publication (often in an abstract) followed by the location where the name is formally described in the main text. The full citation for each author and year of publication is included within the reference section at the back of the text. Each synonym also includes the type genus for family group names, type species for genus group names and type locality for species group names. Within the comments section for ranks above the genus level an effort has been made to include the placement of the taxon within its taxonomic hierarchy by including the named rank above and ranks below, if they were provided, in order to give the reader a sense of the scope and context in which the proposed name was established. For convenience, the taxa that have individual entries do not typically have page numbers included when they are mentioned elsewhere in the text (only author and year). The comments section also outlines a broad taxonomic history of each taxon based on the major reviews. The synonymy for each taxon includes any unique name combination applied by any author for the first time to that taxon. This includes taxon names that have been made in error.

A future taxonomic research section is also included where appropriate. It very often happens that a particular genus or species has not been revised for many years, if ever, since its original description, and we point out if this has occurred and seems to be potentially a problem. We also take the liberty of pointing out, on occasion, when work of taxonomic relevance (especially a DNA phylogeny) has been published but the authors have not themselves ventured to tamper with the ‘accepted classification’ or where further confirmation of a taxonomic decision may be needed. Finally, a list of known homonyms is included within a taxonomic hierarchy for not only mammal groups but also other animal groups.

Acknowledgements

This work would not have been possible without significant assistance from numerous sources. We would like to thank those who helped read sections of this book and suggested numerous changes including Paul Andrew (Taronga Zoo), Ken Aplin (Smithsonian Institution), Joanne Burden (Muséum National d’Histoire Naturelle), Morgan Churchill (University of Wyoming), Mark Eldridge (Australian Museum), Erich Fitzgerald (Museum Victoria), Ewan Fordyce (University of Otago), Kris Helgen (Smithsonian Institution), Harry Parnaby (Australian Museum) and Terry Reardon (South Australian Museum). Stephen Jackson would like to thank several libraries and their associated staff that were very helpful in providing assistance during his visits. These include Carol Gokce, Paul Cooper, Eliza Walsh, Kirsten Marshall, John Rose and Emma Solway from the Natural History Museum in London who provided many of the references and assisted during the first visit to the library in 2003; Nicola Gamba, Paul Cooper, Lisa Di Tommaso, Samantha Gare, Nadja Noel, Kamila Reekie, Sarah Sworder, John Rose and Angela Thresher during the second visit in 2008; and Hellen Sharman, Sarah Sworder, Kamila Reekie, Harriet Campbell Longley, Lisa Atalla, Sarah Stewart, Goulven Keineg, Rachel Whittington, Elinor Skedgell, Lisa Di Tommaso and Lorraine Portch during the third visit in 2012. Thanks also to Therese Nouaille-Degorce and Evelyne Bremond-Hoslet from the Bibliothèque Centrale du Museum National d’Histoire Naturelle in Paris for providing several valuable references. Great thanks also to the staff at the National Museum of Natural History (Smithsonian Institution) libraries in Washington DC including Martha Rosen, Leslie Overstreet, Daria Wingreen-Mason and Kirstin van der Veen who helped him enormously in finding and copying references for this project. Thanks also to the staff of the National Natuurhistorisch Museum in Leiden, including Tom Gilissen, Marianne van der Wal and Agnes Bavelaar for all their help. Many thanks also to the Australian Museum and staff including Anja Divljan, Anina Hainsworth, Sandy Ingleby, Fiona Simpson and Fran Smith. Many thanks to Davide Molone who provided accommodation and interesting discussions during one of the visits to the Natural History Museum in London. Many thanks to Lindell Andrews and Dawn Roberts for providing several important references. Enormous thanks also go to the extraordinary Biodiversity Heritage Library that has allowed so many of the books and journals to be made available. Many thanks to Susan Bell (American Museum of Natural History) for answering various queries and providing several difficult-to-find citations. Ken Aplin provided important information on the taxonomy of various groups, which has been very much appreciated. The University of New South Wales and Professors Mike Archer and Sue Hand are gratefully acknowledged for their all their assistance. Finally, Stephen Jackson sincerely thanks Kerstin, Olivia and James for all their support during the writing of this book.

