The biogeographical history of non-marine mammaliaforms in the Sahul region

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Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

THE BIOGEOGRAPHICAL HISTORY OF NON-MARINE MAMMALIAFORMS IN
THE SAHUL REGION
Robin M. D. Beck
School of Environment & Life Sciences, University of Salford, Salford, M5 4WT, UK
School of Biological, Earth and Environmental Sciences, University of New South Wales,
NSW 2052, Australia
Email: r.m.d.beck@salford.ac.uk
1 Introduction
Today, the biogeographical region comprising Australia, New Guinea and adjacent islands is
the only part of the globe where representatives of all three major extant mammalian clades
occur together, namely monotremes, marsupials and placentals (Wilson and Reeder 2005;
Van Dyck and Strahan 2008; Flannery 1995b). Monotremes (five species) are currently found
nowhere else, whilst >240 described marsupial species comprise ~40% of the total terrestrial
mammal diversity in the region. Placentals dominate the mammal faunas of most continental
landmasses, but in Australia and New Guinea only two placental clades have achieved
moderate diversity: murine rodents (>160 species; ~25% of the total) and bats (>130 species;
~20% of the total). Most other non-marine placental clades seem to have been entirely absent
from Australia and New Guinea prior to human-related introductions during the Holocene.
This unique overall pattern of mammalian biodiversity, so different from that seen elsewhere

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

in the world, has fascinated a long line of researchers, including Darwin and Wallace
themselves.
Evidence from the fossil record reveals further striking patterns and complexities. For
example, Metatheria (the clade which includes modern marsupials) probably originated in the
northern hemisphere by the middle Cretaceous at the latest (~125 Ma; Luo et al. 2003), but
metatherians appear to have been absent from the southern continents, including Australia,
until the latest Cretaceous or earliest Palaeocene, some 40-60 Myr later (Case et al. 2005;
Pascual and Ortiz-Jaureguizar 2007; Kielan-Jaworowska et al. 2004; Woodburne et al. 2014;
Rougier et al. 2011b; Goin et al. 2012b). Prior to the late Oligocene, the fossil record of
mammals and their extinct relatives (collectively, mammaliaforms) in Australia is very poor,
but it includes high-level taxa that are known nowhere else, such as the middle Cretaceous
ausktribosphenids and Kollikodon, and biogeographical enigmas such as the putative
eutherian “condylarth” Tingamarra from the early Eocene (Long et al. 2002; Archer et al.
1999a). The richer Oligo-Miocene record of Australia is dominated by marsupials, including
members of most living families, with monotremes and representatives of multiple bat
families also present (Archer et al. 1999a; Long et al. 2002; Black et al. 2012). However, the
oldest murine rodent fossils from Australia are only ~4 Ma, and the oldest from New Guinea
only 3-3.5 Ma (Godthelp 1999; Aplin 2006; Aplin and Ford 2014), several Myr younger than
the origin of Murinae as a whole (Jacobs and Flynn 2005; Fabre et al. 2013; Schenk et al.
2013).
This fossil evidence, when combined with phylogenies, divergence dates estimated
from molecular and stratigraphic data, and geological information, gives insight into the
biogeographical history of Australian and New Guinean mammaliaforms, and provides clues
as to how the current mammal fauna of the region developed. At finer taxonomic scales,
phylogeographic studies of molecular data are beginning to reveal the roles that

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

environmental change and putative barriers to gene flow have played in shaping the
biogeography of modern species and populations. Nevertheless, major gaps in our current
knowledge - due to factors such as the highly incomplete fossil record of mammaliaforms in
the region, uncertainty regarding the phylogeny and even alpha taxonomy of many taxa, and
a lack of detailed, quantitative biogeographic analyses – mean that numerous uncertainties
remain.
This chapter represents an attempted synthesis of our current knowledge of terrestrial
mammaliaform biogeography in Australia, New Guinea and adjacent islands (specifically, the
region east of Lydekker’s Line; Figure 1d), integrating available fossil, phylogenetic and
geological data. My major focus is above the species level, with extensive discussion of the
fossil record, but I also briefly discuss phylogeographic studies of modern species. I end with
a short summary of the biggest lacunae in our current knowledge, and the prospects for
improving our understanding of the biogeographical history of mammaliaforms in what is
undoubtedly one of the most fascinating and remarkable regions on Earth.
2 Phylogenetic definitions and scope
Table 1 lists the formal phylogenetic definitions of selected clades discussed in this review. I
follow most recent studies (e.g. Meredith et al. 2011; O'Leary et al. 2013; Bi et al. 2014) in
restricting the name Mammalia to the crown-clade only. I use Mammaliaformes to refer to
the more inclusive synapsid clade corresponding to “traditional” definitions of Mammalia
(e.g. Kielan-Jaworowska et al. 2004), following the stem-based definition of Sereno (2006:
table 10.1). Most fossil mammaliaforms found in Sahul to date appear to be members of
Mammalia (Long et al. 2002), but a few (e.g. Kollikodon) may fall outside the crown-clade
(see below).

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Within Mammalia, I follow a crown-clade definition of Theria, that is to say the clade
circumscribed by Marsupialia and Placentalia plus all other fossil taxa within that clade
(Sereno 2006; O'Leary et al. 2013). As in most recent studies (e.g. O'Leary et al. 2013;
Sereno 2006; Vullo et al. 2009), I restrict Marsupialia and Placentalia to the crown-clades,
with Metatheria and Eutheria referring to their respective total clades. Vullo et al. (2009)
proposed the name Marsupialiformes to correspond to the “traditional”, more inclusive
definition of Marsupialia (e.g. Kielan-Jaworowska et al. 2004). Vullo et al. (2009: 19910)
gave an approximate definition of Marsupialiformes: “crown group Marsupialia (extant
marsupials and related extinct fossil taxa) plus all stem marsupialiform taxa that are more
closely related to them, as their sister taxa, than to Deltatheroida and basal Metatheria”.
However, I propose a new, less ambiguous definition here (Table 1). It is unclear whether
only (crown-clade) marsupials reached Sahul, or whether stem-marsupialiforms were also
present (Sigé et al. 2009; Beck 2014). Fossil metatherians from Sahul and elsewhere in
Gondwana that cannot be definitively placed within the crown-clade will therefore be
referred to as marsupialiforms here.
Several clades within Marsupialia are of particular biogeographical relevance, and it
is appropriate to discuss their phylogenetic definitions. The clade Australidelphia was
originally named by Szalay (1982) to include the four Sahulian marsupial orders –
Dasyuromorphia (predominantly carnivorous forms such as quolls, dunnarts, the numbat and
the thylacine), Diprotodontia (“possums”, kangaroos, wombats and the koala),
Notoryctemorphia (marsupial moles) and Peramelemorphia (bandicoots and bilbies) - plus
Microbiotheria (represented today by a single species, the South American “monito del
monte”, Dromiciops gliroides). However, here I follow the apomorphy-based definition for
Australidelphia that I proposed previously (Beck 2012: 717; Table 1). I use the name “crown-
clade Australidelphia” to refer to the clade circumscribed by the five extant australidelphian

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

orders, and propose a formal, node-based phylogenetic definition for this clade (Table 1).
Within crown-clade Australidelphia, the four Sahulian orders appear to form a clade to the
exclusion of Microbiotheria (Beck 2008a; 2011; 2009a; Meredith et al. 2009b; Nilsson et al.
2010), which Archer (1984) named Eomarsupialia. I follow Beck et al.’s (2014, : 132) crown-
based definition for Eomarsupialia.
Marine mammals will not be discussed in this review, because the factors influencing
their biogeographical distributions are obviously very different to those affecting non-marine
forms. Recent reviews discussing the biogeography of marine mammals that occur in the
waters around Sahul include Deméré et al. (2003) on pinnipedimorphs, Fordyce (2006) on
cetaceans, and de Iongh and Domning (2014) on sirenians. I also do not cover the arrival of
humans in the region, nor the various other placental species they introduced; recent
discussions of these topics can be found in, inter alia, Johnson (2006), Helgen (2007),
Davidson (2014), Dennell and Porr (2014) and Prins and Gordon (2014).
3 Geographical and geological context
From a geographical perspective, this review focuses on continental and oceanic landmasses
of the Sahul shelf, delimited by Lydekker’s Line to the west and the Pacific oceanic plate to
the east, including the mainland and adjacent islands of Australia and New Guinea (Figure
1d; Lydekker 1896). However, it should be recognised that this boundary is (like other faunal
lines in the region; Simpson 1977) somewhat arbitrary and that numerous typically
“Sahulian” mammals (e.g. marsupials, “Old Endemic” and “New Endemic” murines) occur
on islands west of Lydekker’s line (Flannery 1995a). I refer to the Australian mainland plus
Tasmania as “Australia”, and Australia plus New Guinea and adjacent islands as “Sahul”. As
discussed below, the major period of uplift of New Guinea does not appear to have occurred

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

until the late Miocene; thus, I refer to Australia only when discussing biogeographical and
geological events prior to this.
An understanding of the geological history of Sahul is key to interpreting the
biogeographical history of the species inhabiting it. The following is a brief review of major
global tectonic events occurring over the known timeframe of mammaliaform evolution
(from the Late Triassic onwards; Kielan-Jaworowska et al. 2004), focusing on those directly
affecting the geological evolution of the region.
The major phase of the break-up of Pangaea commenced with the opening of the
Central Atlantic Ocean ~180-195 Ma, although complete separation of Laurasia and
Gondwana (with the development of a continuous Tethyan Seaway between the two
supercontinents) may not have occurred until the Early Cretaceous (Seton et al. 2012; Torsvik
and Cocks 2013; Lomolino et al. 2010). West Gondwana (what would become South
America and Africa) and East Gondwana (what would become Antarctica, India,
Madagascar, Australia and New Zealand) began to separate ~140-170 Ma, but remained in
contact at their southern ends (Seton et al. 2012; Torsvik and Cocks 2013; Lomolino et al.
2010; Chatterjee et al. 2013); the southern tip of South America and the Antarctic Peninsula
did not separate fully, via opening of the Drake Passage, until well into the Cenozoic (Figure
2; see below). The first major landmass to break away from Gondwana appears to have been
Indo-Madagascar, ~130 Ma (Chatterjee et al. 2013; Seton et al. 2012; Lomolino et al. 2010).
Africa and South America began to separate ~120 Ma, with the separation largely complete
by ~100 Ma (Seton et al. 2012; Lomolino et al. 2010). New Zealand began to break away
from Australia ~80-90 Ma (Lomolino et al. 2010; Seton et al. 2012; Ericson et al. 2014), but
complete separation may not have been achieved until ~52 Ma (Ericson et al. 2014).

