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Full text of "Memoirs of the Queensland Museum"

MEMOIRS 




OF THE 



QUEENSLAND MUSEUM 





BRISBANE 
30 JUNE 1997 



VOLUME 41 

Part 2 



THE PALAEONTOLOGY AND GEOLOGY OF DUNSINANE SITE, RIVERSLEIGH 

DERRICK A. ARENA 



Arena. D.A. 1997:06:30: Palaeontology and geology of Dunsinane Siie, Riversleigh.A^fm- 
oirsofthe Queensland Museum 4l{2): 171-179. Brisbane. ISSN 0079-8835. 

The stratigraphy, fossil assemblages and models of formation and post-deposilional history 
of Dunsinane Site, on the western 'arm' of southern Gag Plateau, are described. The fossil 
vertebrate fauna most resembles the late Oligocene (System A) fauna of White Hunter Site. 
Rich, exceptionally preserved invertebrate and plant (twigs, leaves, reproductive organs) 
assemblages occur in nodules of iron oxide-rich ilourapatite and as fragments derived from 
these nodules and appear to have been preserved by early diagenetic microbially-medialed 
phosphatisation. The sediment is most likely lacustrine and may directly overlie Precambrian 
sediments. Erosion of overlying sediments and severe chemical weathering may have led to 
development of the surface lag of insoluble residue which includes phosphatised bone, 
nodules and other fossil fragments. Relationships of the fossiliferous nodules to the vertebrate 
fauna and surrounding sediments are not fully determined. 

Derrick A. Arena. School of Biological Science. University of New South Wales, New South 

Wales 2052, Australia; received 10 November J996. 



Dunsinane Site was discovered in 1990 during 
exploration of southern Gag Plateau at 
Riversleigh, NW Queensland (Fig. I A). In one 
area the surface was strewn with fragments of 
fossil bone, fossil wood and nodules of rock that 
contained leaves, wood, other plant material and 
invertebrates. Dunsinane Site is the only fossil 
site at Riversleigh containing plants and one of 
the few yielding arthropods (Archer et al., 1994). 
The site was also distinctive in that the fossils 
were associated with a soft, apparently un- 
consolidated sediment, rather than embedded in 
solid limestone. Issues to be resolved regarding 
this site included the nature and provenance of the 
fossilised plant and arthropod material, and the 
relationships of this material to other Riversleigh 
sediments. 

CONCEPTS AND TERMINOLOGY 

Palaeontological concepts, biostratigraphy and 
laxonomic classification follow Archer et al. 
( 1 994) and Greaser ( 1 997). The terms 'Dunsinane 
limestone', 'Dunsinane deposit', 'Dunsinane 
sediment' and 'Dunsinane calcrete' are used here 
informally. 

STRATIGRAPHY AND GEOLOGY 

Representative rock types from Dunsinane Site 
have been examined by powder X-ray diffrac- 
tion, SEM and thin-sectioning techniques. A 
sample of 4 nodules was used for destructive 
analysis. The area around Dunsinane Site is dom- 
inated by 4 main rock types: Precambrian quartz- 



ite, ferruginised deposits, overlying Tertiary 
limestone and the Dunsinane limestone. 

PRECAMBRIAN QUARTZITE 

The grey Precambrian quartzite is a massive, 
thick tabular- bedded, crystalline pure quartz sed- 
iment. This and a laminated chert constitute the 
teirain around Gag Plateau and form the base- 
ment underlying the Dunsinane deposit. 

FERRUGINISED DEPOSITS 

Outcrops of ferruginised deposits generally do 
not exceed 15m in diameter. The ferruginisation 
is apparently related to localised groundwater 
activity. This type of deposit occurs throughout 
the Riversleigh area, particularly along geologi- 
cal boundaries. Around Dunsinane Site such de- 
posits occur in the Precambrian quartzite, 
overlying Tertiary limestone, and at a point at the 
junction of the Precambrian quartzite and the 
Dunsinane sediment, apparently post-dating all 
of these sediments. In general they consist of 
pisolitic iron oxide, and iron oxide- enriched al- 
terations of the sediments in which they occur. 

OVERLYING TERTIARY LIMESTONES 

The hard, micritic. Tertiary limestones directly 
overlie the Dunsinane limestone. These lime- 
stones exhibit vertical changes in colour and fre- 
quency of molluscs and calcareous mud clasls, all 
of which may be construed as primary bedding 
features. Occasional vertebrate bone fragments 
occur with molluscs which are very common and 
well-preserved. Contact of the Tertiary Lime- 
stone with the underlying Dunsinane sediment is 



172 



MEMOIRS OF THE QUEENSLAND MUSEUM 





FIG. 1. A, Southern Gag Plateau and Dunsinane Site Local area map. Study area in B outlined. Contours at 
mAHD. (Southern Gag Plateau map inset from Megirian 1992). B, Dunsinane Site study area indicating the 
positions of Dunsinane, Bernie's Cooking Pot (BCP), Custard Tart (CT) and Sue's Rocky Road (SRR) Sites. 
Siipling=Dunsinane limestone. Contours metres from arbitrary field datum. 



apparently undulalory, and may be unconform- 
able. 

DUNSINANE LIMESTONE 

The Dunsinane limestone ouicrop is ai least 1 50 
m across. It includes the fossil sites Bernie's 
Cooking Pot (BCP), Custard Tart (CT) and Sue's 
Rocky Road (SRR) (Fig. IB), which have pre- 
viously been regarded as separate sites. 

The limestone has been severely weathered, 
resulting in replacement of most of the original 
sediment and its sedimentary structures by soft 
calcrete. Relict unweathered outcrops occur as 
sandy limestone with apparent ilat bedding and 
minute bone fragments. Because the overlying 
limestone is closely associated with these relict 
outcrops, it appears that the weathering event 
post-dates the deposition and erosion of the over- 
lying sediment. 

Because the fine, well-sorted particles and ap- 
parent flat-bedding does not indicate high energy 
flow, the Dunsinane sediment was probably de- 
posited in a pond or lake. This relatively arena- 
ceous sediment was probably deposited under 
near shore conditions in a quartz-rich Precambr- 
ian terrain. 

Fossil bones and nodules are preserved with 
brown iron oxide-rich flourapatite (fluorapat- 
ite=calcium fluoride phosphate; Ca5(P04)3F). 
Energy dispersive X-Ray spectrometry demon- 
strated that iron oxide appears to be included 
within euhedral flourapatite crystals. Flourapatite 
has replaced the majority of structures within 



nodule matrices. Bones are impregnated with 
iron-oxide rich flourapatite, particularly around 
the inner parts. Bones and nodules occur in relict 
in situ sediment and in the surface layers across 
the area of the outcrop. Nodule fragments and 
fragments of fossil wood derived from weathered 
nodules are common. There are also amorphous 
concretions of tlourapalite and sparry calcitc 
deep within the calcrete (Im depth) in some 
places. 

MODEL OF POST-DEPOSITIONAL HJS- 
TORY 

Weathering is a striking feature of the Dunsin- 
ane sediment. The calcrete is most likely autoch- 
thonous because of the relationship between 
relict unweathered outcrops of Dunsinane sedi- 
ment and overlying rock. Local groundwater ac- 
tivity has resulted in ferruginisation in the 
vicinity, although this has not occurred at Dunsin- 
ane Site itself. Meteoric and surface water are the 
most likely agents responsible for chemical 
weathering of the original Dunsinane sediment. 

Severity of weathering has been influenced by 
permeability of the original Dunsinane sediment, 
impermeability of the Precambrian quart/ite and 
Tertiary limestone, the situation oi^ Dunsinane 
Site at the junction of 3 drainage channels, and its 
position near the top of the local drainage net- 
work. 

Water focussed on Dunsinane Site by the local 
drainage system was channelled into the perme- 
able Dunsinane sediment before draining away 



DUNSINANESITE 



173 



into the lower calchment. This caused the chem- 
ical weathering of the limestone responsible for: 
1. apparent subsidence, slumping and surface 
lowering of the sediment; 2, accumulation of a 
surface lag of fossils mineralised with insoluble 
llourapatite; 3, partial dissolution and precipita- 
tion of flourapatite bodies deep in the original 
sediment; and 4, a thin calcareous duricrust which 
has cemented surface debris. 

FLORA AND FAUNA 

VERTEBRATES. Due to weathering and poor 
condition of the fossilised bone, identifiable ver- 
tebrate fossils from Dunsinane Site are relatively 
uncommon. However, there is marked diversity 
implied by the number of groups present. In con- 
trast with many Riversicigh sites, fish and bird 
remains have not yet been found at Dunsinane 
Site. The Vertebrates includes: 

Class Rcptilia 

Order Testudines 

Family Chelonidae 

genus & sp. indet. 

Order Crocodilia 

Family Crocodyhdae 

'!Baru sp. 

Class Mammalia 

Order Marsupiaha 

Suborder Diprotodonlia 

Family Wynyardiidac 

INamilamadeta sp. 

Family Diprotodonlidae 

Subfamily Diprolodontinae 

'^Bematherium sp. 

Subfamily Zygomalurinae 

"INeohelos sp. 

Family Macropodidae 

Subfamily Balbarinae 

Nambaroo sp. nov. 

genus & sp. indet. 

Subfamily Balungamayinae 

IWabularoo sp. 

Order Placenlalia 

Suborder Chiroptera 

Family Hipposideridae 

IBrachipposideros sp. 