Definitions of nomenclatural and bibliographic terms

Absolute tautonomy: The identical spelling of a generic or subgeneric name and the specific or subspecific name of one of its originally included species or subspecies.

Abtheilung (Abth.): Part (German).

Available: Available to be used in nomenclature.

Available name: A scientific name that satisfies the nomenclatural requirements set forth in the International Code of Zoological Nomenclature, International Commission on Zoological Nomenclature (the current Code is the 4th edition, 1999; see ICZN in References). An available name may or may not be the valid name for a taxon.

Band (Bd.): Volume (German).

Berolini: of Berlin, Germany (Latin).

Binominal (binomial) name: The combination of two names, the first being the generic name and the second being the specific name, that together constitute the scientific name of a species. Any interpolated name (i.e. subgeneric name) is not counted as a component of a binominal name.

The Code: International Code of Zoological Nomenclature. First edition, 1961; 2nd edition, 1964; 3rd edition, 1985a; 4th edition, 1999.

Combination (or name combination): The association of a generic name and a specific name to form the name of a species, or of both with a subspecific name to form the name of the subspecies. In synonymy, new combinations do not constitute new names as such.

The Commission: International Commission on Zoological Nomenclature. Executive Secretary, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom; email: [email protected].

Conditionally proposed names (or nomenclatural acts): Conditional proposal of a name means that the author used language to the effect that if the subject specimen or sample proved in the future to represent a different taxon, it should be known as (Scientific name). Names or nomenclatural acts proposed conditionally after 1960 are not thereby made available (ICZN, 1999: Art. 15.1).

Definition: A statement in words that purports to give those characters that, in combination, uniquely distinguishes a taxon.

Description: A statement in words of taxonomic characters of a specimen or a taxon. A description of a nominal taxon when it is established is called an original description.

Diagnosis: A statement in words that purports to give those characters that differentiates the taxon from other taxa with which it is likely to be confused.

Ed./Eds.: editor/editors

Editio: Edition (Latin).

Edn.: Edition

Emendation (emend.): Any intentional change in the original spelling of an available name that is not mandated by the Code.

emend. pro: emendation for [a particular name].

errore pro: inadvertent error for [a particular name].

extralimital: Of an occurrence or a taxon, beyond the geographic limits of this work.

fide: According to.

First reviser: The 1st author to subsequently cite names (including different original spellings of the same name) or nomenclatural acts published on the same date and to select one of them to have precedence over the other(s).

First use of a name: The oldest use of a name.

Genus group: In the hierarchy of classification the group of taxa ranked between the family group and the species group. Contains taxa at the ranks of genus and subgenus. Holmiae: Stockholm, Sweden (Latin).

Haplotype: A combination of alleles that are closely linked on the same chromosome (or in mtDNA), and are [almost invariably] inherited together.

Holotype: A single specimen designated as the name-bearing type of a species or subspecies when it is established, or the single specimen on which such a taxon was based when no type was specified.

Homonym: Each of two or more available names having the same spelling, or differing only in suffix, and denoting different nominal taxa, whether in the family group, the genus group, or the species group. Junior homonyms are unavailable in nomenclature. Homonyms in the species group apply only within the same genus, and may be either primary or secondary; homonyms in the genus and family groups apply throughout the animal kingdom. Variant spellings of what is essentially the same word are deemed to be identical (ICZN, 1999: Art. 58).

Homonymy: The relationship between homonyms; that is to say, the situation when one name is a homonym of another.

ICZN: International Commission on Zoological Nomenclature (see the Commission).

Incertae sedis: Of uncertain taxonomic position.

Incorrect original spelling: An original spelling that is incorrect because of a misspelling or printer’s error, one of two or more different spellings of the same name in the original description, or a spelling requiring a mandatory change.

Incorrect subsequent spelling: Any change in the spelling of an available name other than a mandatory change or emendation (ICZN, 1999: Art. 33.3). An inadvertent alteration of an available name. Incorrect subsequent spellings are not available names unless an author purposely uses the misspelling for a previously undescribed taxon.

Indication: A reference to a description, definition, or figure in another publication, including pre-Linnaean (pre-1758) works. Using a reference in place of a description was a common practice in the late 18th century and during the 19th century, but is not permissible in literature after 1930.