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Isotopic evidence indicates an influx of Pacific seawater into the Atlantic, across the
Drake Passage between the southern tip of South America and the Antarctic Peninsula, ~41
Ma (Scher and Martin 2006), although the Drake Passage may already have been open, but
shallow (<1000 m), as early as ~50 Ma (Livermore et al. 2005; Lawver et al. 2011; Eagles
and Jokat 2014). Deepwater opening of the Drake Passage may not have occurred until ~30
Ma (Eagles and Jokat 2014). Evidence from dinoflagellates and organic geological records
indicates a flow of water through the Tasmanian Gateway between Antarctica and Australia
~49-50 Ma (Bijl et al. 2013). This was followed by deepening of the Tasmanian Gateway
~35.5 Ma (Stickley et al. 2004), leading to the establishment of the Antarctic Circumpolar
Current. Terrestrial vertebrate dispersal between Australia and Antarctica would therefore
seem highly unlikely after 35.5 Ma, and so Australia can be reasonably considered an “island
continent” from this date onwards (Figure 2).
The geological history of New Guinea is complex and as yet incompletely
understood. However, it is now generally accepted that only small areas of land, if any, were
emergent north of the Australian continent until at least the middle-late Miocene (Figure 1a-
c), with the major period of enlargement of the New Guinean landmass occurring over the
last 5 Myr (Quarles van Ufford and Cloos 2005; Hall 2002; Hill and Hall 2003; Westerman et
al. 2012; Toussaint et al. 2014; Baldwin et al. 2012; Hocknull 2009; Cloos et al. 2005).
Particularly significant is that there is no geological evidence for a dry-land connection
between Australia and New Guinea during the Eocene and/or Oligocene, contra Flannery
(1988, : fig. 2; 1995b, : maps 3-4).
Large fluctuations in eustatic sea level over the last ~10 Myr, and particularly the last
~2.5 Myr (Miller et al. 2005; Haq et al. 1987), have resulted in major changes in extent of
exposed land on the Sahul Shelf (Figure 1d). This has led to the repeated formation and
severing of dry-land connections between New Guinea, mainland Australia and adjacent

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

islands over the last 3.5 Myr (Hocknull 2009; Coller 2009). The approximate extent and
duration of these connections can be visualised using Monash University’s “SahulTime”
webpage (http://sahultime.monash.edu.au/; Coller 2009). Of particular importance is that,
over the last ~1 Myr, Australia and New Guinea have more often formed a single landmass
than they have been separated by sea (Bintanja et al. 2005; Hocknull 2009; Coller 2009).
However, it is also crucial to note that, even at periods of lowest sea level (such as during the
Last Glacial Maximum), major marine barriers remained in place between Sahul and the land
masses west of Lydekker’s Line (Figure 1d); there has never been a dry-land connection
between Sahul and Wallacea, or between Wallacea and the landmasses of the Sunda Shelf
(Sundaland), further to the west (Lohman et al. 2011). Thus, dispersal to and from Sahul has
required the crossing of marine barriers since the deepwater opening of the Tasman Gateway
between Antarctica and Australia ~35.5 Ma (see above). This presumably explains why,
among mammals, only murine rodents and bats appear to have successfully dispersed to
Sahul from southeast Asia, despite the existence of highly diverse placental faunas in
Sundaland and the presence of a number of terrestrial placentals besides murines in Wallacea
(Dennell et al. 2014; Flannery 1995a). Marine barriers are obviously less formidable to volant
bats than to terrestrial mammals, and murines appear to be particularly adept at overwater
dispersal (van der Geer et al. 2010; Achmadi et al. 2013).
4 Biogeographical history of non-marine mammaliaform clades
4.1 ?Multituberculata
Multituberculates were conspicuous members of Late Jurassic, Cretaceous and early
Palaeogene mammal faunas of Laurasia (Kielan-Jaworowska et al. 2004; Rose 2006).
Putative Gondwanan multituberculates, by contrast, are very rare and fragmentary, and their

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

biogeographical interpretation correspondingly controversial. To date, a single probable
multituberculate has been described from Australia (Rich et al. 2009a): Corriebaatar
marywaltersae (Figure 3f), represented by a partial dentary preserving a “plagiaulacoid”
premolar from the early-middle Aptian (~115-125 Ma) Flat Rocks site in the Eumeralla
(=”Wonthaggi”) Formation, Strzelecki Group, southern Victoria (Figure 3a). Rich et al.
(2009a) tentatively referred C. marywaltersae to the multituberculate clade Cimolodonta, but
suggested that it might in fact represent a previously unknown mammaliaform lineage. A
third possibility not considered by Rich et al. (2009a) is that Corriebataar is a representative
of the gondwanatherian family Ferugliotheriidae (otherwise known only from the Late
Cretaceous and possibly early Palaeogene of South America; Goin et al. 2012a), as this group
has also been argued to be characterised by the presence of a plagiaulacoid lower premolar
(Gurovich and Beck 2009; but see Pascual et al. 1999; Pascual and Ortiz-Jaureguizar 2007).
Nevertheless, pending further analysis and/or the discovery of additional material, I will
follow Rich et al.’s (2009) preferred interpretation here, namely that Corriebataar is a
cimolodontan multituberculate.
The oldest well-dated multituberculates are from the late Bathonian (~166 Ma) of
England (Butler and Hooker 2005). However, Butler and Hooker (2005) argued for a much
earlier origin of Multituberculata, in the Early Jurassic or even earlier. If this is the case, then
the apparent presence of multituberculates in Gondwana (based on Corriebaatar and other
putative Gondwanan records; Krause 2013; Kielan-Jaworowska et al. 2007; Rich et al. 2009a;
Gurovich and Beck 2009; Parmar et al. 2013) could reflect an initial trans-Pangaean
distribution for Multituberculata and subsequent vicariance (see “Geographical and
geological context” above). Of particular relevance is Indobaatar zofiae from the Kota
Formation in India, which has been described as a probable eobataarid multituberculate
(Parmar et al. 2013). The age of the Kota Formation remains controversial, with estimates

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

ranging from the Early Jurassic to the Early Cretaceous (Parmar et al. 2013; Prasad and
Manhas 2007). However, if the Kota Formation is Early Jurassic, then Indobaatar suggests
that Multituberculata originated prior to Pangaean break-up, and so the presence of
Corriebaatar in the Aptian of Australia is plausibly the result of vicariance.
4.2 Monotremata
Two definitive fossil monotremes have been described from middle Cretaceous sites in
Australia: Steropodon galmani (Figure 3c) from the Albian Griman Creek Formation at
Lightning Ridge (Figure 3a) in northern New South Wales (Archer et al. 1985), and
Teinolophos trusleri (Figure 3e) from the early-middle Aptian Flat Rocks site (Figure 3a) in
southern Victoria (Rich et al. 1999; 2001b). Other putative records of monotremes from the
Cretaceous of Australia are more uncertain. The dentally bizarre Kollikodon ritchiei (Figure
3d), also from Lightning Ridge, was originally identified as a probable monotreme (Flannery
et al. 1995) but more recently has been suggested to be a non-mammalian mammaliaform
(Musser 2006; 2013). A “tachyglossid-like” partial right humerus from the Albian Dinosaur
Cove site (Figure 3a) in southern Victoria has been named Kryoryctes cadburyi, and
tentatively identified as a monotreme (Pridmore et al. 2005). However, given the lack of data
regarding the postcranial morphology of most Mesozoic mammal groups, this identification
should be viewed with caution. Additional specimens from Lightning Ridge, including a
number of edentulous dentaries, have been suggested to represent monotremes (Musser 2013;
Smith 2009), but these have yet to be described in detail. A fossil from Lightning Ridge,
which Rich et al. (1989) suggested might be a maxilla of Steropodon galmani, has more
recently been identified as turtle (Smith 2009).

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

There is currently no Mesozoic record of monotremes outside Australia (Kielan-
Jaworowska et al. 2004; Rich 2008), and it is tempting to interpret this as evidence of a
restricted biogeographical distribution for the group at this time, perhaps as part of a larger
eastern Gondwanan radiation within Australosphenida (Figure 3b; see “Ausktribosphenidae”
below). Additional evidence that the terrestrial vertebrate fauna of Australia (or eastern
Gondwana more generally) may have been biogeographically isolated or relictual during at
least the middle Cretaceous include the temnospondyl Koolasuchus cleelandi from the early-
middle Aptian Eumeralla Formation of Victoria (Warren et al. 1997), and a putative
dicynodont that appears to be from the Aptian Allaru Formation in central Queensland
(Thulborn and Turner 2003), although the exact geological provenance of the latter specimen
is uncertain and its identification as a dicynodont has been questioned (Agnolin et al. 2010).
Outside Australia, the youngest known temnospondyls are from the Jurassic (Schoch 2014),
and the youngest known dicynodonts are from the late Triassic (Fröbisch 2009). Rich et al.
(2009a) also summarised evidence suggesting a clear distinction between the Early
Cretaceous palaeofloras of South America on the one hand and those of Antarctica and
Australia on the other, perhaps reflecting a climate-related filter acting through the Antarctic
Peninsula. However, the exact biogeographical significance of other terrestrial vertebrates -
such as turtles, crocodylomorphs and dinosaurs - from Australian Mesozoic sites is
contentious, in part because most are known from highly fragmentary specimens (see
summary by Poropat et al. 2014). Some authors have recognised clear evidence of climate-
driven provinciality in the middle Cretaceous terrestrial vertebrate faunas of Australia (e.g.
Benson et al. 2012), whereas others have argued that they show close links to those from
elsewhere in Gondwana, particularly South America, implying extensive faunal exchange
during the middle Cretaceous (e.g. Agnolin et al. 2010).

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Focusing specifically on mammaliaforms, direct comparison of the Australian record
with the record elsewhere in Gondwana faces two major difficulties. Firstly, very few
Mesozoic mammaliaform-bearing sites are currently known throughout Gondwana, and only
a small number of specimens have been obtained from them to date (Kielan-Jaworowska et
al. 2004; Rich 2008). Secondly, Gondwanan sites exhibit a disjunct temporal distribution
(Kielan-Jaworowska et al. 2004; Rich 2008): the Australian sites (Figure 3a) are all Aptian-
Albian in age, whereas those from elsewhere in Gondwana are either much older, much
younger, or of uncertain age. Thus, it is difficult to determine whether differences between
the Mesozoic mammaliaform faunas of Australia and those of other Gondwanan landmasses
reflect biogeographical factors or, alternatively, Gondwana-wide changes in faunal
composition through time. Bearing these difficulties in mind, perhaps of greatest significance
is the apparent absence of monotremes in the rich late Cretaceous “Allenian” (= Alamitian
South American Land Mammal Age [SALMA]) faunas of Patagonia (Rougier et al. 2009a;
Bonaparte 1990; Rougier et al. 2009b; Rougier et al. 2011b). Rich et al. (2009a) interpreted
this as evidence that terrestrial vertebrate dispersal between South America and Australia was
unlikely during the middle Cretaceous but more probable during the late Cretaceous, when
falling global temperatures allowed the cool-adapted high-latitude fauna and flora of
Australia-Antarctica to spread to lower latitudes, including into South America.
Remains of a fossil monotreme, Monotrematum sudamericanum, are known from the
early Palaeocene (Peligran SALMA, ~65.7-63.5 Ma; Clyde et al. 2014) “Banco Negro
Inferior” of the Salamanca Formation of Patagonia (Pascual et al. 1992; 2002; Forasiepi and
Martinelli 2003). There seem two plausible interpretations for this. Firstly, monotremes may
have been common to Australia, Antarctica and South America at least during the
Cretaceous, in which case M. sudamericanum is a South American post-KPg survivor. If so,
then the apparent absence of monotremes in the late Cretaceous Allenian faunas of South