Neohelos. Brachipposuieros, large crocodil- 
ians and chelonid turtles which arc known from 
Riversicigh .sediments oi' various Oligocene and 
Miocene ages are thus not useful for more precise 
biocorrelation. Al Riversicigh wynyardiids are 
known only from System A and B faunas (Archer 
ei al.. 1994). Primitive balungamayme and 
balbunne kangaroos such as IWahidaroo and 



Nambaroo respectively (the latter appears to be 
conspecific with a very primitive fonn from Sys- 
tem A While Hunter Site (Cooke, 1997)), arc 
known from Riversicigh System A and B faunas. 
With the exception of Pleistocene Diproiodon 
optatum , diprotodonlines are restricted to System 
A at Riversicigh (Black, 1997). Thus we regard 
the fauna as most likely a System A fauna. 

INVERTEBRATES 

Insect fragments are often quite small, usually 
around 1 mm and include a variety of beetle elytra 
and beetle prothoraces which show similarities to 
those of curculionids (weevils) and buprestids 
(jewel beetles). Curculionids have been found in 
the Upper Site assemblage (Archer et al., 1994), 
although the long history of this group makes 
them of little biostratigraphic use. Termite re- 
mains may also be present. 

A partial gastropod shell has been identitied as 
a probable terrestrial camacnid (W. Ponder pers. 
comm. 10/95). Camaenids which are known trom 
the Mesozoic occur in System A (Archer et al., 
1994) and are likely to occur in Riversicigh sed- 
iments of all ages. 

FLORA 

The plant assemblage consists of small pieces 
of twig-like wood (some stems may originally 
have been 3~4cm in diameter), leaves, seeds and 
reproductive organs. 

The various types of wood imply high diversity. 
Angiosperm wood is present as well as probable 
gymnosperm wood. Leaf fragments are invari- 
ably broad and serrate-margined. Leaf cuticles 
are intact in a number of specimens. 

There are several pneumatophore-like organs 
up to 2cm in diameter with a central aerenchyma 
of elongate cells and a narrow epidermal layer 
with structures resembling lenticels. 

Groups recognised include Proteaceae, 
Casuarinaccae. Myriaceae and possibly 
Epacridaceae (R. Hill pcrs. comm. 9/95, 10/96). 
Proteaceae are evergreen trees and shrubs, and 
are known from the Early Cretaceous to 
Holocene (Hill, 1994). The generally 
xeromorphic Casuarinaccae have a record from 
the Paleocene to Holocene (Hill. 1994). 
Myrtaceae are known from the mid-Palcocene to 
Holocene (Martm, 1994). The Epacridaceae 
which are prommenl in extant scleromorphic flo- 
ras have existed in Australia since the Late Cre- 
taceous (Jordan & Hill, 1996). 

Serrate leaves are not good indicators of a trop- 
ical closed forest origin (R. Hill pers. comm. 



74 



MEMOIRS OF THE QUEENSLAND MUSEUM 




DrN5;iNANESITE 



175 



9/95). Apparent growth rings und false growth 
rings are evident in some woud samples. Some 
rings arc bmad. inferring a long growth season 
which seems lo be icnninalcd abruptly (R. Hill 
pcrs. comtiv 10/95). !n general the associated 
fli>ristics and the liming of fossil {)Ccurrence.s o\' 
epjicrids throughout the Tertiary coincide with 
temperate climatic conditions, and the nature of 
the inacrolbssil record is inct»nsisienl with mod- 
ern tropical or sub-iropicul rainlbrest (Jordan & 
Hill. 1996). Pollen has been recovered from the 
nodules hut not yet analysed. 

palaeoenvironmental 
implications 

Biocorrelalion oj the Dunsinanc Site fauna with 
that of White Hunter Site (via Nombaroo and 
"^Heinathehntv) allows tentative age assessment. 
White Hunter Site has been correlated wiih 
liladunna Fauna! Zone D (Ngama LF. Lake Pal- 
ankarinnal on the ilariid, Kittennfjn nguma 
(Myers & Archer, IS>97). Sediments associated 
with this South Australian fauna have iKen pal- 
aeomagneiically dated at 24.7-25 Ma (Late 
Oligocene; Woodburneei al.. 1994), This interval 
et»iitcides with an 'icehouse' event. clJa»atic con- 
ditions normally characterised by ccxiler, drier, 
seasonal climatic conditions (Frakoseial.. 1987) 
The 'greenhouscV Scehouse* ciimaiic fluclua 
lions \^i the Tertiary arc rellectcd in the -sedmten- 
tary and lerresiriul fossil records of Australia 
(Frakes et al.. 1987; Archer cl al.. 1995). The 
characierisiics of the Dunsinanc Site fK>ra so far 
observed may indicate 'icehouse' conditions. 

TAPHONOMV 

PHOSPHATESATION OF FOSSIL ASSKM- 
BLAGE 

Early diagenelic phosphalisalion. uitually <tss0- 
ciaied with microbial activity, has been identified 
as the mode of preservation ol phosphatic nod- 
ules and excepiionallv preser\ed fossils li.e. Bal- 
son. I9KU; Miiller. i9K5: Pinna. 1985: Seilachcr 
CI al., I9H5; Soudrv ^ Lewv. 1988; Allison, 



1988a, b, c; Martill. 1988, 1989, 1990: Lucas & 
Prcvoi, 1 99 1 ; Briggs St Kear, 1993; Briggs el al.. 
1993). As confirmed by laboratory cxperimiMM^ 
(i.e. Prcvot & Lucas, 1986; Hirschl'eret al.. I9W: 
Briggs et aL 1993: Briggs & Kear. 1993). 
microbiaily mediated phosphalisalion can occur 
within or adjacent to bacteria, and can result in 
the formation of globular apatite microstruciurcs 
that faithfully preserve the structure of organ- 
isms. Nomeiimes at ihc microscopic level. Condi • 
lions under which flouraputiic replacement of 
organic tissue and carbonates may occur are 
(Lucas & Pr(<v6i, 1991); Da concentration of 
organic phosphorous is required in the syslerii 
(i.e. the sedimcni); 2) anoxic conditions which 
support bacteria capable of precipitating apatite; 
3) acidic conditions that destabilise carbonates; 
Ihesecircunislanccsctmimonly occur iniheinler- 
stiiia of phosphate-rich sediments. These condi- 
tions may be enhanced by closure' by a thin film 
ofsedimentorbaclcrial slime, or enclosing struc- 
tures of organisms or sediment (i.e. carapaces, 
pore spaces) that contain the optimal environ- 
ment for apatite precipitation (Krajewski, 1984; 
Seilacherelal.. 1985; Martill. 1988, I989J990; 
Soudry & Lewv. 1988; Hinschlcr et al.. 1990: 
Lucas & PrcvcV, 199L Wilhv & Martill I9<^; 
Briggs et al., 1993; Briggs & Kear, 1993). 

Characteristic tlourapatile microslrucluivs 
very simikir to those found in fossils from else- 
where in the world thought to be preserved in this 
way occur in nodule malcrial examined by SF-M 
(Fig. 2G.H). The lack of di.stortion or crushing uf 
tissues, preservation of the cellular structures of 
leaves (see below ) and presence of organic mate- 
rial in the Dunsinane nodules indicate early dia- 
geneiic. pre -compaction mineralisation. This 
process is suggested as the mode of preservation 
for the 3D arthropod fossils from Upper Site 
(Duncan & Bnggs. 1996). Fragments of algal 
layers in nodule material may indicate thin micro- 
bial filtiis that Nealed* tissues, enclosing condi- 
tions favourable to preservation and encouraging 
and accelerating mmtTalisalion The Dunsinane 
nodule material may have mineralised in a water 



I'lti. 2. A, Dunsinane Site nodules illusiraimg the vanctv of shupcs and sizes. Sca\c bar=!icTn. B. Sonic nodules 
have prolusions (m this case a piece of ftvssil wcK>d). Scale=20mm. C, Leaf fragnicni, D, QMF3I.^()<), a 
Vdiincnsiona! Fruit lenlaliveh assiened to (he Epacridijceae. E. Thin section ot nodule, enlirel) phosphaiiscd 
organic moicrMl Note the undiskmcd arthropod with cuiicle at Joucr-right, Plane polarised light. Sca^ 
r. Transverse st-ction o!" a leaf lamina. The vertically efongytcd cells are palisade cells, beneaih them is imaci 
spongy mesophyil tissue. Plane polarised Itghi. Sc;ile=2(Xl|jini. G, SVM of nodule material showing ihc 
eliur,jcieri>lic microslnicUirc uT early diagenelicull> precipitated nourapaliie replacing organic lisMic. 
Hcalc=20M-!n IL Glohos*? noiirupfitile microsiruclurcs. Ihe p.seudo-hexiigonal cf^stuLs arc flourapatJlc. 
ScaJe=5^lln. 



176 



MEMOIRS OF THE QUEENSLAND MUSEUM 





FIG. 3. Models of formation of the 
Dunsinane Site nodules. A-C 
showing models of the arrival/for- 
mation of nodules. A, Whilst there 
is little evidence of high-energy 
activity in the sediments, erosion 
and transport cannot be ruled out as 
a rounding mechanism. B, Nodules 
may have formed by partial 
mineralisation of a mat of organic 
material. C, Nodules may be de- 
rived from a weathered overlying 
deposit in which they either formed 
or into which they were trans- 
ported. This model is regarded as 
relatively unparsimonious consid- 
ering the partially phosphatised 
vertebrate assemblage is unlikely 
to have been reworked and that 
nodules occur in in situ Dunsinane 
site limestone. Once incorporated 
into the sediment, the '?unconform- 
ably overlying younger Tertiary 
limestone was deposited (D,E). 
Development of a karst terrain (F) 
and erosion of overlying sediment 
allowed weathering of the Dunsin- 
ane deposit (G). Calcretisation 
(medium stippling) and surface 
lowering resulted in the observed 
distribution of outcrops of relict 
and weathered Dunsinane lime- 
stone, overlying Tertiary limestone 
and a residual lag of insoluble fos- 
sils. 