Invalid name: An available name that is not in current use (i.e. for any recognised taxon).

Junior: A more recent name, comparative to previous use or older name.

Junior objective synonym: Of two synonyms, the one that was established at a later date.

Justified emendation: The correction of an incorrect original spelling.

Lapsus calami (pl. lapsus calamarum): ‘A slip of the pen’; when an author uses a name other than the name intended as opposed to a misspelling, typographical error, or printer’s error.

Lectotype: A syntype designated as the single name-bearing type specimen subsequent to the establishment of a nominal species or subspecies. If the original description of a species group name was based on two or more specimens, and no specimen was identified as the holotype, these specimens are syntypes (previously referred to as cotypes). When one of these syntypes is selected as the name-bearing type specimen, it is referred to as the lectotype.

Lieferung (Lfg.): Fascicle (German).

Linnaean tautonomy: The identical spelling of a new generic or subgeneric name established before 1931 and a pre-1758, one-word name cited as a synonym of one of the species or subspecies originally included in that genus.

Livr.: Livraison. Part of a serial issued from time to time.

Logotype: The type by subsequent determination. The historical type of a genus, selected from two or more original species. A genus whose type is selected from two or more original species is logotypic (see O. Cook, 1914: 314).

Lugduni: Lyons, France (Latin).

Mandatory change: A change in the original spelling mandated by the Code: (1) a change in the ending of a specific or subspecific name (such as agreement in gender with the generic name or in number such as a change from -ii to -orum when the species-group name was intended to honour two or more individuals of the same name instead of a single individual); or (2) the dropping of hyphens or diacritical marks such as accents, the separation of diphthongs (e.g. æ or oe), or a change of spelling required (for names published before 1985) such as converting -ü - to -ue- (e.g. mülleri to muelleri).

Monotypic: Represented by a single taxon. A taxon that includes only one subordinate taxon of the next lower rank.

Monotypy: A situation where: (1) the author does not explicitly indicate a type species for a genus or subgenus but lists a single species by an available name (type species by monotypy), or (2) when a species description is based on a single specimen not explicitly designated the holotype (holotype by monotypy). This is not to be confused with the term ‘monotypic’, a term in taxonomy denoting that a genus has only one species, or a species is not divided into subspecies.

Multiple original spellings: Two or more different original spellings for the same name (ICZN 1999: Art. 32.2.1).

Nec: Preoccupied by.

Neotype: The single specimen designated as the name-bearing type of a nominal species or subspecies when a need arises to objectively define the nominal taxon and no name-bearing type is believed to exist.

New combination: The first combination of a generic name and a previously established species-group name.

Nomenclatural act: Any published act that affects the status of a scientific name or its type. Description of a taxon, revision of a species, designation of a type species, restriction of a type locality, designation of a neotype, selection of a lectotype, and so on, are examples of a nomenclatural act.

Nomenclature: A system of names and provisions for their formation and use. Binomial nomenclature is the system of nomenclature whereby species, but no other taxon, are denoted by a binomen, a combination of two names, the first being the generic name and the second the specific name. Nomenclature is a man-made system, designed to serve taxonomy, which is a reflection of the natural world. The two should not be confused.

Nomen dubium (pl. nomina dubia): A Latin term meaning a name of unknown or doubtful application.

Nomen novum (pl. nomina nova): A Latin term used for a new replacement name. A new name that is published to replace an earlier name (and valid only if the latter is preoccupied) and which is expressively proposed as a replacement name; a new name, not to be confused with a new species, or a new genus, etc., which represent new taxa. Commonly applied to names proposed to replace junior homonyms.

Nomen novum pro: New name for [some other name, either a homonym or for some reason ‘preferred’, this being in the older literature].

Nomen nudum (nom. nud): A naked name, a name that has not met the criteria for availability as outlined in the International Code of Zoological Nomenclature; a name that, if published before 1931, was not accompanied by a description, definition, or indication, or if published after 1930, is not accompanied by a statement that purports to give characters differentiating the taxon; or is not accompanied by a definitive bibliographic reference to such a statement; or is not proposed expressly as a replacement for a pre-existing name. A nomen nudum is not an available name, and therefore the same name may be made available later for the same or a different concept; in such a case it would take authorship and date from that act of establishment, not from any earlier publication as a nomen nudem.