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

America is an artefact, which is possible given that monotreme fossils are rare even in
Australia (R. Pian, pers. comm.). Alternatively, M. sudamericanum is the result of a dispersal
event from Australia to South America (presumably via Antarctica), after the Allenian but
before the Peligran, i.e. during the latest Cretaceous or earliest Palaeocene. This is congruent
with Rich et al.’s (2009a) preferred biogeographical hypothesis discussed above, and
coincides approximately with the likely timing of the dispersal of marsupials from South
America to Australia (Beck et al. 2008; Beck 2012, ; see below). In either case, monotremes
are expected to have been present in Antarctica, but have yet to be found; however, the only
known Antarctic terrestrial mammal faunas, from the early-middle Eocene La Meseta
Formation on Seymour Island off the coast of the Antarctic Peninsula, are as yet poorly
known (Gelfo et al. 2014; Reguero et al. 2013). Besides Monotrematum, definitive
monotremes are also unknown from South America.
Focusing now on the Cenozoic record of monotremes in Sahul, it is interesting that
members of this clade have to date not been found in the early Eocene (~54.6 Ma; Godthelp
et al. 1992) Tingamarra Local Fauna (Figure 4a) in northeastern Australia (pers. obv.). Again,
this may be an artefact of sampling. However, a plausible alternative explanation is that
average temperatures at Tingamarra during the early Eocene were too high for monotremes.
Today, the modern platypus, Ornithorhynchus anatinus, does not occur north of the southern
end of the Cape York Peninsula (~15º S), apparently because it is intolerant of the higher
temperatures further north (Nicol 2013; Grant 2007). If the platypus ecomorphotype is
ancestral for crown-clade monotremes (Phillips et al. 2009; 2010; Mirceta et al. 2013; but see
Camens 2010; Musser 2013; ; ; ; ; Ashwell 2013), and if echidnas (family Tachyglossidae)
did not originate until after the early Eocene (Phillips et al. 2009), then it is plausible that
early Palaeogene monotremes were semi-aquatic (Mirceta et al. 2013) platypus-like forms. If
so, they may have had similar thermal physiologies to Ornithorhynchus anatinus. Although

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Australia was ~20° further south during the early Eocene, the Tingamarra Local Fauna
(Figure 4a) only slightly postdates the Palaeocene-Eocene Thermal Maximum, when global
temperatures were as much as 12°C warmer than today (Zachos et al. 2001). Monotremes
may therefore have been restricted to more southerly latitudes at this time, in southern
Australia or perhaps in Antarctica (see also Musser 2013); however, this hypothesis remains
speculative in the absence of additional early Palaeogene mammaliaform-bearing sites and
better sampling.
The oldest Cenozoic monotremes currently known from Australia are Obdurodon
insignis and a second, currently un-named Obdurodon species from the latest Oligocene
Etadunna and Namba Formations in central Australia (Woodburne and Tedford 1975;
Woodburne et al. 1994). Discovery of a nearly complete skull of a third Obdurodon species,
the early Miocene Ob. dicksoni, confirms that Obdurodon was platypus-like in cranial
morphology (Musser and Archer 1998). The recent description of a fourth species (Pian et al.
2013), the middle Miocene or Pliocene Ob. tharalkooschild, largely fills the temporal gap
between the older Obdurodon species and the earliest remains of Ornithorhynchus, which are
reportedly Pliocene in age (Rich 1991; Musser 2013). Thus, there is evidence for the
continued presence of monotremes in mainland Australia since the late Oligocene.
There are five living species of living monotreme: the platypus Ornithorhynchus
anatinus, the short-beaked echidna Tachyglossus aculeatus, and three species of long-beaked
echidna: Zaglossus attenboroughi, Z. bartoni and Z. bruijni (Wilson and Reeder 2005; Van
Dyck and Strahan 2008; Flannery 1995b; Augee et al. 2006; Flannery and Groves 1998).
Recent point estimates for the divergence between Ornithorhynchus and the tachyglossids
Tachyglossus and Zaglossus based on molecular data are 27.7-37.8 Ma (Phillips et al. 2009;
Meredith et al. 2011), but confidence intervals (CIs) on this divergence vary markedly

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.
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depending on the analysis (composite CI = 13.3-103.1 Ma). Thus, it is unclear whether or not
the Ornithorhynchus-tachyglossid split occurred before or after the opening of the Tasman
Gateway between Australia and Antarctica; however, it probably occurred before the
emergence of the major New Guinean landmass (see “Geographical and geological context”
above). The split between Tachyglossus and Zaglossus, was estimated by Phillips et al.
(2009) to be 5.5 Ma (95% highest posterior distribution = 1.8-10.6 Ma).
Today, Ornithorhynchus anatinus occurs only along the eastern side of Australia, at
latitudes >15º S, and in Tasmania (Grant 2007; Gongora et al. 2012; Kolomyjec et al. 2013).
It has undergone range reduction since European colonisation, and may now be extinct in
South Australia (apart from an introduced population on Kangaroo Island) and throughout
much of the Murray-Darling Basin (Grant 2007). However, it appears never to have occurred
naturally in Western Australia. O. anatinus is known from the southern end of the Cape York
Peninsula, but (as discussed above) is not found further north, apparently because the ambient
temperature is too high (Nicol 2013; Grant 2007). If so, this may explain why O. anatinus
seems never to have been present in New Guinea.
Mitochondrial sequence data indicate the existence of two major clades within O.
anatinus: an Australian mainland clade, and a second clade comprising individuals from
Tasmania and King Island (Gongora et al. 2012). The divergence time between these two
clades was estimated by Gongora et al. (2012) at ~0.7-0.94 Ma, which is long before the last
dry-land connector between Tasmania and the mainland was severed (~14 ka; Lambeck and
Chappell 2001). Gongora et al. (2012) therefore suggested that these clades diverged in
mainland Australia, with subsequent extinction of the Tasmanian mitochondrial haplotype on
the mainland. Recovery of ancient DNA from subfossil O. anatinus specimens should allow
testing of this hypothesis. The north-eastern Queensland population is also genetically quite
divergent (Gongora et al. 2012), and mitochondrial sequences and microsatellites indicate

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

additional geographic structuring within populations, with different clusters coinciding with
major river drainages (Gongora et al. 2012; Kolomyjec et al. 2013). O. anatinus exhibits
Bergmann’s rule across its latitudinal range as a whole; however, at smaller spatial scales
(e.g. within a single river basin), an inverse relationship between temperature and body size
(i.e. the opposite of Bergmann’s rule) is observed (Furlan et al. 2011). In southeastern
Australia, both lower rainfall and higher temperatures were found to be associated with larger
sized individuals by Furlan et al. (2011).
Tachyglossus aculeatus occurs in both Australia and New Guinea, with five
subspecies currently recognised (Augee et al. 2006; Griffiths 1978). These subspecies appear
to show clear biogeographic structuring (Augee et al. 2006; Griffiths 1978): T. a. acanthion
in the Northern Territory, northern Queensland, inland Australia and Western Australia; T. a.
aculeatus in eastern New South Wales, Victoria and southern Queensland; T. a. lawesii in the
lowlands of New Guinea and possibly also the rainforests of northeastern Queensland; T. a.
multiaculeatus in mainland South Australia and Kangaroo Island; and T. a. setosus in
Tasmania. Intriguingly, body size in T. aculeatus corresponds to the reverse of Bergmann’s
rule, with the subspecies occurring at highest latitudes (the Tasmanian T. a. setosus) being
smallest (Augee et al. 2006). The presence of T. aculeatus lawesii in New Guinea is probably
best explained as a relatively recent dispersal from Australia, given that the same subspecies
is reportedly present in northeastern Queensland (Griffiths 1978). Dry-land connectors have
formed repeatedly between Australia and New Guinea since the major emergence of New
Guinea ~5 Ma, and have been the rule rather than the exception over the last ~1 Myr (see
“Geographical and geological context” above), which presumably facilitated dispersal of T.
aculeatus, perhaps as part of a shared Austral-East Torresian mammal fauna (Lavery et al.
2013; see below).

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.
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The three living species of long-beaked echidna occur only in New Guinea, with Z.
bruijnii occuring in the west, Z. bartoni in central and eastern regions, typically at high
elevations, and Z. attenboroughi in the Cyclops Mountains in the island’s north (Flannery
1995b; Flannery and Groves 1998; Helgen et al. 2012; Baillie et al. 2009). Flannery and
Groves (1998) identified four subspecies within Z. bartoni, with three of them forming a
longitudinal cline in body size along the New Guinean central cordillera: the smallest, Z. b.
smeenki, in the east, the largest, Z. b. diamondi, in the west, and the intermediate-sized Z. b.
bartoni between them. A fourth subspecies, Z. b. clunius, is known from the Huon Peninsula
in the east of the island. Recognisable long-beaked echidnas (most likely Zaglossus sp.) are
depicted in Aborginal rock art, possibly late Pleistocene in age, from Arnhem Land in the
Northern Territory of Australia (Helgen et al. 2012). In addition, Helgen et al. (2012)
discussed a long-beaked echidna specimen in the Natural History Museum, London that was
apparently collected in 1901 in the West Kimberley region of northern Western Australia,
and which they identified as Z. bruijnii (currently endemic to western New Guinea). Helgen
et al. (2012) also discussed accounts by Aboriginal people living in the East Kimberley that
may refer to Zaglossus, and suggest that it may still occur in the Kimberley region.
Depending on the exact relationship between the putative Kimberley Zaglossus specimen to
New Guinean populations, this raises the possibility of a complex biogeographical history for
the genus, particularly given the existence of putative Zaglossus specimens from multiple
Plio-Pleistocene sites in Australia (discussed below).
Turning now to the fossil record of tachyglossids, possibly the oldest known remains
are from a gold mine in New South Wales (Dun 1895), which has been suggested to date to
the middle Miocene, but which may in fact be Pleistocene in age (Augee et al. 2006; 2013;
Musser 2006). This material has been referred to the modern genus Zaglossus or to the
extinct genus Megalibgwilia¸ as Z. or M. robusta, by different authors (Griffiths et al. 1991;

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Augee et al. 2006; Musser 2003; 2013; Musser 2006; Long et al. 2002). The next oldest
tachyglossids are from Plio-Pleistocene deposits in mainland Australia. Of these, the short-
beaked forms have been referred to the extant species Tachyglossus aculeatus (although these
putative T. aculeatus specimens are markedly larger than modern individuals; Augee et al.
2006; Pledge 1980). The long-beaked forms have been referred to the extant genus Zaglossus
and/or the fossil genus Megalibgwilia (Augee et al. 2006; Musser 2006; Murray 1978a,
1978b), None of the long-beaked material from Australia has been referred to a living
Zaglossus species, apart from postcranial remains from the Pleistocene Henschke’s Quarry
Cave at Naracoorte in South Australia, which Murray (1978b) provisionally referred to the
living Z. bruijnii; however, Murray (1978a) subsequently referred this material to Zaglossus
sp., and Pledge (1980) suggested it probably represents Tachyglossus. Hocknull (2009) also
reported a manual ungual of an indeterminate tachyglossid from the middle Pleistocene (at
least 330 ka; Hocknull et al. 2007) of Mt Etna in northern Queensland. Tachyglossids (and
other monotremes) are not known from the pre-Pleistocene of New Guinea, although the
fossil record is poor, with only a single mammal-bearing deposit known, namely the middle
Pliocene Otibanda Formation (Plane 1967; Flannery et al. 1993; Long et al. 2002).
A better understanding of the biogeographical history of tachyglossids is prevented by
current uncertainties regarding their taxonomy and phylogeny. The alpha-taxonomy of the
living genera Tachyglossus and Zaglossus is in need of revision and testing with molecular
data. Broadscale phylogeographic analyses of molecular sequence data and associated
estimates of divergence times are also required to clarify when T. aculeatus dispersed to New
Guinea, and whether there was a single or multiple dispersals. Likewise, the biogeographical
history of Zaglossus will remain unclear until the relationships and divergence times between
the New Guinean Zaglossus species, the Plio-Pleistocene long-beaked echidnas of mainland
Australia (Zaglossus and Megalibgwilia species) and the Kimberley Zaglossus specimen