DIJNSINANESITE 



177 



hodj thai had Ma^^niUcd possibly because ul a lack 
of Ircshwaicr input; a proximal acidic anoxic 
sediment such as peal or inHuKcs of nulrienls 
causing microbial bUioms that used up available 
oxygen and increased available organic pliospho- 
rous. As partially decaying organic matter accu- 
mulated on and in the substrate, reducing 
con(litit)ns wouM develop leading lo accumula- 
tion ol iron hydroxides ami dcslabilisation oi 
carbonates (Allison I^^SMblandaprolifcralionor 
anaerobic nncrubes. Phosphaiisalion may have 
occuned in tissues in the substrate and close to it. 
until hahed by exhaustion of available phosphate 
supplies, or dilation of the phosphaierieh me 
dium perhaps by an inHux of freshwakT supply- 
ing tixygcn and dissolvtjd euJcjum carbonaic 

PLANT ASSEMBLAGE 

Plant organ assemblages composed of twigs, 
leaves and reproductive organs such as those ui 
Dunsinanc Site (Fig. 2C,D) are generally re- 
garded as accumuJahons with limited lateral wmd 
transport (Colhnson. l983;Spicer. 1980. 1989}. 
Plant assemblage?; accumuliiied at the site of 
gTowUi usually contain roots, wood fragments 
and plant bases; catasin*phic aeeumulaiions are 
poorI>' sorted and contain comfxjncnts of ail types 
(Cullinson, l9S3;S|>icer. 1989). 

Leaves nio.si likely to be preserved in excellent 
condition are those that Tail directly into water, 
since coniaci wiih the ground usually dritsiically 
reduces the chances of leaf preservation in an 
aquatic environment (Ferguson, 1985; Spiccr. 
1989, 1991). Dissolution of anti-fungal, anti-rni- 
erobial and structural compounds within leaves 
begins immediately after immersion or contact 
with the ground, increasing susceptibility to mi- 
crobial aiiack and structural collapse within 24 
hours (Ferguson. 1985;Spiccr, I9S9. 1991 ). l^af 
liner is generally dispersed within S{)m of the 
parent vegetation (Ferguson. 1985; Spleen 1989, 
1991 1. Hill & Gibson (1986) found thai the ma- 
jority of leaves collected Irom a lake lied were 
from species occurring wiihin 50m of the shotc. 
Leaves in the Dunsinanc assemblage with limited 
immersion damage, such as the exceptionally 
pnL*served leaf with iniaci cellular detail (Fig. 
210. may have been derived from vegetation thai 
oceuired within 50in of the site of pies^'tvaiion. 
Since most leaves lake hours or dav.s to sink 
(Ferguson. 1985; Will & Ciibson. 1986; Spicer, 
19S(). 1989. 1991). buoyant organs such as twigs 
and seeds are prevalent in the assemblage and arc 
ofien in quiie >;oO(I coiuliiion showing little abra- 
sion or decomposition, and tlie insect material is 



so plentiful and diverse these factors may point 
to the accumulation bemg m shallow nearshore 
water. 

The plant material thus^ appears to be part of a 
proximal assen)l)lage. possibly occurring as an 
immersed mat of vegetation. The process ol 
mineralisation that preserved the plant material 
nuisi have occurred in the early stages of degra- 
dation smce leaves have limited strmttural dam- 
age and intact cellular siruciurc. in particuliirly 
spongy mesophyll tissue which is generally the 
most susceptible to breakdown (Spicer. J 989). 
The pneumalophore-likc organs in the nodule 
a,sscmblage could indicate anoxic conditions 
close to the site of preservation. 

PROVENANCE AND FORMATION OFTHC 
PHOSPHATISED NODULES 

The Dunsinanc Siie nodules (Fig. 2A,B) may 
have ibrmcd in a variety oi' ways. Their sulv 
mundcd shape is possibly caused by transport luil 
ihis IS no( supported by ihe huk of any other 
evidence of high-energy flow. More lightweight 
fresh clods of organic material may have been 
sub-rounded by transport and subsequently de- 
posited and phosphalised- However, it is debal- 
able whether the nodule material had been subject 
to u sufilcient compression and con-solidation, 
given the fre5hness and lack of distortion ot some 
fossil structures. The ncKlules may have formed 
at Dunsinanc Siie. as parts of a mat of fresh 
organic material resting on the sediment surface. 
The radiation of mineralisation frotn points 
wiihin the mat may have produced the somewhat 
a>unded, but otherw ise variably sized and shaped 
nodules. Roundness ol nodules may also be ai- 
iribuied to weathering whilst embedded in tiie- 
scdiment and after exposure, resulting in contin- 
uous exloliation of outer layers. 

MOI^ELS FOR ACCUMULATION AND 
PROVENANCE AND FORMATION OF NOD- 
ULES (Fig. 3). Ailochlhony <_>f ihe nodules n)\is\ 
he considered because of their peculiar mineral- 
ogy. Inirufonnalional nodules may have formed 
in an overlying sediment which was weathered, 
resulting in the descent of sub-rounded nodules 
to the surface of the Dunsinanc sedifueni and 
bunal by younger Tertiary lacu.strine scdimcni 
(hence the unconhjrming contact). Ilowevcr.llu^ 
is highly unlikely considering that nodules and 
other flourapalile concretions ocear within in $ita 
Dunsinanc sediment. 

The shared mineralogy of the nodules and ver- 
tebrate fossils is compelling wIkh voobidcjing 



m 



MEMOIRS OF THE QUEENSLAND MUSEUM 



conicmporanciiy. The phosphaiisation process 
can lend lo target only those li-ssues which have a 
high organic phosphorous content (Balson. 19^0; 
A!!i.son. 1988 a.ci. suggesiing ihai Dunsinanc 
Site bones, which show varying degrees of phos- 
phate enrichment, may have heen suhjeci to this 
pmcess when they were Iresh and still retained 
soTue organic coiUcnl. This is suggested hy ihc 
greater cnrichmeni of the inner pans of bone 
which would have contained high conceniraiions 
ol" organic maienaL The extreme damage lo the 
Dunsinanc Site hone, bui Lack of evidence of 
rcw<.»rking or trampling (particularly m the case 
ol a partially phosphatiscd, extremely damaged 
Wnttafht'rmm skull with denlanes articulated.) 
tnay indicate damage by Ihe highly acidic condi- 
tions in which the plant material was preserved. 
This would have softened and dissolved bone in 
the nearby subsUatc. Nevertheless, the evidence 
IS ambiguous aiid^i lliis stage. 

CONCLUSIONS 

Dunsinane Site coniatns a probable System A 
vcnebfalc fauna which luay be icntativcly dated 
at 24.7-25 Ma. The fossils from Dunsinane Site 
are preserved with iron nxide-rich llourapiiiiie. 
which appears to have precipitated as the result 
of early diageneiic microbially mediated phos- 
phalisation. The Dunsinane sediment probably 
formed in a low-energy environment, and has 
been severely weathered, resulting in a lag o( 
insoluble fossil material on the surface. Rclaiion- 
shijjs of the llora and invertebrate faiuia to the 
other components of the site arc as yet unre- 
solved. 

ACKNOWLFDGITMENTS 

Atnongst ihc many people lo whom I owe grat- 
itude for discussion, advice, technical informa- 
tion and access lo resources lac: Michael Archer, 
Henk Godihclp, Su/anne Hand, Robert Hill, 
Gregory Jordan, .lane Heath, Bob Mcsibov. Al- 
berto Albani, Bernard C(x)kc, Philip Crca.scr, 
Robert Jones, Winston Ponder, Anna Gillespie. 
Siephao Williams. Mel Dickson. Helcne Manin. 
Peter Athcrdcn, Rad Flossman, Michael de Mol 
und fellow students. 

LITERATURE CITED 

ALLISON, P.A. 1988a. Tapfionomy of the Eocene 
UiiidunC:iuybii>ta.P;ilae(>nlt>l(>gy31: 11)74-1 100. 
1988b. Phosphati/cd soft-bodied squids from the 
Jurahiic 0.\iord Clay. Lelhaia 2 1 : 403-4 1 0. 



l^XSe. KtmservaiLiigeTslaucn: cause and clai^sKt- 
cation. Paleobiologv 14: 131-144 

ARCHER, M.. HAND, S.J. Ac GODTHELP. H. 1094. 
Riversleigh. 2nd ed. lRccd:Sydncy), 
1995. Tenirirv environmcnlul und bioiiv change iii 
Australia. Pp, 77-90. In Vrha, E.S.. Denton.Ci.H.. 
Partridge. T.C. & Buicklc. L.H. (cdsi. Palcoeli- 
maie and evolution. \^Jlh cmphahi> on human 
ongins. (Vale University Press: New Haven). 

B.\LSON. PS. 1980. The uogin and evolution ot Tcr- 
iiar> phosphontes from eo-stem Englaml Inurnal 
of the Geological Society of Uindon 137. 723- 
729. 

BLACK, K. 1997, Diversity and hiostraligraphy t»f ttte 
OiproiodimtLiidtM y}\' Riversleigh. iioilliwchtcm 
Queensland. Memoirs of the Queensland Museam 
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BRIGGS. DEG . KEAR. A.J., MARTILL, D.M. & 
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CREASER. P 1997. Oligocene-Miocenc sctlinicnis of 
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DUNCAN, !J. & BRIGGS. DE.G 1996. Ihrcc di- 
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HIRSCHLER. A.? LUCAS, J & mJBERT. J 199(1. 
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JORDAN. G.J, & HILL. R^S 1996 The fossil record of 
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DUNSINANESITE 



179 



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MARTILL, D.M. !988. Preservation of fish in the 
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MARTJN. H.A. 1994. Australian Tertian, phyiogeog- 
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WILBY. PR. & MARTILL, D.M. 1992. Fossil fish 
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WOODBURNE. MO.. M.\CFADDEN, B.J., CASE, 
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POWER. J.D., WOODBURNE. J.M., 
SPRINGER. K.B. 1994. Land mammal 
biostratigraphy and magnciostratigraphy of the 
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483-515. 



anew species of palorchestidae (marsupial! a) from the late 

middle to early late miocene encore local fauna. 

riversleigh. northwestern queensland 

Karen black 

Black. K.. I*jy7:0fi:3l). A new species of Palnrchcstidae (Marsupialia)frum the late mitldie 
toearlv line Miocene Encore Local Fauna. Riverslci^h. nonhwcsiern Qucensiand. Memoirs 
of thv'Queensicmd Museum 41(2): 181-185, ISSN 007^-8S35. 