Nomen oblitum (pl. nomina oblita): A Latin term meaning a forgotten name. A name that has not been used since 1899; such a name is not allowed to displace a junior synonym or homonym in prevailing usage.

Nominal taxon: A taxon denoted by an available name.

Nominotypical: The nominal taxon at a subordinate rank within the family group, the genus group, or the species group that contains the name-bearing type of that group.

Non: Not.

nov. = novus, novum: New.

Objective synonym: A name whose synonymy with another is not merely a matter of opinion, because the two are based on the same type material.

Original designation: The designation of the type of a nominal taxon when it is established.

Original description: See Description.

Operational Taxonomic Unit: A group, or sample, or single specimen, for the time being assumed to be a valid taxon for purposes of an analysis.

Pace: ‘With peace’, meaning with all due respect.

Polytypic: A taxon represented by two or more subordinate taxa of the next lower rank. This is not to be confused with its use in taxonomy, referring to a taxon which has two or more subordinate taxa.

Preoccupied name: A name that is unavailable because it is: (1) a generic name of the same spelling (junior homonym) as a generic name previously used for a different animal; or (2) a species or subspecies name of the same spelling originally combined with the same generic name as an earlier described name, even if that species-level name is now used with a different genus.

Preoccupied: A homonym.

Primary homonym: Each of two or more homonyms in the species group, originally combined with the same generic name.

Principle of Priority: The principle that the valid name of a taxon is the oldest available name applied to it provided that the name is not invalidated by any provision of the Code or by any ruling by the Commission.

Pro: Before.

Recte: Correctly.

Renaming: The act of providing a replacement name for a preoccupied name. Note that a different name combination does not constitute a renaming.

Scientific name: A name treated as a Latinised name for a taxon (usually cited in combination with the author and date of publication).

Secondary homonym: Each of two or more homonyms in the species group, originally combined with different generic names but subsequently combined with the same generic name.

Senior: An older name, comparative with a more recent name

Senior synonym: Of two synonyms, the one that was established at an earlier date

Sensu: In the sense of.

Sensu lato: In the broad sense.

Sensu stricto: in the strict (narrow) sense.

Separate: A copy (reprint or offprint) of an article printed separately from the journal or periodical in which the article appears. If separates are printed and distributed before the printing and distribution of the journal or periodical, they are referred to as preprints and usually bear an earlier date and different pagination. Today scientific publications normally print separates (usually called reprints) after the printing and distribution of the book, journal, or periodical in which the article appears. Separates qualifying as reprints have the same pagination found in the publication in which they appear. The advance distribution of separates after 1999 does not constitute publication for the purposes of zoological nomenclature.

Species group: In zoological classification, the lowest-ranking group of taxa, the names of which are regulated by the Code. The species group includes all taxa at the ranks of species and subspecies.

Subjective synonym: A name based on a type specimen thought to belong to the same taxon as another name.

Subsequent designation: Designation of type species for a genus group taxon in another publication after the genus-group taxon was described.

Subsequent spelling: A subsequent spelling that is not the same as the original spelling is an emendation, an incorrect subsequent spelling, or a mandated change.

Syntype: Each specimen of a type series from which a holotype (at time of publication) or a lectotype (subsequent to publication) has not been selected. The syntypes collectively constitute the name-bearing type. Syntypes are sometimes referred to as cotypes, a term that should not now be used in zoological nomenclature.

Tautonymy: The use of the same word for the name of a genus and of one of its originally included nominal species or subspecies. Linnean tautonomy is the identical spelling of a new generic or subgeneric name and pre-1758 name cited as a synonym of only one of the species or subspecies originally included in that genus.

Tautotype: The type of a genus designated because its [species-group] name is identical in spelling with the genus-group name.

Taxon (pl. taxa): A taxonomic unit, whether named or not, considered to comprise a population or group of populations of organisms usually inferred to be phylogenetically related and have characters in common that differentiate the unit from other such units. A taxon encompassing all included taxa of lower rank.