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

(Helgen et al. 2012) are clarified. To this end, a thorough revision and formal phylogenetic
analysis of living and fossil long-beaked echidnas is sorely needed; in particular, the validity
of Megalibgwilia, distinct from Zaglossus, needs to be tested. Ideally, such an analysis should
incorporate DNA sequence data from multiple representatives of extant New Guinean
Zaglossus species, and (if DNA can be obtained) from the Kimberley specimen and subfossil
material of long-beaked echidnas from Australia.
4.3 Ausktribosphenidae
Ausktribosphenos nyktos and Bishops whitmorei are tribosphenic mammals known from
multiple partial lower jaws from the early-middle Aptian Flat Rocks site (Rich et al. 2001a;
Rich et al. 1997), with a species of Bishops (possibly B. whitmorei) also known from the
similarly-aged Eric the Red West site (Rich et al. 2009b). Ausktribosphenos and Bishops are
currently classified as the only named representatives of the family Ausktribosphenidae (Rich
et al. 1997; 2001a; Kielan-Jaworowska et al. 2004). It has been proposed that
aukstribosphenids are eutherians (Rich et al. 1997; 2001a; Woodburne et al. 2003). If so, this
would pose something of a biogeographic conundrum: the majority of fossil evidence
supports a Laurasian origin for Eutheria (Kielan-Jaworowska et al. 2004; Ji et al. 2002; Luo
et al. 2011), and there is no evidence of either eutherians or metatherians in the late
Cretaceous Allenian faunas of southern South America (see above). Both eutherians and
metatherians were widespread and diverse throughout Laurasia during the Cretaceous
(Kielan-Jaworowska et al. 2004), and were also highly diverse in early-middle Palaeocene
faunas known from South America (Marshall and Muizon 1988; Muizon 1991; Gelfo et al.
2007; Bonaparte et al. 1993; Muizon and Cifelli 2001; Goin et al. 1992), suggesting that

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

therian mammals in fact first entered Gondwana during the latest Cretaceous or early
Palaeocene (see below).
However, most studies have concluded that Ausktribosphenos and Bishops fall outside
Theria, and are members of a group that independently evolved a tribosphenic dentition (Luo
et al. 2001; Luo et al. 2002; Kielan-Jaworowska et al. 2004; Davis 2011). In support of this
conclusion, most published phylogenetic analyses (e.g. Luo et al. 2001; Luo et al. 2002;
Kielan-Jaworowska et al. 2004; Rougier et al. 2007; Bi et al. 2014) place Ausktribosphenos
and Bishops in a clade with monotremes and three other tribosphenic mammals from the
Jurassic of Gondwana: Henosferus molus and Asfaltomylos patagonicus from the Toarcian of
South America (Rougier et al. 2007; Rauhut et al. 2002; Martin and Rauhut 2005), and
Ambondro mahabo from the Bathonian of Madagascar (Flynn et al. 1999). This clade was
named Australosphenida by Luo et al. (2001).
Relationships within Australosphenida have yet to be fully resolved and are likely to
remain so pending the discovery of more complete material of this enigmatic group; as such,
the biogeography of the group is also uncertain. However, based on the Early Jurassic age of
Henosferus and Asfaltomylos (see Cúneo et al. 2013), Australosphenida originated prior to
the break-up of Gondwana. Recent phylogenetic analyses suggest that ausktribosphenids and
monotremes form a clade to the exclusion of Asfaltomylos and Ambondro (Rougier et al.
2007; Bi et al. 2014), raising the possibility that Ausktribosphenidae+Monotremata
represents a localised eastern Gondwanan or Australian radiation of australosphenidans.
Certainly, it is interesting that australosphenidans are unknown from Cretaceous deposits in
South America (notably the diverse Allenian faunas), and so it is possible that the group had
become extinct in western Gondwana by the late Cretaceous but survived in the east, along
with other possibly relictual taxa such as the temnospondyl Koolasuchus (see the discussion
in “Monotremata” above).

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.
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4.4 Marsupialiformes
The oldest generally accepted metatherian is Sinodelphys szalayi from the 125 Ma Yixian
Formation in northeastern China (Luo et al. 2003). The subsequent description of the
apparent eutherian Juramaia sinensis from the Upper Jurassic Daohugou Beds in the same
region by Luo et al. (2011) appears to push the age of the Metatheria-Eutheria split back to a
minimum of ~160 Ma. However, doubts as to both the affinities and age of Juramaia have
been raised (Jansa et al. 2014, : supporting information; Sullivan et al. 2014). Recent
molecular estimates for this divergence are broad, spanning 140.5-215.3 Ma when confidence
intervals are taken into account (dos Reis et al. 2012; 2014; Meredith et al. 2011).
Critically, whereas metatherians and eutherians are common elements of middle-to-
late Cretaceous faunas in Laurasia, they have not been found in similarly-aged faunas in the
southern hemisphere (Kielan-Jaworowska et al. 2004), with the exception of a questionable
marsupialiform known from a single partial molar from the Maastrichtian of Madagascar
(Krause 2001; but see Averianov et al. 2003), and a few eutherians from the Maastrichtian of
India (Prasad et al. 1994; 2007; Rana and Wilson 2003; Khosla and Verma 2014). The entire
published Mesozoic mammaliaform record in Australia is restricted to four Aptian-Albian
localities (Figure 3a) – Lightning Ridge in New South Wales, and the Dinosaur Cove, Flat
Rocks and Eric the Red West sites in southern Victoria - and only seven taxa have been
named from these. Thus, the apparent absence of therian mammals from the Mesozoic of
Australia could plausibly be an artefact of incomplete sampling. Of greater significance is the
lack of therians from the much richer Mesozoic mammal record of South America. Prior to
the late Cretaceous, the South American record is relatively sparse (Rougier et al. 2011b).
However, the comparatively rich late Cretaceous Allenian faunas of southern South America

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

lack any trace of therians, preserving instead a diverse, apparently highly endemic
mammalian fauna dominated by non-therian “dryolestoids” and gondwanatherians (Rougier
et al. 2009a; Bonaparte 1990; Rougier et al. 2009b; Rougier et al. 2011b). The simplest
explanation for the lack of therians in these diverse Allenian faunas, and one assumed by
numerous authors (e.g. Szalay 1994; Pascual and Ortiz-Jaureguizar 2007; Pascual 2006; Goin
et al. 2012b; Beck 2008b), is that they were genuinely absent from South America at this
time. Given that the most likely point of entry of therians into Gondwana was from North
America to South America (Case et al. 2005), their apparent absence from South America
during the Allenian is likely an indication that they were also absent from at least those parts
of Gondwana in direct contact with South America at this time, namely Antarctica and
Australia.
The precise age of the Allenian faunas is unknown, but they are most likely late
Campanian-early Maastrichtian in age (Pascual and Ortiz-Jaureguizar 2007; Rougier et al.
2009a; Rougier et al. 2009b). A few mammaliaform fossils are known from a younger South
American Cretaceous site, the middle Maastrichtian Pajcha Pata locality in Bolivia (Gayet et
al. 2001); they include at least one “dryolestoid”, but definitive therians have not been found
there (Rougier et al. 2011b, ; pers. obv.). However, the Pajcha Pata fauna is still too poorly
known to confidently rule out the presence of therians. A conservative maximum bound on
the age of entry of therians (i.e. marsupialiforms and eutherians) into South America is
therefore the maximum age of the Campanian, 83.6 Ma, based on a conservative maximum
age of the Allenian. The diverse mammal fauna from the “Banco Negro Inferior” of the
Salamanca Formation in southern Argentina includes marsupialiforms and eutherians, as well
as non-therian taxa (Gelfo et al. 2007; Bonaparte et al. 1993; Goin et al. 1992). This fauna
forms the basis of the Peligran SALMA, and has recently been dated as early-middle Danian
(~65.7–63.5 Ma; Clyde et al. 2014). This date provides a minimum age for the dispersal of

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

therians from North America to South America. Thus, the probable time of dispersal of both
marsupialiforms and eutherians from North to South America can be constrained to between
83.6 and 63.5 Ma.
A combination of geological, fossil and phylogenetic evidence suggests that the
presence of marsupialiforms in Australia is the result of dispersal from South America, across
Antarctica, prior to the deepwater opening of the Drake Passage and Tasman Gateway
(Woodburne and Case 1996; Kemp 2005; Beck 2008b, 2012; Beck et al. 2008; Lawver et al.
2011). The oldest marsupialiforms known from Australia are from the Tingamarra Local
Fauna in southeastern Queensland (Figure 4a; Beck 2012; Beck et al. 2008; Beck 2014;
Godthelp 1999; Sigé et al. 2009), which has been radiometrically dated as 54.6 Ma (Godthelp
et al. 1992). This provides a minimum date for the dispersal of marsupialiforms to Australia.
The maximum likely date for dispersal can be set at 83.6 Ma, based on the maximum likely
date for the entry of marsupialiforms into South America (see above).
As discussed by Kemp (2005: 218-221) and Beck (2008b, 2012), a key issue is
whether there was: 1) a single dispersal by marsupialiforms (restricted to crown-group
marsupials) from South America to Australia only, which would imply that major dispersal
barriers were already in place by the late Cretaceous-early Palaeogene (the “single dispersal”
model); 2) multiple independent dispersals between South America and Australia, implying
less severe dispersal barriers (the “multiple dispersals” model) ; or 3) a single, relatively
continuous marsupialiform fauna stretching from at least the southern part of South America,
across Antarctica to Australia (collectively, the Austral Kingdom; Morrone 2002; Goin et al.
2007; Goin et al. in press; Aragón et al. 2011) during at least the early part of the Palaeogene,
implying weak or absent dispersal barriers (the “continuous fauna” model). Under the
“continuous fauna” model, faunal differences between South America and Australia would

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

have developed subsequently as a result of vicariance - most plausibly the deepwater opening
of the Drake Passage and Tasman Gateway - and differential extinction (Beck 2012).
Recent molecular analyses of modern marsupials suggest that the four modern
Sahulian marsupial orders form a clade, Eomarsupialia (see Table 1), with respect to the three
extant South American marsupial orders (Didelphimorphia, Paucituberculata and
Microbiotheria), which form a paraphyletic assemblage outside Eomarsupialia (Beck 2008a;
Meredith et al. 2009b; Meredith et al. 2011; Nilsson et al. 2010; Mitchell et al. 2014;
Meredith et al. 2009a). An obvious interpretation of this pattern is that the presence of
marsupials in Sahul is the result of a single dispersal event, from South America, with an
ancestor giving rise to the entire Sahulian marsupial radiation, i.e. the “single dispersal”
model discussed above (Meredith et al. 2009a; Meredith et al. 2009b; Nilsson et al. 2010).
However, for this to be true, all fossil marsupialiforms from Sahul must also be part of the
same radiation, and recent studies indicate that this is not the case. Specifically, material from
the early Eocene Tingamarra Local Fauna (Figure 4a) – namely specimens referable to
Djarthia murgonensis (Figure 4i) and also an isolated calcaneus (QM F30060; Figure 4f) that
represents a second taxon - falls within Marsupialia but outside crown-clade Australidelphia
(which includes the South American Dromiciops) in published phylogenetic analyses (Figure
4b), suggesting that the “single dispersal” model can be rejected (Beck 2012; Beck et al.
2008). Although yet to be included in formal phylogenetic analyses, the Tingamarran species
Thylacotinga bartholomaii, Chulpasia jimthorselli (Figure 4c) and Archaeonothos
henkgodthelpi (Figure 4e) also show no clear evidence of belonging to Eomarsupialia: T.
bartholomaii and C. jimthorselli show closest similarities to C. mattaueri (Figure 4d) from
the late Palaeocene or early Eocene of Peru (Sigé et al. 2009), while A. henkgodthelpi
resembles Kasserinotherium tunisiense from the early Eocene of Tunisia and Wirunodon
chanku from the ?middle-late Eocene of Peru (Beck 2014). Thus, monophyly of the modern