A single palorchesiid M' from the Encore Local Fauna. Riversleigh, norihwesiern Ou^'^»is- 
land is described as Palorcheues nnnlKs sp, nov. In size and morphology, it is intermediate 
between the M' ot middle Miocene Pn^patonhcstes twvaculucfphalus froni System C 
deposits, Rivcrsleigti and the Bullock Creek Local Fauna, Northern Territory, and that of* 
Pcflorchc'itf.s painei trom the late Miocene Alcooia Lc»ca] Fauna. Northern Territory. These 
rrlalionships support an early laie Miocene age for the Encore Local Fauna and confirm that 
PropiUorchesies is the sister-group of Pah?rchesrts. Consequently, the monophyly of 
Palorchesiidae is cast further in doubt. Species o\ Ngupakuldiu and Pitikantia may be more 
appropriately regarded a.s plesiomoqihic members of Dipiolodontidae. 
UPalorcheitiihie. Palorchenes, Ptvpalorchi'sivs, lute Miocene, Hiversleigfi 

Karen Black. School of Bloh^ical Science, UniverslnofNe\\ South Wules, NewSourh Wales 
2052, Australia: 4 November 1996. 



A recently discovered upper molar from Encore 
Site on the Gag Plaieau, Rivcrsleigh has in- 
creased the lale Oligoccne to late middle Miocene 
material of Palorchestinae to 9 specimens. This 
paucity of maierial. which prior to 1 ^86 consisted 
exclusively of the highly derived Pulorcliesies, 
has made resolution of relationships within Ihe 
family difficult. Although Pahnhestes annlus 
sp. nov, is only known from an isolated M'. it 
adds substantially lo phylogenetic understanding 
within the family. 

On the basis oi verlebrale stage-of-cvolulion 
hiijcorrelaiion the Encore Local Fauna is cur- 
rently regarded as late middle to early late 
Miocene (approximately lOMa; Archer et al.. 
K)95). Taxa from Encore Site are more derived 
than these characteristic of Riversleigh's upper 
System C assemblages yet plesiomorphic relative 
to related ia\a nl the late Miocene Alcooia Local 
Fauna, Northern Territory (Archer el al.. 1995). 
The species described below supports an early 
late Miocene age*. 

Institutional abbreviations used here are as fol- 
lows: QMF, Queensland Museum palacontologi- 
cal collection; CPC, Commonwealth 
Palaeontologicai Collection al the Australian 
Geological Survey Organisation. Canberra; 
NTMP. An Gallery and Museum of the Northern 
Territory palaeontologicai collection: SAMP. 
South Australian Museum; UCMP, University of 
California. Berkeley. Cusp nomenclalure follows 
Archer ( 1984) and Rich et al. ( 1978) except that 
ihcirhypoconc of upper molars is Ihe melaconule 



following Tedford & Woodburnc { 19S7). Molar 
number homology follows Lueketl (1993). 
Hieher level syslemalic nomenclature follows 
Aplin& Archer (1987). 

SYSTEMATIC PALAEONTOLOGY 

Order DIPROTODONTIA Owen. 1866 
Suborder VOMBATIFORMES Woodburne. 

1984 

Infraordcr VOMBATOMORPHIA Aplin ifc 

Archer. 1987 

Supcrfamily DIPROTODONTOIDEA 

Archer <!<: Barthoiomai. 1978 

Family PALORCHESTIDAE Tate, I94« 

emend. Archer & Bariholomai, 1978 

Palorcheste^ Owen, 1873 

TYPE SPECIES. Pulorrhesiesazaet Owen. 187.3. 
OTHER SPECIES P. panm De Vis, 1 895; P. poinei 
Woodburne. 1967: P. selesiiae Maekness. 1995. 

Palorchestesanulussp- noy. 
<Figs 1-2. Table I) 

VIATERI AL. Holotype. QMF30792, a right .M* imSK- 
ing the posterior cingulum and anterior and posterior 
roots from the late middle Miocene to early lute 
Miocene Encore Local Fauna, on the Gag Plateau. 
Rivcrslcigh. 

ETYMOLOGY. Latin anultis. link; refers to its being 
a structural link between Propalorcheste.% and Pal- 



1«2 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE I. Measurements (mm )otpalofchcstid M' 



Species 


No 


Lei-igih 


Anicrior 
width 


Posierior 
witiih 


Pnhrchesies 
anulus 


QMF3n792 


17! 


13.5 


13.0 


Pr. gottticulus 


NTMP895-1 


14J 


11.5 


10.9 


OMF30883 


15.3 


- 


- 


QMF:U38H4 


162 


12.6 


11.0 


QMF:ohi: 


15.2 


- 


10.8 


cularephuias 


NTMPK62-27 


168 


1.^.2 12.0 


P.\ek'sitae 


QMF 12455 


22.6 


16.6 16.9 




UCMP 70553 R 


16.5 


13.6 


13 8 


UCMP70.S53L 


16.8 


14.4 


137 


P. fmintt 


UCMP70.S50 


16.7 


13.9 


13 7 1 




UCMP 6652 ( 


17.8 : MO 13.2 1 


CPC6752 18-2 * M3 - \ 


P pnr\>u,f 


g.\1F784 


20-7 


15.7 15 4 


QMF 1 1476 


- 


15.4 15 3 


0MF2%.A 


19.3 


14,9 


M4 


Q\\F}1\9 


19.3 


\S0 


14.2 


QMF2%7 ; 19.4 


1.56 


15,6 


QMF2%5 


20.9 




14,5 


P- ozael 


QMR72 


26.6 


21 V ' 21.5 


QMF3837 


25, S 


20.7 19.7 


P3K^70 


28 3 


21.a 21,4 


P3l37i 


28-3 


22.6 21,9 


P>1372 


26.1 


22,9 21.9 



orchcsm and to ihe disiincl midlink, a character of 
Pat arches tes. 

COMPARISON. Palorchesivs wwhn iliffers 
from P. painei in being proporiionalely narrower 
anieriorjy and posterii^rly. in its poorly developed 
lingual cingulum. more open transverse median 
valley lingually. less lightly V-shaped IransverhC 
median valley in lingual view and less well-dii- 
vclopedhmdlink. 

Palorchesres anulus differs from P, parvus. P. 
selestiiie and P. az.ael in bemg smaller; in havmg 
generally less well-developed links; in having a 
.shallower, more open transverse median valley: 
in having a more buccally po.silioned midlink; in 
having a less well-developed, lower, less bucc- 
ally extensive fi.c. in lacking the anterohuccal 
cingulum) anterior cingulum (compared with Ihe 
high, loph-iike anleri4)r cingulum in bt»lh P. 
parvus and P. aztu'l} and consequently, in lacking 
the deep vidleys formed between Ihe anterior 
cingulum and (he anicnor base of ihc protoioph. 

Pidorchestes anulus differs from both P 



parvus and P. azoel: in lacking the second 

medial foreiink; in having a less well - devel- 
oped midlink which is deeply V-shaped, its 
respective anterior and posterior crcsis 
meeting lowcnn the transverse median val- 
ley (and more buccally) than the well-devel- 
oped structure in both P. panus and P. 
azael', in lacking Ihc second buccal midlink. 
and having only a poorly-developed acces- 
sory crest extending anteriorly from a me- 
dial point on the melaloph; in havmg u 
poorly-developed lingual cingulum; in lack- 
ing a buccal cingulum: and in having well- 
developed posiparaconal and 
postmetaconal crests extending posteriorly 
from the apices o[ the paracone and 
metacone respectively. 

Paiorchestes anulus differs from P. 
selesiiae: in having a short crescenlic lin- 
gual cingulum; in lacking the anlerolingual 
foreiink; in lacking the secondary midlink 
and Ihe minor lingual midlink; and m having 
less-crenulatcd enamel at the ha.se of the 
protoioph and meialoph. 

Palorchesies anulus differs from P. 
parvus in having a straighlcr, less crescentic 
melaloph and in having a less crenulale 
transverse median valley. 

Paiorchestes anulus differs IVom P. azael 
in lacking the well- developed lingual mid- 
link; in having a belter-developed posterior 
cingulum; and in having a hindlink devel- 
oped. 