Taxonomy: The theoretical study of classification, including its bases, principles, procedures, and rules. It includes the classification and naming of animals.

Tome: Volume (French).

Tomus: Volume (Latin).

Type: A tern used alone, or forming part of a compound term, to denote a particular kind of specimen or taxon.

Type locality: The geographical place of capture or collection of the name-bearing type of a nominal species or subspecies.

Typotypical: [of a specimen] originating from the type locality of the species or subspecies to which it is thought to belong.

Unavailable: Not available for use in nomenclature.

Unavailable work: A publication in which names and nomenclatural acts are rejected for nomenclatural purposes. The work may be unavailable because it was published before 1758, or because the author was not consistently binomial, or (after 1950) was published anonymously, or contained a disclaimer, or because the Commission has ruled it to be unavailable.

Unjustified emendation: An intentional alteration of the original spelling of an available name that is not mandated.

Valid name: An available name that is used as the current name for a taxon. A valid name is a term applied in nomenclature to mean only the name by which a taxon is currently identified.

Vernacular name: A name of an animal or animals in a language used for general purposes as opposed to a Latinised name proposed only for zoological nomenclature.

Vide: See.

Virtual tautonomy: The nearly identical spelling, or of the same origin or meaning, of a generic or subgeneric name and the specific or subspecific name in a binomen or trinomen. The term virtual tautonomy is not regulated by the Code.

Abbreviations

†:   Extinct.

Φ:  Extralimital taxon.

Ω:  Introduced taxon.

Taxonomy of the mammals of Australia

CLASS MAMMALIA Linnaeus, 1758

SUBCLASS PROTOTHERIA Gill, 1872

ORDER MONOTREMATA Bonaparte, 1832 sensu Bonaparte, 1838

Family Ornithorhynchidae J. Gray, 1825 sensu Burnett, 1830 [Platypus]

Ornithorhynchus anatinus (Shaw, 1799) Platypus

Family Tachyglossidae Gill, 1872 [Echidnas]

Tachyglossus aculeatus (Shaw, 1792) Short-beaked Echidna

† Zaglossus bruijni (Peters & Doria, 1876) Western Long-beaked Echidna

SUBCLASS THERIA Parker & Haswell, 1897

SUPERLEGION TRECHNOTHERIA McKenna, 1975

LEGION YANGOTHERIA Chow & Rich, 1982

SUBLEGION CLADOTHERIA McKenna, 1975

INFRALEGION ZATHERIA McKenna, 1975

INFRACLASS TRIBOSPHENIDA McKenna, 1975

SUPERCOHORT MARSUPIALIA IIliger, 1811 sensu Cuvier, 1816

COHORT AUSTRALIDELPHIA Szalay, 1982

ORDER DASYUROMORPHIA Gill, 1872 sensu Aplin & Archer, 1987

Superfamily Dasyuroidea Goldfuss, 1820 sensu Marshall et al., 1990

Family Dasyuridae Goldfuss, 1820 sensu Owen, 1839 [Dasyurids]

Subfamily Dasyurinae Goldfuss, 1820 sensu Marshall et al., 1990

Dasycercus blythi (Waite, 1904) Brush-tailed Mulgara

Dasycercus cristicauda (Krefft, 1867) Crest-tailed Mulgara

Dasykaluta rosamondae (Ride, 1964) Kaluta

Dasyuroides byrnei Spencer, 1896 Kowari

Dasyurus geoffroii Gould, 1841 Western Quoll

Dasyurus hallucatus Gould, 1842 Northern Quoll

Dasyurus maculatus (Kerr, 1792) Spotted-tailed Quoll [Tiger Quoll]