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Sahulian marsupial radiation relative to modern South American marsupials appears to be the
result of extinction of non-eomarsupialian lineages in Australia, some time between the early
Eocene and late Oligocene.
As I have noted previously (Beck 2012), distinguishing between the other two
possibilities, namely the “multiple dispersals” and “continuous fauna” models, is difficult
given deficiencies in the available fossil record, particularly from the early Palaeogene of
Australia and Antarctica. However, the apparent absence of typically “South American”
marsupialiforms, such as the dentally distinctive polydolopids (which are also common in the
middle Eocene La Meseta Fauna of Antarctica; Chornogubsky et al. 2009), and also eutherian
groups such as “meridiungulates” and xenarthrans, from Tingamarra and younger Australian
sites argues against the “continuous fauna” model (Beck 2012). Similarly, recent molecular
estimates of divergence times indicate that the modern Sahulian marsupial orders had
diverged from each other by the middle Eocene at the latest (Beck 2008a; Mitchell et al.
2014; Meredith et al. 2011; Meredith et al. 2009a; Meredith et al. 2009b), but unequivocal
members of these orders have not been found in any South American site or the La Meseta
Fauna. Thus, on present evidence, the “multiple dispersals” model appears most likely.
However, the number of dispersals is unclear, and it is uncertain whether they were all from
South America to Australia, or whether dispersal(s) in the reverse direction also occurred. For
example, it is possible that the presence of microbiotherians in western Antarctica and South
America is the result of a back-dispersal from Australia (Beck et al. 2008; Beck 2012).
The “multiple dispersals” model implies the presence of dispersal barriers between
South America and Australia during the late Cretaceous-early Palaeogene, preventing the
formation of a single continuous mammalian fauna (Beck 2012, 2008b). I have previously
speculated (Beck 2008b) that Antarctica may have posed such a barrier due to: 1) the narrow,
mountainous, high-latitude connection between western and eastern Antarctica; 2) low

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

temperatures (particularly at the highest latitudes), even during the “greenhouse” conditions
of the Late Cretaceous and early Palaeogene (Poole et al. 2005; Francis and Poole 2002;
Kemp et al. 2014), and possibly (3) the Valdivian-type flora that extended from southern
South America across Antarctica and into Australia (see also Case et al. 1988). Based on this,
I suggested that trans-Antarctic dispersal by terrestrial mammals during the latest Cretaceous
and early Palaeogene would have been more likely for those taxa characterised by a high
basal metabolic rate, cold tolerance, the ability to hibernate, small body size and arboreal
adaptations (Beck 2008b).
Turning now to the modern Sahulian radiation, a key issue has been to clarify the
precise biogeographical relationships between the Australian and New Guinean marsupial
faunas. Flannery (1988) proposed that the development of a seaway between the two
landmasses led to the marsupial faunas of the two landmasses becoming distinct by the early
Miocene at the latest, and that a dry-land connection was not re-established until the
Pleistocene. However, this scenario conflicts with current evidence (summarised in
“Geographical and geological context” above) that: 1) the majority of the New Guinean
landmass has become emergent only in the last 5 Myr (Figure 1a-c); and 2) that dry-land
connectors have formed repeatedly between Australia and New Guinea since at least 3.5 Ma
(Figure 1d). Several studies have used molecular estimates of divergence dates between
modern Australian and New Guinean taxa to infer the timing of putative dispersal events
between the two landmasses, and determine whether they conform to Flannery’s (1988)
model (e.g. Aplin et al. 1993; Kirsch and Springer 1993; Westerman et al. 2012; Mitchell et
al. 2014; Meredith et al. 2010; Raterman et al. 2006). The most recent and comprehensive of
these is that of Mitchell et al. (2014), who observed that most of their inferred dispersals
between Australia and New Guinea can be dated as having occurred within the last 5 Myr,
after the major emergence of the New Guinean landmass. This is congruent with Murray’s

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

(1992) conclusion that Plio-Pleistocene zygomaturine diprotodontids known from New
Guinea are the result of dispersal from Australia during the middle-to-late Pliocene.
However, Mitchell et al. (2014) argued that the divergence times of three
predominantly New Guinean marsupial clades – namely 1) a clade within Phalangeridae
comprising the genera Ailurops, Phalanger, Spilocuscus and Strigocuscus; 2) the Murexia
sensu lato dasyurid clade (which encompasses the genera Murexia, Micromurexia,
Murexechinus, Paramurexia and Phascomurexia; Van Dyck 2002; Krajewski et al. 2007;
Groves 2005a); and 3) peroryctid bandicoots (Echymipera, Microperoryctes, Peroryctes and
Rhynchomeles) – suggest early dispersals from Australia to New Guinea, probably some time
between 9 and 11 Ma. Given the apparent synchronicity of these dispersals, Mitchell et al.
(2014) considered that they are unlikely to have been over marine barriers, contra current
geological evidence that there was probably no land connection between New Guinea and
Australia until after the major period of uplift of New Guinea ~5 Ma (see “Geographical and
geological context” above).
There are, however, at least two alternative explanations for the presence of these
three “old” clades in New Guinea. Firstly, and perhaps more prosaically, it may reflect
problems with the molecular clock analyses implemented by Mitchell et al. (2014), resulting
in overestimated divergence dates; many current molecular clock models appear unable to
fully account for rapid changes in the rate of molecular evolution, and have likely
overestimated the ages of certain divergences in published studies (Kitazoe et al. 2007;
Dornburg et al. 2012; Waddell 2008; Steiper and Seiffert 2012; Dornburg et al. 2014).
Indeed, such issues should be borne in mind when considering all of the molecular
divergence dates presented in this chapter.

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Perhaps a more interesting alternative explanation is that the apparent antiquity of
these three clades is due to the extinction of Australian relatives. Support for this
interpretation comes from the rich middle Pleistocene vertebrate fauna from Mount Etna in
northern Queensland, which has revealed the presence of multiple typically “New Guinean”
taxa that have gone extinct in Australia within the last 280 ka (Hocknull et al. 2007; Hocknull
2005; Cramb et al. 2009; Hocknull 2009). These include probable representatives of the three
“old” New Guinean clades identified by Mitchell et al. (2014), most notably: 1) Phalanger
gymnotis (see Hocknull 2009); 2) a dasyurid that Cramb et al. (2009) identified as cf.
Micromurexia habbema; 3) three peramelemorphians that appear most similar among living
bandicoots to Peroryctes and Microperoryctes (see Hocknull 2005; 2009). Thus, it seems that
the current lack of representatives of these three clades in Australia – with the exception of
the peroryctid Echymipera rufescens and the phalangerids Phalanger mimicus and
Spilocuscus maculatus, all of which appear to represent recent divergences from New
Guinean conspecifics – is the result of extinction, rather than prolonged biogeographical
isolation from New Guinea. Congruent with this interpretation, several other marsupials from
middle Pleistocene deposits at Mt Etna have also been argued to be more closely related to
New Guinean than to Australian species among living taxa, including species of
Dendrolagus, Dactylopsila and Pseudochirops (Hocknull 2005; Hocknull 2009). Also of
relevance is the presence at Mt Etna of several highly distinctive taxa that appear to represent
major lineages that are now entirely extinct. These include the phascolarctid Invictokoala
monticola (see Price and Hocknull 2011) and representatives of three as-yet-undescribed
dasyurid genera (Cramb et al. 2009). This further demonstrates that the modern fauna of
Australia has been shaped by the complete loss of several major marsupial lineages within the
last 280 ka.

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

In fact, it is perhaps more appropriate to consider northeastern Australia and New
Guinea, as well as islands such as the Moluccas, as essentially having a shared fauna
throughout at least the Pliocene and early Pleistocene, intermittently divided during
interglacial periods; this corresponds to Lavery et al.’s (2013) concept of the Austral-East
Torresian zoogeographic province, spanning the northeast of Australia and southwest of New
Guinea (see also Helgen 2007; Flannery 1995b). If members of this shared fauna then
became extinct in Australia, perhaps with the onset of more xeric conditions 280-205 ka
(Hocknull et al. 2007), this might explain the relatively ancient divergences between
predominantly New Guinean and predominantly Australian sister-taxa found in molecular
studies of modern species, without recourse to early (and geologically implausible) dispersals
to New Guinea of the kind proposed by Mitchell et al. (2014).
Focusing specifically on the modern New Guinean marsupial fauna, Helgen (2007)
represents a recent, comprehensive overview of the taxonomy and distribution of marsupials
and other mammals on the island. However, there is a need for detailed phylogeographic
studies, both to clarify species boundaries and to determine the biogeographical factors that
have shaped their distributions. For example, they would allow further testing and refinement
of the Oceanic, Tumbanan and Austral zoogeographic provinces recognised by Lavery et al.
(2013; see also Flannery 1995b; Helgen 2007), and help understand the role that uplift and
altitudinal gradients have played shaping marsupial diversity on the island (Macqueen et al.
2011; Meredith et al. 2010; Helgen 2007). One of the few such studies published to date is
Macqueen et al.’s (2011) phylogeographic analysis of New Guinean pademelons (Thylogale
spp.). Macqueen et al. (2011) found evidence for the existence of “western” and “eastern”
Thylogale clades that do not correspond to the current species taxonomy (Flannery 1995b;
Groves 2005b) or to Flannery’s (1992) recognition of northern, central and southern groups;
instead, the major split coincides with the Ramu-Markham and Watut-Tauri valleys, in the

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

east of New Guinea. Additional genetic structuring within the western and eastern Thylogale
clades is complex, and likely reflects both geological (e.g. recent uplift) and climatic (e.g.
vegetational changes between glacial and interglacial periods) factors (Macqueen et al. 2011).
It remains to be seen whether other New Guinean marsupial species show congruent or
contrasting biogeographical patterns.
Published phylogeographic studies of Australian marsupials have been far more
numerous (e.g. Pope et al. 2000; Brown et al. 2006; Cooper et al. 2000; Spencer et al. 2001;
Potter et al. 2012; Neaves et al. 2009; Malekian et al. 2010; Macqueen et al. 2012; Pavlova et
al. 2010; Pope et al. 2001; Eldridge et al. 2011; Hazlitt et al. 2014; Firestone et al. 1999), and
cannot be fully summarised here. These studies have uncovered complex biogeographical
patterns, which have been interpreted to be the result of numerous different factors, including
the existence of past and present biogeographical barriers to gene flow, climate change, and
habitat contraction and expansion. Perhaps most striking is that biogeographic patterns found
in these studies appear to be highly individualistic, with little evidence of commonalities
between different marsupial species. For example, the Burdekin Gap (an area of dry
woodland separating the Wet Tropics of Queensland from mesic forest habitats further south)
appears to have formed a major barrier to gene flow in the yellow-bellied glider (Petaurus
australis; Brown et al. 2006) but not in the red-legged pademelon (Thylogale stigmatica;
Eldridge et al. 2011), even though both are forest-adapted species. Similarly, Potter et al.
(2012) found evidence of a deep divergence between populations of brachyotis-group rock
wallabies either side of the East-West Kimberley Divide, whereas similar phylogeographic
structuring was not observed in the scaly-tailed possum (Wyulda squamicaudata; Potter et al.
2014), despite the fact that both are rock-dwelling species.
These individualistic biogeographic patterns are perhaps unsurprising, given species’
differing dispersal abilities, niche requirements and other aspects of basic biology.