DESCRIPTION- Tooth rectangular, bilophod- 
oni. consisting o\' an unlerior protoioph connecl- 
mg ibe proU)cone wiih the paracone, and a 
posterior metaloph connecting the metacone with 
the metaconuie. Protiiloph antcnorly convex; 
metaloph slightly more linear, wiih its lingual end 
dclleclcd posteriorly. Metaconuie highest cusp; 
the paracone and prolocone subequal in height; 
metacone lowest cusp (taking into account slight 
wear on the apices o\ the major cusps). Anterior 
cingulum well-defined but low on the anterior 
base ol the crown, extending lingually from ihe 
anterohuccal tooth margin to the anlerolingual 
base of the prolocone. Lingual cingulum short, 
poorly dcllned. connecting the poslerolingual 
biisc of ihe prolocone to the antcroiingual base of 
the metaconuie. Posterior cingulum not pre- 
served (but suggested by the short crest at thtr 
poslerolingual base of the metaconuie). 
Foreiink well-developed. eMentiing anteriorly 



MIOCENE PaLORCHESTID, ri versleigh 



\K 




FIG. I Palon'hesfeiamtlussp. nov. Holoiype, QMF307*^2: Occlasal siereopairofrighi M*. Bar indicates IQmm. 



iVom Ibc apex of the protoloph at a point slightly 
lingual fo ihc parucone apcv, meeting ihc anterior 
eingulum at the parastylar corner oi' tooth. Two 
accessory crests (or incipient hnks) poorly-de- 
fined: one originating at the paracone and lading 
down the anterohLiccai lace ol the crown; the 
second originating Ironi the proioloph ai a point 
slightly lingual to the main forelink, extending 
anteriorly and slightly buccally, along the Umgi- 
Uidinal axis oi' the tooth, terminating in the valley 
between the anterior base ofihe proit)Ioph and the 
anterior cingulum. Single midlink lormed by the 
junction of respective anterior and posterior 
crests tiom the melaloph and protoloph meeting 
low in the transverse median valley (making tlie 
link sharply V-shaped in lateral view) approxi- 
mately 4mm from the buccal tooih margin. An 
additional moderately -developed crest extending 
anteriorly from ihc apex of the metaloph into the 
transverse median valley but without a connect- 
ing crest from the protoloph. Hindlink well-de- 
veloped, extending posteriorly and slightly 
lingually i'rom the metaloph, approximately 5mm 
lingual to the buccal looth margin. A thickening 
in the enamel (the posterior meiaconule buttress) 
posterior to the mctaconuleapex hut probably not 
developed into a crest. A similar buttress on the 
posterior Hank of the protocone. 

DlSCLiSSJON. Palorchesttds are rare, fragmen- 
tary components of Tertiary fossil assemblages. 
Until recently, the family consisted of only the 
primitive, generalised, late Oligocene N^^apakal- 
dia and Pinkamki, and the derived, highly 
specialised late Miocene to late Pleistocene Pal- 
onhestes. The large temporal and niorphotogicul 
gaps separating these groups has made relaiKin- 
ships within the family difficult 10 resolve. Stirton 
(19671 recognised 4 suhfainilies within the 
Diprotodontidae and included N^ainikaldia and 



Fitiktvuia in the Paloivhcslinac (later raised la 

family status) based i>n similarities in basicrania! 
morphology to Falorchesre.\\ However, these 
supposed apomorphies are al.so shared with the 
Diprotodoniinae and have since (e.g. Archer 
I9iS4i been intepreted as symplesiomorphic 
within Vi)mbaiomorphia. Consequently. Archer 
( 1 984 \ concluded the Palorcheslidae was not mo- 
nophyleiic, a view later confirmed by Murray 
iI9S6;1990), with his description oi Pro- 
palonhesies dentitions and cranial fragments 
from the middle Miocene Bullock Creek Local 
Fauna. Northern Territory, and .several Oligo- 
Miocene sites at Ri\ ersleigh. Murray ( 1 990) con- 
cluded {.haiPropalonhesu's is the picsiomorphic 
sisier-iaxon oi^ Polorchesres and demonstraied a 
structural transition froin the sclenodoni 
wynyardiid molar pallcrn to the bilophodonl pal- 
orchcslid molar pallern. He further concluded 
thai Nf^apiikatditi and Pdikanna, having sup- 
pressed their selenodonl heritage, .show closer 
affinities to the fully bilophodonl diproiodonlids 
than palorchesiids. Preliminary analyses of late 
Oligocene and Miocene diproiodonlids and pal- 
orchesiids from Rivcrsleigh further suggest that 
N^itpakaldiu and Pitikontia shoulil be regarded 
as primitive members of Diproiodonlidac. 

Pulorthe\tcs antdits supports a Propalor- 
chcstt'\IPalor< hesies sislcr-group relationship 
and confirms doubts (Archer & Bartholomai. 
197K; Archer. 1984: Mun-ay. I'M): Mackness. 
1995)uboui the monophyly of the lamily.TheM* 
o\' P. aniilus is intermediate in a number of key 
features between (he middle Miocene Pro- 
palonhfMesnavacnlacvpliidirs from the BulNvk 
Creek Local Fauna, and System C deposits at 
Rivcrsleigh. and Pnlnrchesfcs painei from {\\c 
late Miocene Alcooia Local Fauna. The Encore 
M' consistently groups with Pmpah>nlwsjes 
noviwulaifphains and P. patm'i falling w Khin Iho 



84 



MEMOIRS OKTIin QUEENSLAND MUSEUM 





A 












/v. porUwulm 






24 r 


Pr. ftomatia/-ephalHS 










P. pamet 


^ 




22 




L P. parvus 












m P. iDtulus 


£ 
^ 


20 




o p. azael 
• P. selesliuc 


o 


s 


18 


- 






o 






> 




£ 


IS 




A fl 




c 






A 




< 


14 

12 

1 


a 




., 1 ... I ... 1 




1 \j 
14 


16 18 20 22 24 


26 2B 30 








Length 





B 



20 



■o 


ia 
















* 


r 




























♦ 


o 


16 - 














to 










.". 






% 
















£ 


14 
12 

in 


_ 

a 


o 


It 

a 

1 1 > 


X 
! . 


6 

1 i. 1 


,A 


J^i. i— ; 



14 16 18 20 22 24 26 28 30 
Lenglh 



C 
22 

20 

16 
16 
14 
12 
10 



GO 






10 



12 



14 16 16 20 
Antenor width 



22 



24 



FiG. 2. Bivariaie plois of M* tooth dimensions for 
species of Pahrchesies antl Propalonhe'ites: A. 
length against anterior widih; B. length againsi pi>s- 
icriurwidlh; C, an teriorwidlh against posterior width. 
Scale in mm. 



size range of both species (Fig. 2A-B). Propor- 
tionally (Fig. 2C), however. P. amdus groups 
more closely with P. painei Along the Pro- 
palorchestes-Palonhestes morphociine (Fig. 
2C) there is a noticeable shift towards a squaring- 
up of the molar crown. Propalonhestes molars 
arc more elongate than wide, and trapezoidal in 
occlusal view, a feature most obvious in the mo- 
lars of the plesiomorphic Pr. poniiculus. In con- 
trast, the posterior width of the M' of the highly 
derived P, azael. is similar to its anterior width, 
giving the tooth a more rectangular profile in 
occlusal view. The intitial stages of this transition 
are evident within P. anulus. The melaloph of M' 
is less convex than in Propalorchestes and ap- 
proaches the length of the protoloph. thus increiis- 
ing the posterior width of the mohir cK)wn. This 
feuiure is retlecled in the position of P. anulus on 
the morphociine (Fig. 2C) and is indicative of its 
derived state relative to Propalarchcstes. 

Other features of the M' that indicate P. anulus 
is derived with respect to Propalanhestes in- 
clude its well-developed forelink and accessory 
forelink and well-developed hindlink; a higher, 
stronger midlink; a more open transverse median 
valley; well- developed convex posterobuccal 
postparaconal and poslmelaconal crests; well-de- 
veloped buttresses on the posterolingual face of 
the protocone and mctaconule; a less convex 
melaloph; and a well-developed parastyle con- 
nected to the protoloph by the forelink. Mackness 
(1^95) listed the well-developed midlink on M' 
as the single synapomorphy o( Palorchesfes as 
opposed to Propalorchestes. The Encore species, 
with a strong, high tnidlink, is included in Pal- 
orchesies as a primitive member of the genus, 
rather than as a derived species of Pro- 
palorchestes. 

The Encore deposit is regarded to be most 
probably early late Miocene in age {Archer el al., 
1995). Slage-of-evolution biocorrclation of mar- 
supial laxa including vombaiids (Krikmann 
(pers. comm.), propleopine kanganxis (Wroc, 
1996), koalas (Archer el al., 1995), dasyurids 
( Wroe. this volume) and ihylacoleonids (Gilles- 
pie, this volume) suggest the Encore Local Fauna 
lies somewhere between Riversleigh's upper 
System C assemblages and the laic Miocene Al- 
cooia Local Fauna and is probably around 1 Ma. 
The presence of P. anulus al Enctire Site, struc- 
turally intermediate between the middle Miocene 
Pi. novaculacephalus and the late Miocene P. 
painei, further substantiates an early late Miocene 
aj^e. 



MIOCENE PALORCHESTID. RIVERSLEIGH 



185 



ACKNOWLEDGEMENTS 

I thank Mike Archer and Peter MuiTay who 
critically read a draft of this paper. I thank Anna 
Gillespie, Henk Godthclp and Steve Wroc for 
assistance with slage-of-evolution biocorrclation 
of the Encore Local Fauna. Vital support for 
research at Riversleigh has come from the Aus- 
tralian Research Grant Scheme (grants lo M. 
Archer); the National Estate Grants Scheme 
(Queensland) (grants to M. Archer and A. 
Bartholomai); the University of New South 
Wales; the Commonwealth Department of Envi- 
ronment, Sporis and Territories; the Queensland 
National Parks and W^ildlife Service; the Com- 
monwealth World Heritage Unit; ICl Australia 
Pty Ltd; the Australian Geographic Society; the 
Queensland Museum; the Australian Museum; 
the Royal Zoological Society of New South 
Wales; the Linnean Society of New South Wales; 
Century Zinc Pty Ltd; the Riversleigh Society 
Inc.; and private supporters including Elaine 
Clark, Margaret Beavis, Martin Dickson, Sue & 
Jim Lavarack and Sue & Don Scotl-Orr. Vital 
assistance in the field has come from many hun- 
dreds of volunteers as well as staff and postgrad- 
uate students of the University of New South 
Wales. Skilled preparation of most of the 
Riversleigh material has been carried out by 
Anna Gillespie. The author also thanks Drs Peter 
Murray and Michael Archer for critically reading 
a draft of this manuscript. 