Dasyurus viverrinus (Shaw, 1800) Eastern Quoll

Parantechinus apicalis (J. Gray, 1842) Dibbler

Pseudantechinus bilarni (Johnson, 1954) Sandstone Pseudantechinus

Pseudantechinus macdonnellensis (Spencer, 1895) Fat-tailed Pseudantechinus

Pseudantechinus mimulus (Thomas, 1906) Carpentarian Pseudantechinus

Pseudantechinus ningbing Kitchener, 1988 Ningbing Pseudantechinus

Pseudantechinus roryi N. Cooper et al., 2000 Tan Pseudantechinus

Pseudantechinus woolleyae Kitchener & Caputi, 1988 Woolley’s Pseudantechinus

Sarcophilus harrisii (Boitard, 1841) Tasmanian Devil

Subfamily Phascogalinae Gill, 1872 sensu Marshall, 1990

Antechinus adustus (Thomas, 1923) Rusty Antechinus

Antechinus agilis Dickman et al., 1998 Agile Antechinus

Antechinus argentus Baker et al., 2013 Silver-headed Antechinus

Antechinus arktos Baker et al., 2014 Black-tailed Antechinus

Antechinus bellus (Thomas, 1904) Fawn Antechinus

Antechinus flavipes (Waterhouse, 1838) Yellow-footed Antechinus

Antechinus godmani (Thomas, 1923) Atherton Antechinus

Antechinus leo Van Dyck, 1980 Cinnamon Antechinus

Antechinus minimus (É. Geoffroy, 1803) Swamp Antechinus

Antechinus mysticus Baker et al., 2012 Buff-footed Antechinus

Antechinus stuartii Macleay, 1841 Brown Antechinus

Antechinus subtropicus Van Dyck & Crowther, 2000 Subtropical Antechinus

Antechinus swainsonii (Waterhouse, 1840) Dusky Antechinus

Phascogale calura Gould, 1844 Red-tailed Phascogale

Phascogale pirata Thomas, 1904 Northern Phascogale

Phascogale tapoatafa (Meyer, 1793) Brush-tailed Phascogale

Subfamily Planigalinae Archer, 1982 sensu Marshall et al., 1990

Planigale gilesi Aitken, 1972 Gile’s Planigale

Planigale ingrami (Thomas, 1906) Long-tailed Planigale

Planigale maculata (Gould, 1851) Common Planigale

Planigale tenuirostris Troughton, 1928 Narrow-nosed Planigale

Subfamily Sminthopsinae Archer, 1982 sensu Marshall et al., 1990

Antechinomys laniger (Gould, 1856) Kultarr

Ningaui ridei Archer, 1975 Wongai Ningaui

Ningaui timealeyi Archer, 1975 Pilbara Ningaui

Ningaui yvonnae Kitchener et al., 1983 Southern Ningaui

Sminthopsis archeri Van Dyck, 1986 Chestnut Dunnart

Sminthopsis bindi Van Dyck et al., 1994 Kakadu Dunnart

Sminthopsis butleri Archer, 1979 Butler’s Dunnart

Sminthopsis crassicaudata (Gould, 1844) Fat-tailed Dunnart

Sminthopsis dolichura Kitchener, et al., 1984 Little Long-tailed Dunnart

Sminthopsis douglasi Archer, 1979 Julia Creek Dunnart

Sminthopsis fuliginosus (Gould, 1852) Grey-bellied Dunnart

Sminthopsis gilberti Kitchener et al., 1984 Gilbert’s Dunnart

Sminthopsis granulipes Troughton, 1932 White-tailed Dunnart

Sminthopsis hirtipes Thomas, 1898 Greater Hairy-footed Dunnart

Sminthopsis leucopus (J. Gray, 1842) White-footed Dunnart

Sminthopsis longicaudata Spencer, 1909 Large Long-tailed Dunnart

Sminthopsis macroura (Gould, 1845) Stripe-faced Dunnart

Sminthopsis murina (Waterhouse, 1838) Common Dunnart

Sminthopsis ooldea Troughton, 1965 Ooldea Dunnart

Sminthopsis psammophila Spencer, 1895 Sandhill Dunnart

Sminthopsis virginiae (de Tarragon, 1847) Red-cheeked Dunnart

Sminthopsis youngsoni McKenzie & Archer, 1982 Lesser Hairy-footed Dunnart

Family Myrmecobiidae Waterhouse, 1841 [Numbat]

Myrmecobius fasciatus Waterhouse, 1836 Numbat

Family Thylacinidae Bonaparte, 1838 [Thylacine]