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Nevertheless, they may not be easily predictable: for example, Eldridge et al. (2014) found
evidence of strong genetic structuring across the Carpentarian Barrier (an area of semi-arid
grassland in northern Australia separating more mesic savannah woodland habitats further
west and east) in the common wallaroo (Macropus robustus), even though the species is a
habitat generalist that is continuously distributed throughout the region. Conversely, the
antilopine wallaroo (M. antilopinus) showed little genetic differentiation across the
Carpentarian Barrier, despite the fact that it is a tropical woodland specialist that does not
occur in the region of the barrier (Eldridge et al. 2014). In fact, the phylogeographic
structuring observed in M. robustus and M. antilopinus in northern Australia is the reverse of
a priori predictions based on the known biology and distributions of the two species
(Eldridge et al. 2014). Similar studies of other marsupial species may reveal further surprises.
Two factors are likely to complicate future phylogeographic studies of modern
Sahulian marsupials. Firstly, it is clear that during the late Pleistocene-Holocene, numerous
New Guinean (Helgen 2007; Aplin et al. 1999b; Aplin and Pasveer 2006) and Australian
(Johnson 2006; Burbidge et al. 2008; McKenzie et al. 2007) species have gone entirely
extinct. Many others have radically reduced distributions, with the loss of entire populations,
at least some of which likely represent major, evolutionarily distinct lineages. Conversely,
there is also evidence that current distributions of some New Guinean marsupials have been
influenced by human-mediated introductions (Macqueen et al. 2011; Heinsohn 2010). As a
result, studies that use molecular and/or distributional data from current populations only may
be misled as to the likely biogeographical history of the species in question; in fact, this
might explain the lack of evidence for common biogeographical patterns in phylogeographic
studies published to date. Once again, evidence from the fossil and subfossil record, and also
from museum specimens collected during the early years of European colonisation (before
the extinction or extreme range reduction of many small- and medium-sized marsupial

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

species in Australia) will be critical; this might be simply as distributional data (as used, for
example, in the biogeographic analyses of Burbidge et al. 2008 and Lavery et al. 2013) or, if
obtainable, as ancient DNA.
4.5 ?Condylartha
Godthelp et al. (1992) described an isolated tribosphenic lower molar (Figure 4h) from the
early Eocene Tingamarra Local Fauna (Figure 4a) that they named Tingamarra porterorum.
They tentatively referred Tingamarra to “Condylarthra”, a non-monophyletic assemblage of
eutherians, some of which likely gave rise to the various “ungulate” groups (Rose 2006;
Archibald 1998). “Condylarths” are first known in South America from the early Palaeocene
(Peligran SALMA; Gelfo et al. 2007; Gelfo 2007; Clyde et al. 2014), probably the result of
dispersal from North America, at around the same time that marsupialiforms seem to have
taken the same route (Muizon and Cifelli 2001, ; see "Marsupialiformes" above). Thus,
dispersal of this group to Australia, again presumably along the same route taken by
marsupialiforms (namely from South America, via Antarctica), seems plausible.
However, the single known tooth of T. porterorum does not show close similarities to,
and is also markedly smaller than, any known South American “condylarth” (Muizon and
Cifelli 2000; Gelfo 2007; Gelfo et al. 2007; Gelfo and Sigé 2011). Furthermore, eutherians
besides bats are otherwise entirely absent from Australian fossil deposits until the first
appearance of murine rodents ~4 Ma (Long et al. 2002; Archer et al. 1999b); if T. porterorum
is indeed a eutherian, this lineage appears to have gone extinct in Australia before the late
Oligocene. Another lower molar that shows a very similar overall morphology to T.
porterorum but is distinctly larger has been found at Tingamarra but remains undescribed
(Godthelp et al. 2001). Study of this material, together with an isolated petrosal from

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Tingamarra that apparently represents a small non-volant eutherian, should shed further light
on this biogeographically puzzling taxon (Beck et al., in prep.).
4.6 Chiroptera
Besides the putative “condylarth” Tingamarra (see above) and murine rodents (see below),
bats (order Chiroptera) are the only eutherians known to have reached Australia without
human assistance. This is perhaps unsurprising: as the only truly volant mammals, bats are
excellent dispersers, and they are often the only native mammals on otherwise isolated
landmasses such as oceanic islands (Flannery 1995a; Simmons 2005; van der Geer et al.
2010). Hand (2006) provided a recent and comprehensive overview of Sahulian bat diversity
and biogeography, and the following is largely a summary of this work.
Recent phylogenetic analyses of placental mammals place bats within the
superordinal clade Laurasiatheria, together with artiodactyls, perissodactyls, carnivorans,
pangolins and eulipotyphlan ‘insectivores’ (Meredith et al. 2011; O'Leary et al. 2013). As the
name suggests, current evidence suggests that this superorder probably originated in Laurasia
(Springer et al. 2011). However, the oldest record of bats in Australia is also one of the
earliest globally, namely Australonycteris clarkae from the early Eocene Tingamarra Local
Fauna (Figure 4g; Hand et al. 1994); only a few European bats from the earliest Eocene may
be slightly older (Tabuce et al. 2009). Indeed, bats appear to have achieved an essentially
global distribution early in the Eocene (Smith et al. 2012).
Based on unpublished postcranial material, A. clarkae appears to have been fully
volant (S. J. Hand, pers. comm.). Its phylogenetic relationships have yet to be fully assessed,
but preliminary analyses indicate that it lies outside crown-clade Chiroptera (Hand and Beck,

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

in prep.). If so, it represents an early dispersal to Australia unrelated to later dispersals by
crown-clade forms (Hand 2006). It is possible that Australonycteris followed the same
dispersal route into Australia as did marsupialiforms and the ‘condylarth’ Tingamarra (and
possibly, in the opposite direction, the monotreme Monotrematum), namely from South
America, via Antarctica. However, the oldest known bats from South America are younger
(~48-52 Ma) than A. clarkae, and do not appear to be closely related to it (Tejedor et al.
2005). Furthermore, given that bats are clearly capable of dispersing over large distances
overwater, there seems no reason why Australonycteris need necessarily have taken a
terrestrial route to Australia. Further study of early bats from other landmasses, such as those
from the early Eocene Vastan fauna of India (Smith et al. 2007), and their incorporation into
broadscale phylogenetic analyses, should help clarify their biogeographical relationships.
Until then, the closest relative(s) and likely biogeographical origin of Australonycteris will
remain obscure.
After Australonycteris, the fossil record of bats (and, indeed, other mammals) in
Sahul is blank until the late Oligocene. However, representatives of at least eight crown-clade
families are known from Sahul: Mystacinidae, Hipposideridae, Megadermatidae, Molossidae,
Emballonuridae, Vespertilionidae, Rhinolophidae and Pteropodidae (Hand 2006). Of these,
most are known from Oligo-Miocene sites in Australia (Hand 2006). However,
emballonurids are first known from the Rackham’s Roost site at Riversleigh World Heritage
Area in northwestern Queensland (Hand 2006), which was originally suggested to be
Pliocene in age based on biocorrelation, but on the basis of radiometric dates now appears to
be early Pleistocene (Woodhead et al. 2014). Pteropodids also have no pre-Pleistocene
Australian record (Hand 2006).
Based on their known fossil and modern distributions, it seems plausible that the
presence of these families in Sahul is the result of dispersal from Sundaland, via Wallacea

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

(Hand 2006), with the notable exception of Mystacinidae (discussed below). However,
uncertainty remains regarding the phylogeny and even the alpha-taxonomy of many of these
families (e.g. Lavery et al. 2014; Reardon et al. 2014; Almeida et al. 2014), limiting the
biogeographical inferences that can be drawn at present. In addition, it is clear that the fossil
record of bats in Sahul exhibits extreme sampling biases, with the vast majority of specimens
known only from the Riversleigh World Heritage Area (Hand 2006). Resolving the
biogeographical relationships of these families – for example, determining the extent to
which Sahul itself has acted as a centre of diversification (e.g. for Hipposideridae; Hand and
Kirsch 1998) - will require more comprehensive phylogenetic analyses that integrate
molecular and fossil evidence.
Unlike the other bat families discussed above, the presence of mystacinids in
Australia is unlikely to be the result of dispersal from Sundaland (Hand 2006). Mystacinidae
is a member of the superfamily Noctilionoidea (Teeling et al. 2005; Meredith et al. 2011), the
other members of which are largely restricted to the southern hemisphere (Simmons 2005;
Gunnell et al. 2014; Czaplewski and Morgan 2012). Noctilionoids are entirely unknown from
Sundaland or Wallacea (Simmons 2005; Czaplewski and Morgan 2012; Gunnell et al. 2014).
Today, Mystacinidae is represented by a single extant species, M. tuberculata, which is
endemic to New Zealand; a second New Zealand species, M. robusta appears to have gone
extinct within the last 50 years (Simmons 2005). At least two mystacinid species were
present in the early Miocene St Bathans fauna, Central Otago, on New Zealand’s south
island, demonstrating that the group has been present in New Zealand for at least the last 16
million years (Hand et al. 2013). However, at least four mystacinid species are known from
late Oligocene to middle Miocene deposits in Australia (Hand et al. 2005). Recent molecular
estimates suggest that Mystacinidae diverged from other noctilionoids 41.0-52.9 Ma (Miller-
Butterworth et al. 2007; Meredith et al. 2011); this largely postdates recent estimates for the

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

separation of New Zealand from Antarctica (Ericson et al. 2014, ; see "Geographical and
geological context" above), suggesting that, regardless of the precise origin of mystacinids,
their dispersal either into or out of New Zealand was probably overwater. However, the fossil
record and phylogeny of Mystacinidae are still too poorly known to determine whether the
family originated in Australia and dispersed to New Zealand, vice versa, or whether it
originated on another landmass, such as Antarctica (Hand et al. 2013; Kirsch et al. 1998;
Teeling et al. 2003; Hand et al. 1998).
4.7 Murinae
Today, murine rodents comprise >160 species in Sahul, ~25% of the total native mammal
fauna (Musser and Carleton 2005; Van Dyck and Strahan 2008; Flannery 1995b; Aplin and
Ford 2014; Aplin 2006; Breed and Ford 2007). This number is likely to rise considerably as a
result of ongoing fieldwork, taxonomic revisions, and DNA sequencing of collected
specimens uncovering cryptic diversity. At least five species (Mus musculus/domesticus and
at least four species of Rattus) present in Sahul are the result of human-mediated
introductions (Aplin and Ford 2014, : table 10.3), and will not be discussed here. The
remaining extant species are recognised as comprising two distinct groups: the “Old
Endemics”, numbering at least 140 species in at least 35 genera, within the tribe Hydromyini;
and the “New Endemics”, comprising at least 20 species, all within the genus Rattus (Figure
5; Musser and Carleton 2005; Van Dyck and Strahan 2008; Flannery 1995b; Aplin and Ford
2014; Aplin 2006; Breed and Ford 2007).
Based on current fossil and molecular evidence, Murinae probably originated in
southeast Asia during the middle Miocene (Schenk et al. 2013; Jacobs and Flynn 2005; Fabre
et al. 2013). Recent large-scale molecular phylogenies indicate that the “Old Endemics” and