LITERATURE CITED 

APLIN, K. & ARCHER. M. 1987. Recent advances in 
marsupial systemalics with a new syncretic clas- 
sification. Pp. xv-lxxii. In M Archer (ed.) Pos- 
sums and opossums: studies in evolution. (Surrey 
Beally and Sons and the Royal Zoological Society 
of New South Wales: Sydney) 

ARCHER, M. 1984. The Australian marsupial radia- 
tion. Pp. 633- 808. In Archer, M. & Clayton, G. 
(eds). Vertebrate zoogeography and evolution in 
Australasia. (Hesperian Press: Perth). 

ARCHER, M & BARTHOLOMAI, A. 1978. Tertiary 
mammals of Australia: a synoptic review. Al- 
cheringa2; 1-19. 

ARCHER, M., HAND, S.J. & GODTHELP, H. 1995. 
Tertiary environmental and bioiic change in Aus- 
tralia. Pp. 77-90. In Vrba, E.S., Denton, G.H., 
Partridge, T.C. & Burckle, L.H. (eds), Palaeocli- 
mate and evolution with emphasis on human t)ri- 
gins. (Yale University Press: New Haven). 



GILLESPIE. A. 1997. Priscileo wskeUyae sp. nov. 
{Thylacoleonidae, Marsupialia) iVom the 
Oligocene-Miocene of Riversleigh, northwestern 
Queensland. Memoirs of the Queensland Museum 
41:321-327. 

LUCKETT, W. P. 1993. An ontogenetic assessment of 
dental homologies in therian mammals. Pp. 182- 
204. In Szalay, F.S., Novacek, M.J. & McKenna, 
M.C. (eds), Mammal phytogeny: Mesozoicdiffer- 
enliaiion. multituberculaies, monotremes. early 
therians and marsupials. (Springer-Vcrlag: New 
York). 

MACKNESS. B. 1995. Palorchesies selesuae, a new 
species of palorchestid marsupial from the early 
Pliocene Bluff Downs Local Fauna, northeastern 
Queensland. Memoirs of the Queensland Museum 
38(2): 603-609. 

MURRAY, P. 1986. Pwpalorchesies novaculo- 
cephalus gen. et sp. nov., a new palorchestid 
(Diprotodontoidea: Marsupialia) from the middle 
Miocene Camtield Beds. Northern Territory, Aus- 
tralia. The Beagle. Occasional Papers of the 
Northern Territory Museum of Arts and Sciences 
3(1): 195-211. 

MURRAY, P. 1990. Primitive marsupial tapirs {Pro- 
palorchestes novaculacephalus Murray and P. 
ponticuhts (Marsupialia: Palorchestidae sp. nov.) 
from the mid-Miocene o'i north Australia. The 
Beagle. Records of the Northern Territory Mu- 
seum of Arts and Sciences 7: 39-5 1 . 

RICH, T.H., ARCHER, M. & TEDFORD, R.H. 1978. 
Raemeoifierium yaikolai gen. ct sp. no\'.. a primi- 
tive diprotodontid from the medial Miocene of 
South Australia. Memoirs o^ the National Mu- 
seum of Victoria 39: 85-91. 

STIRTON, R.A. 1967. The Diprotodonlidae from the 
Ngapakaldi Fauna, South Australia. Bureau of 
Mineral Resources, Geology and Geophysics Bul- 
letin 85: 1-44. 

TEDFORD, R.H. & WOODBURNE, M.O. 1987. The 
Ilariidac, a new family of vombatifomi marsupials 
from Miocene strata of South Australia and an 
evaluation of the homolgy of molar cusps in the 
Diprotodonlidae. Pp. 401-418. In Archer, M. 
(ed.). Possums and opossums: studies in evolu- 
tion. (Surrey Beatty & Sons and the Royal Zoo- 
logical Society of New South Wales: Sydney). 

WROE. S. 1996. An investigation of phylogeny in the 
giant extinct ral kangaroo Ekahadeui (Pro- 
pleopinae, Potoroidae, Marsupialia). Journal o'[ 
Paleontology 70(4): 681-690. 

WROE. S. 1997 Mayigrip/ms orbits gen. el sp. nov. a 
Miocene dasyuromorphian from Riversleigh, 
northwestern Queensland. Memoirs of the 
Queensland Museum 41: 439-438. 



UiVliRSlTY AND BIOSTRATIGR.'VPH^' OF THE DIPROTODONTOIDEA OF 
RTVERSLBIGH. NORTHWESTERN QUEENSLAND 

KAREN BLACK 



Black, K. !907;06:30: Divcrsily andbiosiraligraphy u! theDiprotodonloideaolRiversleigh. 
norihwtvsicmO'tccnslanci. Me(f\oin of the Qaeenslcmd Museum 4H2): 187-192. Brisbane. 
ISSN 0079-8835. 

The diversity und distrihution of Riversieigh's Dipruiodonloidca is discussed with rcspccl 
lo ciirrcnl uildcfsiaitdmg uf the local straiigr;iphic sequence. The mnrc plesiomorphic 
diproiodoniid and pa!orchest>d laxa urc resiricicd to the older Sysieivi A Local Fauiuis. 
M'o/it'/cnand PropalorchesfvsT^wgt ihruu^houi SysicniN A. B midC exhibit iug ;i tnorphocl- 
ine. Stage-of-evoluliun biocoiTclation with oihcr Ausiralian and New Giunean Ternary 
mummal I'aunas is in accord with previously e&iabllshed biosiiatigruphie hyputhcscs. How- 
ever .some basal Riverslcigh deposits may predaic (he central Ao'^tralian Wipajiri and 
Eladunna Fnrmalions. Diproi odontoid diversity is; high ni Sysieni A deposus. low iii System 
B and rclalivcly high in System C. Abundance is high in SysiL-m C^ deposits and low in 
Systems A and B Only two la.va extend into the Pleistocene. Patterns of diversity and 
abundance are discussed in view ol^ Australia's changing Cainozoic climate. 
U\Diprotodoniidue , Palorche^tidae, Rtverslct^h, Tertiary. 

Karen BlacK School of Biological Science. University c^NewSoHih Wtt/es, NewSouih HWA'.v 
2052, Australia: received 4 November 1996. 



The paucity orradionictricany dated Australian 
Tcniary freshwater fossil deposits has resulted in 
marsupial stagc-of-evolulion biochronology 
being ihc most commonly employed method of 
dating fossil assemblages (Wuodburne el al., 
I'J85). FundametUal to this tnethod is an adequate 
understanding of phyiclic succession within the 
ta\on (Megirian, 1994). Chronologic and phy- 
letic succession within diproiodontoid lineages 
arc well documented (e.g. Stirion et al., 1967; 
Murray. 199Ub; Murray el al. 1993). Conse- 
quently, the group has played a major role in 
previous biochronologic analyses (e.g. Stirton ct 
al.. 1967: Woodbtirncei al.. 1985; Woodbumeel 
al., 1993; Archer ctal., 1994; Archer etal., 1995) 
and has contributed significantly to the construc- 
tion of the chronological framework of 
Australia's Tertiary mammal faunas. 

Similarly^ because of limited siratigraphic data 
and the lack of radiometric dates, much of the 
current chronologic sequence o\ Riversieigh's 
local faunas has been constructed on the basis of 
vertebrate biocc^rrclalion (Archer el al.. 1989), 
Recent reappraisal of diprotodontoid material 
from Rivcrsleigh indicates at least 9 genera and 
I K species. This paper oullines the diversity and 
distribution of diprotodontoids throughout the 
Riversleigh Systems and discusses their bearing 
on current biostratigraphic understanding, 
tiioc(irrelatu:m of Riversieigh's Oligocene- 
Miocene deposits with other Australian fossil 
assemblages is also discussed, 



Liihostraiigraphy and Systems terminology 
follow Archer el al. ( 1989, I994al. Cusp nomen- 
clature lolhms Archer (1984) and Rich el al. 
(1978) Molar hontology ts (hat proposed hy 
LuckctH 1993 k Picinolariiumbcr follows Flower 
(1867) Higher level systematic nomcnelaiurc 
follows Aplin & Archer ( 1987). 

BIOSTR/XTIGRAPHY 

Figure 1 lists the diprotodt)nlids and pal- 
orcheslids from different local faunas withm 
Riversieigh's siratigraphic units. These units are 
defined hy Archer et al (1989. 1994a): System 
A, late Oligocene lo early Miocene: System B. 
eariy to middle Miocene; System C. middle to 
early laie Miocene; and Pleistocene assemblages. 

Silvahistms wichaelhini is the most picsio- 
morphic zygomaiurine recognised {with the pos- 
sible exceplion of Haemeotheriinn yatkolai Rich 
ci al-, 1978) and may be antecedent to (he entire 
/ygomaturine radiation (Black & Archer, 1997a). 
Four species of Neohelos are recognised ( P. Mur- 
ray pcrs. comm). Neohelos sp. nov. I, a stnall 
picsiomorphic form; N. firarensis a medium- 
sized moderately derived form from the 
Kuljamarpu Local Fauna, South Australia; 
Neohelos sp. nov. 2. a large deri ved formfrom the 
Bullock Creek Local Fauna. NT. anieccdcnt lo 
Kolopsis torus (Woodburne, 1967) o\' the laic 
Miocene Alcotita Local Fauna. NT; and Neohelos 
sp- nt)v. 3, a highly derived form structurally 



188 



MEMOIRS OF THE QUEENSLAND MUSEUM 









SYSTEV 




A 






SYSTEM B 


SYSTEM C 


PL 

El 




L 


MID 


H 


LOW 


MID 


HIGH 


C+ 


LOCAL FAUNAS ♦ 


D 


B 



H 


B 
R 


S 
B 


J 

A 


V 
IP 


A 
L 


W 
H 


W 
W 


C 
S 


N 
G 


D 
T 


M 
M 


i 

N 


R 
T 


G 


D 
I 


J 
JS 


H 
H 


J 
J 


90 


J 
C 


D 
S 


C 




E 

N 


T 

E 


S. michaelbirti 






1 


















































S. johnnilandi 














2 










































Silvabestius sp. 