† Thylacinus cynocephalus (Harris, 1808) Thylacine

ORDER NOTORYCTEMORPHIA Kirsch, 1977 sensu Aplin & Archer, 1987

Family Notoryctidae J. Ogilby, 1892 [Marsupial Moles]

Notoryctes caurinus Thomas, 1920 Northern Marsupial Mole

Notoryctes typhlops (Stirling, 1889) Southern Marsupial Mole

ORDER PERAMELEMORPHIA Ameghino, 1889 sensu Aplin & Archer, 1987

Superfamily Perameloidea J. Gray, 1825 sensu Van Dyck & Strahan, 2008

Family Chaeropodidae Gill, 1872 sensu Groves, 2005 [Pig-footed Bandicoot]

† Chaeropus ecaudatus (W. Ogilby, 1838) Pig-footed Bandicoot

Family Peramelidae J. Gray, 1825 sensu Van Dyck & Strahan, 2008 [Bandicoots]

Subfamily Echymiperinae McKenna & Bell, 1997 sensu Van Dyck & Strahan, 2008

Echymipera rufescens (Peters & Doria, 1875) Long-nosed Echymipera

Subfamily Peramelinae J. Gray, 1825 sensu Kirsch et al., 1997

Isoodon auratus (Ramsay, 1887) Golden Bandicoot

Isoodon macrourus (Gould, 1842) Northern Brown Bandicoot

Isoodon obesulus (Shaw, 1797) Southern Brown Bandicoot

Isoodon peninsulae Thomas, 1922 Cape York Brown Bandicoot

Perameles bougainville Quoy & Gaimard, 1824 Western Barred Bandicoot

† Perameles eremiana Spencer, 1897 Desert Bandicoot

Perameles gunnii J. Gray, 1838 Eastern Barred Bandicoot

Perameles nasuta É. Geoffroy, 1804 Southern Long-nosed Bandicoot

Perameles pallescens Thomas, 1923 Northern Long-nosed Bandicoot

Family Thylacomyidae Bensley, 1903 sensu Archer & Kirsch, 1977 [Bilbies]

Macrotis lagotis (Reid, 1837) Greater Bilby

† Macrotis leucura (Thomas, 1887) Lesser Bilby

ORDER DIPROTODONTIA Owen, 1877

SUBORDER VOMBATIFORMES Woodburne, 1984 sensu Aplin & Archer, 1987

INFRAORDER PHASCOLARCTOMORPHIA Aplin & Archer, 1987

Family Phascolarctidae Owen, 1839 [Koala]

Phascolarctos cinereus (Goldfuss, 1817) Koala

INFRAORDER VOMBATOMORPHIA Aplin & Archer, 1987

Family Vombatidae Burnett, 1830 sensu Dawson, 1983 [Wombats]

Lasiorhinus krefftii (Owen, 1872) Northern Hairy-nosed Wombat

Lasiorhinus latifrons (Owen, 1845) Southern Hairy-nosed Wombat

Vombatus ursinus (Shaw, 1800) Bare-nosed Wombat

SUBORDER PHALANGERIDA Aplin & Archer, 1987

Superfamily Burramyoidea Broom, 1898 sensu Aplin & Archer, 1987

Family Burramyidae Broom, 1898 sensu Aplin & Archer, 1987 [Pygmy-possums]

Burramys parvus Broom, 1895 Mountain Pygmy-possum

Cercartetus caudatus (Milne-Edwards, 1877) Long-tailed Pygmy-possum

Cercartetus concinnus (Gould, 1845) Western Pygmy-possum

Cercartetus lepidus (Thomas, 1888) Little Pygmy-possum

Cercartetus nanus (Desmarest, 1817) Eastern Pygmy-possum

Superfamily Petauroidea Bonaparte, 1832 sensu Aplin & Archer, 1987

Family Petauridae Bonaparte, 1832 sensu Baverstock, 1984 [Striped Possum, Leadbeater’s Possum, and Lesser Gliders]

Subfamily Dactylopsilinae Kirsch, 1977 sensu Edwards & Westerman, 1992

Dactylopsila trivirgata J. Gray, 1858 Torresian Striped Possum

Gymnobelideus leadbeateri McCoy, 1867 Leadbeater’s Possum

Subfamily