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

the “New Endemics” are each nested within otherwise predominantly southeast Asian clades
(Figure 5; Schenk et al. 2013; Rowe et al. 2008). This topology implies one dispersal by the
ancestor of the “Old Endemics” and another, later dispersal by the ancestor of the “New
Endemics” into Sahul, most likely from Sundaland, via Wallacea (Schenk et al. 2013; Rowe
et al. 2008; Fabre et al. 2013; Aplin and Ford 2014). Both dispersal events must have
involved crossing of marine barriers (Figure 1d; see “Geographical and geological context”
above). The presence of both “Old Endemics” (e.g. species of Melomys; Flannery 1995a)
and “New Endemics” (e.g. Rattus morotaiensis from Halmahera in the north Moluccas; Fabre
et al. 2013) in Wallacea suggests back-dispersals from Sahul by both groups. It seems likely
that further collecting and analysis of modern, subfossil and fossil murines from the region
will complicate this story still further (Aplin and Ford 2014; Aplin and Helgen 2010).
Godthelp’s (2001) report of a “Potwarmus-like ‘dendromurine’” and a second taxon that
“seems to have affinities with the widespread, primitive Chiropodomys group from South
East Asia” from the Rackham’s Roost deposit (probably early Pleistocene in age; Woodhead
et al. 2014) at the Riversleigh World Heritage Area is particularly intriguing. Today,
dendromurines occur only in Africa, whereas Chiropodomys species are found in mainland
Asia and Sundaland (Musser and Carleton 2005), and there is otherwise no Recent or fossil
record of either group from Sahul. However, these potentially highly significant taxa have yet
to be formally described.
The likely times of arrival of the “Old Endemics” and the “New Endemics” in Sahul
can be broadly constrained based on the known fossil record and molecular estimates of
divergence times. As noted by Aplin and Ford (2014), the apparent absence of murines in the
diverse Hamilton Fauna (currently estimated as 4.46 Ma; Turnbull et al. 2003) in western
Victoria suggests that they had probably failed to reach Australia by this time. The oldest
well-dated evidence of murines in Sahul is indeterminate material from the Bluff Downs

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Local Fauna in northeastern Queensland (Aplin and Ford 2014, : table 10.2), which has a
minimum radiometric date of 3.6 Ma (Mackness et al. 2000), with material from a slightly
younger (3.6-2.8 Ma) deposit on Barrow Island including representatives of modern
Australian “Old Endemic” genera (Pseudomys and Zyzomys; Aplin and Ford 2014: table
10.2). Thus, the likely time of the first entry of murines into Australia can be constrained to
between 4.46 and 3.6 Ma. Fabre et al.’s (2013, ; see Figure 5) and Schenk et al.’s (2013)
pooled molecular estimates suggest entry into Sahul by the ancestor of the “Old Endemics”
after 9.6 Ma (the maximum age of divergence from their nearest non-Sahulian sister-taxon)
but before 5.5 Ma (the minimum age of divergence between the Sahulian lineages). The fossil
record and molecular dating are both subject to potentially misleading biases (see above);
however, assuming that both sources of evidence are accurate in this case, this implies that
the “Old Endemics” were present in New Guinea for at least 1 Myr before reaching Australia
(see also Aplin and Ford 2014; Rowe et al. 2008, : 97).
Schenk et al. (2013: 852) found some evidence for an increase in diversification in a
subclade comprising the predominantly Australian Pseudomys-, Mesembriomys- and
Uromys-groups (see Aplin and Ford 2014, : table 10.1), and hypothesised that this may
reflect the first colonisation of Australia by the “Old Endemics”. If so, Schenk et al.’s (2013)
estimated divergence dates suggest that this occurred 4.8-4.2 Ma, which is nicely congruent
with the fossil evidence discussed above. Given that murines must have crossed water
barriers to enter New Guinea, a dry-land connector may have been unnecessary for at least
some lineages to reach Australia. However, dispersal of others (e.g. the arboreal rainforest
specialist Pogonomys) from New Guinea to Australia may have been facilitated by the
repeated development of land connections from the latest Miocene onwards (Figure 1d; see
“Geographical and geological context” above).

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

The arrival of the “New Endemics” in Sahul was clearly a more recent event (Figure
5). The oldest securely-dated and identified Rattus fossils from Australia are from Speaking
Tube Cave at Mt. Etna in eastern central Queensland (Cramb 2012; Cramb and Hocknull
2010), which is estimated to be ~280 ka (Hocknull et al. 2007); Rattus is notable by its
absence from older Mt Etna sites, despite their rich mammal faunas that include multiple
“Old Endemic” murine species (Cramb 2012). However, molecular estimates for the first
arrival of Rattus in Sahul are considerably older than this (Rowe et al. 2011a; Fabre et al.
2013; Schenk et al. 2013): after 3 Ma (the maximum age of divergence from their nearest
non-Sahulian sister-taxon) but before 0.85 Ma (the minimum age of divergence between the
Sahulian lineages). Again, making the questionable assumption that both the fossil record and
the molecular divergence dates are accurate, this implies that the “New Endemics” also
experienced a prolonged period (>500 ka) in New Guinea before dispersing to Australia.
The biogeographical relationships between Australian and New Guinean murine
species appear highly complex. Among the “Old Endemics”, the molecular phylogeny of
Rowe et al. (2008) implies at least nine dispersals between Australia and New Guinea (five
from New Guinea to Australia, two from Australia to New Guinea and two equivocal), while
Aplin and Ford (2014: table 10.3) identified at least 14 (11 from New Guinea to Australia,
and three from Australia to New Guinea). Among the “New Endemics”, Aplin and Ford
(2014: table 10.3) identified three dispersals, two from New Guinea to Australia and one in
the reverse direction. This pattern is perhaps unsurprising given the repeated formation and
severing of dry-land connections between Australia and New Guinea over the last 5 Myr.
However, as for marsupials (see “Marsupialiformes” above), it may be that northern Australia
and southern New Guinea (corresponding to the Austral-East Torresian zoogeographic
province recognised by Lavery et al. 2013) had an at least partially shared murine fauna for
much of the Plio-Pleistocene, intermittently divided during interglacial periods, and that

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

faunal differences developed as a result of differential extinction with the onset of more xeric
conditions in northern Australia ~280 ka. The reported presence in middle Pleistocene (>280
ka) deposits at Mt Etna of Abeomelomys (currently a New Guinean endemic; Cramb 2012)
and species of Pogonomys and Uromys that appear more closely related to New Guinean than
Australian species among living taxa (Hocknull 2005; Hocknull 2009; Hocknull et al. 2007)
is congruent with this interpretation. However, adequate testing of this hypothesis will require
considerable improvements in the Plio-Pleistocene fossil records of northern Australia and
New Guinea.
Assuming a prolonged period in New Guinea before dispersing to Australia (see
above), both the “Old Endemics” and “New Endemics” presumably radiated initially in
predominantly mesic environments. However, in Australia multiple “Old Endemic” lineages
have evolved to occupy the xeric habitats, most within the Pseudomys-group (including the
species-rich Pseudomys and Notomys), but also a member of the Mesembriomys-group,
Leporillus. By contrast, most “New Endemic” species are restricted to mesic, coastal
environments mainly in eastern Australia; however, R. villosissimus and (at least until
recently) R. tunneyi are both known from wetter areas of central Australia.
To date, only Australian murines have been subjected to phylogeographic studies.
Bryant and Fuller’s (2014) DNA sequence and microsatellite analysis of the “Old Endemic”
Melomys cervinipes, which occurs along the east coast of Australia, found evidence that a
number of recognised potential biogeographical barriers (the Brisbane Valley Barrier, the St
Lawrence Gap and the Burdekin Gap) have influenced the phylogeographic structure of the
species, as have habitat fragmentation and contraction, local extinctions and later re-
expansions. Rowe et al.’s (2011b) study of another “Old Endemic”, the Hastings River mouse
(Pseudomys oralis) found evidence of two major mitochondrial lineages that appear to have
diverged 300-900 ka, with the area of lineage overlap corresponding to the northern limit of

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

the Macleay–McPherson Overlap Zone (Burbidge 1960; Ebach et al. 2013). Strikingly,
however, there is no indication from Rowe et al.’s (2011b) results that the Brisbane Valley
Barrier further north has posed a barrier to gene flow in Pseudomys oralis, contra Bryant and
Fuller’s (2014) findings regarding Melomys cervinipes. Although many more
phylogeographic studies are required, this hints at the likelihood that biogeographical patterns
within Australian (and probably also New Guinean) murines are likely to be highly species-
specific, as they appear to be in marsupials.
Like small- and medium-sized marsupials, the native murines of Australia have
suffered severely in post-European colonisation, with the introduction of invasive placental
species (particularly cats and foxes) and widespread habitat modification (Johnson 2006;
Burbidge et al. 2008; McKenzie et al. 2007). As a result, several species have gone extinct
and many others have seen enormous range reductions. New Guinean species also appear to
have experienced extinctions and range reductions, particularly on islands off the New
Guinean mainland (Aplin et al. 1999a). As already discussed, the loss of entire populations
(at least some of which likely represent evolutionarily distinct lineages) is likely to misled
biogeographical studies that are restricted to extant representatives of a particular species.
Again, inclusion of distributional data (as in Burbidge et al. 2008 and Lavery et al. 2013) and
(if possible) ancient DNA from subfossil and fossil specimens will be critical for an accurate
understanding of the biogeographical history of Sahulian murines.
4.8 Questionable records
Clemens et al. (2003) described a fossilised vertebrate tooth (AMF 118621) from the Albian
Griman Creek Formation at Lightning Ridge in New South Wales, as a probable upper molar
of a “dryolestoid” mammal. The oldest “dryolestoid” from South America is Cronopio

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

dentiacutus from the Cenomanian La Buitera locality in southern Argentina (Rougier et al.
2011a), and “dryolestoids” were highly diverse in the Late Cretaceous Allenian faunas of
southern South America (Rougier et al. 2011b; Rougier et al. 2009a; Rougier et al. 2009b;
Bonaparte 1990). Thus, the apparent presence of at least one “dryolestoid” in Australia
during the middle Cretaceous would not be surprising, and would provide the closest link yet
between the Mesozoic mammaliaform faunas of Australia and South America, which
otherwise appear to show high level endemism (Rich 2008; Rich et al. 2009a). However, it is
unclear whether AMF 118621 is indeed a “dryolestoid”, or even a synapsid. Perhaps most
concerning is its very large size: at 12.2 mm long by 14.0 mm wide, it is far bigger than the
upper molars of any definitive Mesozoic dryolestoid (Clemens et al. 2003). Furthermore, as
noted by Clemens et al. (2003), some crocodyliforms are known to have evolved complex,
superficially mammal-like cheek teeth (including Cretaceous forms from Gondwana;
O'Connor et al. 2010). It is entirely possible that AMF 118621 does not represent a
mammaliaform.
Mastodon australis and Notelephas australis were described and identified as
proboscideans by Owen (1845, 1882) based on specimens apparently collected from
Australia: M. australis from “further in the interior than [the caves] of Wellington Valley”
(presumably inland New South Wales) and N. australis from the Darling Downs in
southeastern Queensland. Among mammals, proboscideans have excellent overwater
dispersal capabilities (van der Geer et al. 2010; Johnson 1980): in 1856, an elephant was
reported to have swum 48 km to land after falling overboard in the Atlantic (Johnson 1980),
and remains of the fossil proboscidean Stegodon are known from multiple islands in
Wallacea (Dennell et al. 2014; van der Geer et al. 2010), undoubtedly the result of overwater
dispersal(s). However, Stegodon fossils have not been found east of Lydekker’s Line, and
their apparent absence from New Guinea is perhaps the biggest argument against their

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

reaching Australia, at least as live individuals. The decaying, gas-filled carcass of a large-
bodied, thick-skinned mammal such as a proboscidean might be capable of being transported
prolonged distances overwater (Archer 1999), but this would not explain why Mastodon
australis at least was reportedly found far inland. Owen’s (1845, 1882) identification of these
specimens as proboscideans has not been questioned (Archer et al. 1999b), but doubts have
been raised to whether they were genuinely collected in Australia (Longman 1916) . Pending
careful reassessment of the known material and reported provenance of Mastodon australis
(known from a single molar, apparently now lost) and Notelephas australis (known from a
fragmentary tusk held at the Natural History Museum, London), or the discovery of
additional specimens, they will remain biogeographical enigmas.
5 Some concluding thoughts: problems and prospects
As the above summary should make abundantly clear, there are numerous areas of
uncertainty in our understanding of the biogeography of mammaliaforms in Sahul, most of
which are due to a sheer lack of evidence. Inevitably, this leads to the all-too-familiar cry of
“more data is required”. But what precisely is needed, and how likely is this to be obtained?
Our understanding of the complex geological history of Sahul, as well as related aspects such
as sea level and palaeoclimate, will undoubtedly increase with further study. Ever improving
sequencing technology will allow researchers to obtain sequence data from multiple
individuals of extant taxa increasingly quickly and cheaply. Remarkable advances in methods
for extracting and sequencing ancient DNA allow sequence data to be acquired increasingly
effectively from subfossil and fossil remains, with the latest methods successfully obtaining
genuine sequences from specimens >400 ka (Orlando et al. 2013; Dabney et al. 2013; Meyer
et al. 2014). If such methods can be successfully applied to subfossil and fossil remains from