1 
















































10 






Neohetos sp. nov. 1 


1 


1 




1 


1 














































N. tirarensis 


1 






1 












1 


I 


1 




1 


1 








1 


















Neohelos sp. nov. 2 


































I 






4 
















Neohelos sp. nov. 3 


h— — 








































2 














Ni. lavarackorum 
































1 


1 


1 








24 


1 


2 








Nifiibadon sp. 












































1 












Zygomaturine 
gen. nov. 


























9 






























B. angulaiu/n 


1 


1 


2 




4 


1 






3 






































Bemaiherium sp. 
















1 








































Ngapakaldia sp. 










I 














































Pr poriticulus 


I 






1 


2 


1 








1 


1 


1 










I 






















Pr. novaciiia 
cephalus 






































1 


I 
















Palorchesies anulus 




















































1 




P. Qzael 






















































1 


Diprotodon opiaiuiu 






















































1 


TOTAL 


5 


2 


2 


3 


4 


2 


I 


1 


1 


2 


2 


2 


1 


1 


1 


1 


3 


J_ 


-} 


2 


1 


2 


1 


1 


1 


1 


2 



FIG. 1 . The distribution of diprotodontoid taxa through the Riversieigh sequence. Numbers indicate the minimum 
number of individuals for each species found in a given local fauna. PLEI=Pleistocene. System B heading is 
subdivided into: L=Low, MID=Middle and H=HighLocal Fauna Abbreviations: AL, Alsite; 90, Alan's Ledge 
1990; BO, Burnt Offering; BR, Bone Reef; CO, Cleft of Ages (tentatively regarded as System C); CS, Camel 
Sputum; D, D-Site; DI, Diprotodonl Site; DS, Dome Site; DT, Dirk's Towers; EN, Encore Site; G, Gag Site; 
H, Hiatus Site;HH, Henk's Hollow; IN, Inabeyance; JA, Jeanette's Amphitheatre; JC, Jim's Carousel; JJ, Jaw 
Junction; JJS, Jim's Jaw Site; MM, Mike's Menagerie; NG, Neville's Garden; RT, Ringtail Site; SB, Sticky 
Beak Site; TE, Terrace Site: WH, White Hunter; VIP. VIP Site; WW, Wayne's Wok. 



transitional between Neohelos sp. nov. 2 and 
Kolopsisyperns (Murray et al, 1 993) from the late 
Miocene Ongeva Local Fauna, Waite Formation. 
Bematherium sp. is known from a single maxil- 
lary fragment which is plesiomorphic relative to 
B. angiilum. A new unnamed species of 
Ngapakaldia is more derived than the central 
Australian N. tedfordi but plesiomorphic relative 
to N. bonythoni. Fragmentary material of 
Zygomaturine gen. nov. has an uncertain phylo- 
genetic position. Nimbadon sp. is similar to N. 
lavarackorum in size, molar morphology and cra- 
nial profile; however, differences in upper pre- 
molar morphology may imply a more 
plesiomorphic position within the genus. How- 
ever, I doubt whether this single skull is indica- 
tive of a new species. Extreme intraspecific 
variation in P-^ morphology is common among 
Tertiary zygomaturines (e.g. Neohelos spp.; P. 



Murray pers. comm.) with reduction or loss of 
premolar cusps a relatively common phenome- 
non. Comparable cranial material for Ni. lav- 
arackorum has yet to be processed. Propalor 
chestes ponticulus (Murray, 1990b) is plesio- 
morphic relative to P. novaculacephalus. Pro- 
palorchestes novaculacephalus occurs in the 
Bullock Creek Local Fauna; Murray (1990b) in- 
dicated that this material is more derived than the 
Riversieigh specimens. Palorchestes anulus is 
intermediate between Pr. novaculacephalus and 
Palorchestes painei from the late Miocene Al- 
coota Local Fauna (Black, 1997). Palorchestes 
azael and Diprotodon optatum , the most derived 
members of Palorchestidae and Diprotodontidae. 
respectively, are known from Terrace Site, which 
has been radiocarbon dated at approximately 
23,900-F/- 4,100-2,700 years BP (Davis & 
Archer, 1997). 



DIVERSITY OF DIPROTODONTIDS AND PALORCHESTIDS 



189 



DISCUSSION 

AGREEMENT WITH PROPOSED STRATIG- 
RAPHY. The dislribution of diprolodonle^ids 
throughout the Riversleigh systems is generally 
consistent with Archer et al/s (1989, 1994a, 
1995) proposed stratigraphic framework. The 
most plesiomorphic forms are restricted to the 
older System A local faunas, and more derived 
taxa become more abundant in Systems B and C. 

A similar pattern oi' distribution is evident 
within lineages. There is a gradual evolution in 
cranial and dental morphology accompanied by 
an increase in body size within Neohelos and 
PropalorchesTes. respectively, through the 
Riversleigh sequence (Murray, 1990b; pcrs. 
comm.). Although some temporal overlap is evi- 
dent, generally Neohelos sp. nov. 1 is the domi- 
nant form in System A, N. tirarensis is most 
abundant in System B. Neohelos sp. nov. 2 spans 
low to high System C and Neohelos sp. nov. 3 is 
unique to high System C Jaw Junction Site. 

A similar succession is exhibited by pal- 
orchestids. Plesiomoq")hic Propalorchesies pon- 
ticulus is relatively common in System A and B 
Sites, and is succeeded by the more derived Pr. 
novae uiacephal us in mid-high System C Sites. 

INTRACONTINENTAL COMPARISONS. The 

Riversleigh assemblages contain the most diverse 
array of diproiodontids and palorchestids of any 
single region on the continent. Of the 9 genera, 
Neohelos, NimbadofU Ngapakatdia, Propalor- 
chesies and Palorchesfes are also know n from the 
Oligocene-Miocene in central and northern Aus- 
tralia. The biochronological potential of some of 
these genera is, however, limited, because of 
uncertainty about interspecific afi'inilies. 
Neohelos, Propalorchesies and Palorchesfes. are 
biochronologically most significant. 

Neohelos tirarensis from System B is close to 
the type material from the Kutjamarpu Local 
Fauna suggesting age equivalence; this is sup- 
ported by other shared taxa such as Wakiewxikie 
/^/H'iYva(Godthelpclal.. 1989). Pa//V/ra( Archer, 
1994), V/akaleo oldfieldi (A. Gillespie pers. 
comm.), Namilamadeta: Namharoo (Archer ct 
al., 1989) and Litokoala (Black & Archer. 
1997b). 

Neohelos sp. nov. I in Systems A-C is more 
plesiomorphic than N, tirarensis suggesting thai 
some oi' Rivcrsleigh's stratigraphic units predate 
the KutjamarpuLocal Fauna. Neohelos from fau- 
nal /ones D-E of the Etadunna Formation have 
not been specifically identified (Woodburnc et 



al, 1993). Murray (pers. comm.) suggests thai if 
this material is found to represent tirarensis then 
some of Riversleigh's older deposits may predate 
mammal-bearing Etadunna Formation. 

Myers & Archer (1997) correlated White 
Hunter Site (System A) and MammaUm Hill 
(Zone D. Etadunnal Fomiation). the laller 24.7- 
25 MY BP (Woodburne et al.. 1993). 

INgapakaldia sp. nov. in System A extends the 
generic distribution from South Australia. This 
species appears to be more derived than N. 
tedfordi from the Ngapakaldi and Ngama Local 
Faunas (Etadunna Fmn) and the Tarkarooloo 
Local Fauna (Namba Fmn), yet plesiomorphic 
relative to N. honythoni from the Ngapakaldi 
Local Fauna. Ngapakaldia also occurs in faunal 
zones C. D and E of the Etadunna Fmn. As with 
Neohelos, species level identification is neces- 
sary before Ngapakatdia will be o^ use in 
biocorrelation 

Propalorchesies novaculacephalus and 
Neohelos sp. nov. 2 in low to mid System C 
assemblages confirms previous hvpotheses 
(Archer etal., 1989; 1994a; 1995) that 'they are of 
a similar age to the Bullock Creek Local Fauna 
(Fig. 2). Correlation is further supported by the 
shared Nimbacinns dicksoni (Muirhead &. 
Archer, 1990), Wakaleo vanderleiiri (Munay & 
Megirian, 1990) and Balharoo sp. (Flanncry et 
al., 1983). 

Neohelos sp. nov. 3, the largest and most de- 
rived species oi' Neohelos, occurs in the Jaw 
Junction assemblage suggesting that this high 
System C deposit is younger than the Bullock 
Creek Local Fauna. Neohelos sp. nov. 3 exhibits 
an upper third premolar morphology that antici- 
pates the condition ftiund in the more derived 
zygomalurine Kolopsis, which first appears in the 
Alcoota and Ongeva Local Faunas of the Waile 
Fmn, Northern Territory. Ancestor-desccndenl 
relationships are well supported for the 
Neohelos! KolopsislZygomatunis clade (Stirton 
etal., 1967; Murray etaL, 1993). The presence of 
Neohelos material, structurally transitional be- 
tween Bullock Creek's N. sp. nov. 2 and (he 
Waite Formation's Kolopsis further suggests this 
correlation making some of Riversleigh's high 
System C deposits, such as Jaw Junction Site, 
younger than the Bullock Creek Local Fauna but 
older than late Miocene deposits ol' the Waile 
Formation. 