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Sahul, they will open up entire new vistas of research, enabling extinct species and
populations to be incorporated directly into phylogenetic and phylogeographic analyses based
on molecular data (e.g. Stiller et al. 2014; Austin et al. 2013; Sheng et al. 2014). If ancient
DNA proves unobtainable from particular specimens, methods that obtain amino acid
sequences from proteins such as collagen may still allow phylogenetically useful molecular
data to be acquired (Buckley 2013). Ongoing collecting is likely to result in the discovery of
new mammalian species and also additional populations of currently known species,
particularly in New Guinea and adjacent islands (Helgen 2007), which is likely to lead to
revision and refinement of current biogeographical hypotheses.
Improvements in the Sahulian fossil record, however, may prove more elusive,
particularly for the Mesozoic and early Palaeogene. Despite continuing exploration, the entire
published Mesozoic record of mammaliaforms in Australia is from four middle Cretaceous
localities (Kielan-Jaworowska et al. 2004; Long et al. 2002; Rich and Vickers-Rich 2004;
Rich et al. 2009b): the Aptian Dinosaur Cove, Flat Rocks and Eric the Red West sites in
southern Victoria, and the Albian Griman Creek Formation at Lightning Ridge in northern
New South Wales (Figure 3a). Similarly, there is only a single early Palaeogene mammal-
bearing fossil site known in Australia, the early Eocene Tingamarra Local Fauna (Figure 4a;
Godthelp et al. 1992; Long et al. 2002), to fill the 85 million year old gap between the four
Aptian-Albian sites and the richer fossil record of mammals known from the late Oligocene
onwards. Given that these pre-Oligocene sites are so few in number and that only a handful
of mammalian taxa that have been described from them to date, it is unclear whether the
apparent absence of particular groups from these sites (e.g. the apparent lack of monotremes
at Tingamarra) is an accurate reflection of the mammaliaform fauna of the time, or simply an
artefact of the very limited sampling to date. Thus, biogeographical inferences drawn from

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

the pre-Oligocene mammaliaform record of Australia should be treated with particular
caution.
Progress is also likely to come from the development and use of new, quantitative
methods of phylogenetic and biogeographic analysis. Particularly promising are methods that
allow the phylogenetic relationships and divergence times of fossil and extant taxa to be
calculated simultaneously (Pyron 2011; Ronquist et al. 2012), although the accuracy of these
methods remains to be fully established (Beck and Lee 2014). Also of interest are new
parametric methods for analysing biogeography within an explicitly statistical, model-based
framework (Ree et al. 2005; Ree and Smith 2008; Matzke 2013, 2014; Ree and Sanmartin
2009). Such methods: 1) permit both vicariance and dispersal; 2) take into account
divergence times/temporal branch lengths and (if specified) changes in connectivity between
areas through time; 3) allow the best fitting model(s) to be identified using explicit model
selection criteria. These methods are starting to be applied to Sahulian mammaliaform clades
(Fabre et al. 2013; Mitchell et al. 2014; Westerman et al. 2012).
As noted repeatedly throughout this review, incorporation of evidence from the fossil,
subfossil and historical record – whether simply as distributional data, or via inclusion of
extinct taxa in phylogenetic and phylogeographic analyses, ideally in the form of ancient
DNA sequences – is likely to be critical for accurate reconstruction of biogeographical
histories (see also Lieberman 2002; Buerki et al. 2013). Specifically, complete extinction of
species or populations may explain biogeographical patterns that are otherwise hard to
explain, such as the “old” New Guinean marsupial clades found by Mitchell et al. (2014) and
Tasmanian platypus clade found by Gongora et al. (2012). Thus, active collaboration between
palaeontologists and researchers working on living taxa is likely to prove at least as important
as simple improvements in data and methods.

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

The prospects are therefore good for major improvements in our understanding of at
least some areas of Sahulian mammaliaform biogeography. At the same time, however, the
modern mammal fauna of Sahul is under threat, with numerous species experiencing severe
and ongoing declines (Woinarski et al. 2011; Woinarski et al. 2010; George 1979). Let us
hope that current and future biogeographical studies do not become monuments to yet more
vanished populations and species.
6 Acknowledgements
My thanks to the editor, Malte Ebach, for inviting me to contribute this review. I thank
Guillermo Rougier, Erich Fitzgerald, Tom Rich, Kris Helgen and John Schenk for discussion
and for supplying me with references. Pierre-Henri Fabre, Mike Archer, Peter Trusler, Ken
Aplin and Fred Ford generously provided me with images that have been used in some of the
figures. I am particularly grateful to Scott Hocknull for his thorough and constructive review.
My thanks also to Sue Hand, Rebecca Pian, Julien Louys, Karen Black and Mike
Woodburne, all of whom read earlier drafts of this chapter and gave extremely helpful
comments. Financial support for my research on Australian mammaliaform biogeography has
been provided by the Leverhulme Trust (via Study Abroad Studentship SAS/30110), Phil
Creaser and the CREATE fund at the University of New South Wales (via a CREATE
scholarship), the National Science Foundation (via grant DEB-0743039, in collaboration with
Rob Voss at the American Museum of Natural History), and the Australian Research Council
(via Discovery Early Career Researcher Award DE120100957).
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Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

Figure 1. Circum-Antarctic palaeo-depth models for the seafloor at 61, 52, 43, 25, 10 and 5
MYA, illustrating the opening and development of the Drake Passage (between South
America and Antarctica) and the Tasmanian Gateway (between Antarctica and Australia).
Reprinted from Palaeogeography, Palaeoclimatology, Palaeoecology 231, Brown et al.,
“Circum-Antarctic palaeobathymetry: Illustrated examples from Cenozoic to recent times”,
pp 158–168, Copyright 2006, with permission from Elsevier.
Figure 2. a-c Palaeotectonic evolution and emergence of New Guinea over the last 10 Ma
(green, land; dark blue, deep sea; lighter blue, shallow sea; red white brick, calcareous
plateaus possibly exposed at times; orange, highland; grey, high altitude above 2,800 m).
Reprinted by permission from Macmillan Publishers Ltd: Nature Communications 5,
Toussaint et al., “The towering orogeny of New Guinea as a trigger for arthropod
megadiversity”, Copyright 2014. d Geography of Sahul region showing maximum land
extent during Pleistocene glacial maxima. Modified from Aplin and Ford (2014: fig. 10.1).
Lydekker’s Line is indicated.
Figure 3. a Locations of known Mezozoic mammaliaform-bearing fossil sites in Australia (1,
Lightning Ridge, northern New South Wales [Albian]; 2, Dinosaur Cove, southern Victoria
[early-middle Aptian]; 3, Eric the Red West, southern Victoria [early-middle Aptian]; 4, Flat
Rocks, southern Victoria [early-middle Aptian]). b Composite phylogeny of
Australosphenida, based on Rougier et al. (2007) and Bi et al. (2014). Terminals are colour-
coded according to their known biogeographic distributions. Extinct taxa are identified by
daggers. Note that the Sahulian terminals (in red) form a single clade to the exclusion of non-
Sahulian terminals (see main text). c Holotype of the monotreme Steropodon galmani (AM

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

F66763, partial right dentary preserving m1-3) from Lightning Ridge (see Archer et al.,
1985). d Holotype of Kollikodon ritchiei (AM F96602, partial right dentary preserving m1-3)
from Lightning Ridge (see Flannery et al., 1992). e Holotype of monotreme Teinolophos
trusleri (NMV P208231, partial left dentary preserving m?2) from Flat Rocks (see Rich et al.,
1999; 2001; 2005). Illustration - P. Trusler. f Holotype of ?cimolodontan multituberculate
Corriebaatar marywaltersae (NMV P216655; partial left dentary preserving p4 and anterior
root of m1) from Flat Rocks (see Rich et al., 2009). Illustration - P. Trusler. g Holotype of
ausktribosphenid Ausktribosphenos nyktos (NMV P208090, partial right dentary preserving
p6 m1-3) from Flat Rocks (see Rich et al., 1997). Illustration - P. Trusler. h Holotype of
aukstribosphenid Bishops whitmorei (NMV P210075, partial left dentary preserving p1-6 m1-
3 (see Rich et al., 2001). Illustration - P. Trusler.
Figure 4. a Location of Australia’s only early Palaeogene mammal-bearing fossil site, the
early Eocene Tingamarra Local Fauna, near Murgon in southeastern Queensland. b
Phylogeny of Metatheria based on maximum parsimony analysis of a 260 morphological
character matrix with relationships constrained using a “molecular scaffold”, modified from
Beck (2012: fig. 6a). Terminals are colour-coded according to their known biogeographic
distributions. Extinct taxa are identified by daggers. Note that the Sahulian terminals (in red)
do not form a single clade to the exclusion of non-Sahulian terminals, and hence the ‘single
dispersal’ model for the presence of marsupialiforms in Sahul can be rejected (see Beck,
2012 for full details). c Holotype of the bunodont marsupialiform Chulpasia jimthorselli (QM
F50411, left M1 or 2) from Tingamarra (see Sige et al., 2009). d Holotype of the bunodont
marsupialiform Chulpasia mattaueri (CHU 30, left M1 or 2) from the ?late Palaeocene
Chulpas locality, Laguna Umayo red mudstone unit, Peru (see Sige et al., 2009). e Holotype
of the faunivorous marsupialiform Archaeonothos henkgodthelpi (QM F53825, left M2 or 3)

Beck, R. M. D. (accepted). The biogeographical history of non-marine mammaliaforms in the Sahul region. In
M. C. Ebach (Ed.), Handbook of Australasian biogeography. Boca Raton, Florida, USA: CRC Press.

from Tingamarra (see Beck, 2014). f Calcaneus of an “ameridelphian” marsupial (QM
F30060) from Tingamarra (see Beck, 2012). g Holotype of the bat Australonycteris clarkae
(QM F19147, left m?2) from Tingamarra (see Hand et al., 1994). h Holotype of the
?condylarth eutherian Tingamarra porterorum (QM F20564, right lower molar) from
Tingamarra (see Godthelp et al., 1992). i Holotype of the australidelphian marsupial Djarthia
murgonensis (QM F31458, left partial dentary preserving m2-4) from Tingamarra (see
Godthelp et al., 1999; Beck et al., 2008).
Figure 5. Phylogeny of murid rodents based on a Bayesian relaxed molecular clock analysis
of one mitochondrial (cytochrome b) and two nuclear (growth hormone receptor exon 10;
interphotoreceptor retinoid binding protein exon 1) protein-coding genes using BEAST,
modified from Fabre et al. (2013: fig. 3).Branches are colour-coded according to their known
biogeographic distributions, and the predominantly Sahulian “Old Endemics” and “New
Endemics” are indicated (see Fabre et al., 2013 for full details).

SAHUL
Madagascar
South America
h
abc d
ef
g

ab
Eurasia + North America
Eurasia
South America
North America
SAHUL
cd
ef
g
h
i
South America + North America