This is also true of Riversleigh's Encore assem- 
blage which, on the basis of its derived and often 
unique fauna, is believed to be early late Miocene 
in age (Archer et al., 1995). This is based on the 



190 



MEMOIRS OF THE QUEENSLAND MUSEUM 



10.4 

MY 



16.3 
MY 



23.3 

MY 



LATE 
MIOCENE 



MID 

MIOCENE 



FARiA 

vuo< em: 



LAIE 

Ol IGOCENE 



KUTJAMARPU 



NGAMA 



TARKAROOLOO 



NGAPAK.ALDI 



DiTJIMANKA 



ERICMAS 



PINPA 



ALCOOTA 



BULLOCK 
CREEK 



ENCORE 



SYSTEM C^ 



SYSTEM B 



SYSTEM A 



SOUTH AUSTRALIA 



NORTHERN 
TERRITORY 



RIVERSLEIGH 
QUEENSLAND 



Fig. 2. Tentative correlation of Rivcrsleigh's Oligo-Mioccne fossil deposits with the Etadunna and Wipajiri 
Formations, South Australia and the Alcoota and Bullock Creek local faunas, Northern Territory based on 
diprotodonloid slage-of-cvolulion comparisons. 



presence of several taxa which exhibit adapta- 
tions to the onset ol drier climates during the late 
Miocene as well as taxa whose lineages extend 
into Pliocene and Pleistocene times. These in- 
clude: Palorchestes aniilus which is structurally 
antecedent to P. painei from the late Miocene 
Alcoota Local Fauna (Black, 1997); a small 
Phascolarctos\ a hypselodont vombalid related to 
the late Cainozoic Warendja wakefielcli (Archer 
et al., 1995); a thylacoieonid intermediate be- 
tween Wakaleo vanderleuri and the late Miocene 
Wakaleo alcootense (A. Gillespie pers. comm.); 
a giant rat kangaroo, Ekaltadeta jamiemulvanei 
(Wroe, 1996), which is possibly annecetant to 
Pleistocene Pmpleopus; and Mayigriphus 
orbusCWroe. 1997), a dasyuromorphian with 
some features correlated with drier environments 
in modern dasyurids. 

DIVERSITY AND FAUNAL CHANGE. Sys- 
tem A sites contain the highest number of con- 
temporaneous diprotodonloids with 5 species at 
Site D and 4 at Sticky Beak Site (Figs 1 -2 ). A drop 
in diversity is evident in System B with only 1 or 
2 species per site. Diversity increases slightly in 
low to middle System C assemblages with gener- 
ally 1-2 contemporaneous diprotodontoid spe- 



cies. Conversely, upper System C deposits ex- 
hibit a decline in diprotodontoid diversity. If we 
consider the small number of taxa being analysed 
such changes in diprotodontoid diversity may not 
be significant. However, a similar decline in both 
family- and generic-level diversity is evident in a 
number of other marsupial groups during the 
middle to late Miocene. This decline may be 
related to the late Miocene onset of 'icehouse' 
climatic conditions resulting in the regional col- 
lapse of rainforest and subsequent spread of open 
forest and woodland/savanna (Archer et al., 
1994b; 1995). 

Replacement of rainforest habitat by more open 
forest may have benefited select diprotodontoids 
which is reflected in an increase in species abun- 
dance for some System C Local Faunas. Rela- 
tively high 'minimum number of individuals' 
estimates for species o[ Nimbadon and Neohelos 
sp. nov. 1 have been recorded from AL90 and 
Cleft of Ages Sites respectively. This may sug- 
gest that individuals of these species were roam- 
ing in mobs, a feature characteristic of slow 
moving medium- to large-sized herbivores in rel- 
atively open environments. 

It is also feasible that high abundance in the 
above System C assemblages, and conversly low 



DIVERSITY OF DIPROTODONTIDS AND PALORCHESTIDS 



I9J 



abundance in System A-B assemblages, may rep- 
resent sampling or laphonomic biases. Likewise, 
the absence of diprotodonlines in System B and 
C may be an artefact of incorrect taxonomic 
assignment. Even so, several zygomaturine spe- 
cies recovered from System B and System C 
assemblages appear to have developed a number 
ofdiprotodontine- like features of their dentition. 
These include the reduction of the paraslyle and 
loss of the hypocone on P-' and a reduced 
paracristid on Mi. A similar phenomenon has 
occurred in the Alcooia Local Fauna where dental 
and cranial morphology of the zygomaturine Al- 
kwertathehum webbi is convergent on that of the 
diprolodontinc Pynunios alcootense (Murray, 
1990a). In general, zygomaturines display a 
higher diversity and abundance in Tertiary fossil 
assemblages than diprotodonlines. This may re- 
fiect their greater ability to adapt to changing 
environments than their diprolodontinc counter- 
parts. It is further rellected in Rivcrslcigh's Sys- 
tem B and System C Local Faunas, where 
diversifying zygomaturines have subsequently 
radiated into vacant niches occupied by 
diprotodontines during the late Oligocene. 

Diprolodontoids are not known from 
Rivcrslcigh's Pliocene assemblages. They seem 
to have declined markedly in generic diversity 
throughout the Australian Pliocene. Significant 
faunal turnover is characteristic of most marsu- 
pial families during the late Miocene and early 
Pliocene (Archer et al., 1995). 

The drop in diversity of diprotodontid and pal- 
orchestid species continues into the Pleistocene, 
with only one memberof each family represented 
(Fig. 1). Both species are common throughout 
Australia's Quaternary fossil deposits, yet both 
are ihe lasl of their respective lineages. This de- 
cline in diprotodonloid diversity may be a conse- 
quence of unsuccessful competition with rapidly 
diversifying mesic and xeric macropodoids 
( Archer el al.. 1994). 

ACKNOWLEDGEMENTS 

I wish to thank Mike Archer and Peter Murray 
who critically read a draft of this manuscripi and 
Anna Gillespie. Henk Godthelp and Steve Wroe 
for ihcir assistance with taxon correlations. Vilal 
support for research at Riversleigh has come from 
the Australian Research Grant Scheme (to M. 
Archer); the National Estate Grants Scheme 
(Queensland) (toM. Archer and A. Bariholomai); 
the University of New South Wales; the Com- 
monwealth Department of Environment, Sports 



and Territories; the Queensland National Parks 
and Wildlife Service; the Commonwealth World 
Heritage Unit; ICI Australia Ply Lid; ihe Austra- 
lian Geographic Society; the Queensland Mu- 
seum; the Australian Museum; ihe Royal 
Zoological Society of New South Wales; Ihe 
Linnean Society of New South Wales; Century 
Zinc Ply Ltd; the Riversleigh Society Inc.; and 
private supporters including Elaine Clark. Mar- 
garet Beavis, Martin Dickson, Sue & Jim Lav- 
arack and Sue & Don Scotl-Orr. Vital assistance 
in the field has come from many hundreds of 
volunteers as well as staff and postgraduate stu- 
dents of the University of New South Wales. 
Skilled preparation of most of the Riversleigh 
material has been carried out by Anna Gillespie. 

LITERATURE CITED 

APLIN, K. & ARCHER, M. 1987. Recem advances in 
marsupial sysicmaiics with a new syncretic clas- 
sificalion. Pp. xv-lxxii. In Archer. M. (cd.). Pos- 
sums and opossums: studies in evolution. (Surrey 
Beatty &. Sons & Royal Zoological Society of 
New South Wales: Sydney). 

ARCHER. M. 1984. The Australian marsupial radia- 
tion. Pp. 633-808. In Archer, M. & Claylon, G. 
(eds). Vertebrate zoogeography and evolution in 
Australasia. (Hesperian Press: Perth). 

1994. The ringtail possums (Marsupiulia, 
Pseudochciridae) of Riversleigh. Riversleigh 
Symposium Abstracts, p.24-25. 

ARCHER, M.. GODTHELP, H., HAND. S.J. & 
MEGIRIAN. D. 1989. Fossil mammals of 
Riversleigh, northwestern Queensland: prelitni- 
riiiry overview of biosiratigraphy, correlation and 
environmental change. The Australian Zoologist 
25:35-69. 

ARCHER. M.. HAND, S.J. & GODTHELP. H. 1994a. 
Riversleigh. 2rid ed.iReed Books: Sydney). 
1994b. Patterns in the history of Australia's mam- 
mals and inferences about palacohabitals. Pp. 
80-103. In Hill, R.S. (ed.). History of the Austra- 
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1995. Tertiary environmental and biotic change in 
Australia. Pp. 77-90. In Vrba. E.S.. Denton.G.H.. 
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BLACK, K. 1997. A new species of Palorchestidae 
(Marsupialia) from the late middle to early lale 
Miocene Encore Local Fauna, Riversleigh, north- 
western Queensland. Memoirs of the Queensland 
Museum 41: 181-185. 

BLACK, K. & ARCHER. M. 1997a. Silvahesiius gen. 
nov. a primitive zygomaturine (Marsupialia, 
Diprotodonlidae) from Riversleigh, northwestern 



192 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Queensland. Memoirs oflhe Queensland Museum 
41: 193-208. 
1997b. Nimiokoala gen. nov. ( Marsupial ia, 
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Queensland, vvilh a revision oi Litokoala. Mem- 
oirs of the Queensland Museum 41 : 209-228. 

DAVIS, A.C. & ARCHER, M. 1997. Palorchestes 
azciel (Marsupialia, Palorchcstidae) From the late 
Pleistocene Terrace Site Local Fauna, 
Riversleigh, nonhweslem Queensland. Memoirs 
of the Queensland Museum 41: 31 5-320. 

FLANNERY, T. F, ARCHER, M. & PLANE, M.D. 
1983. Middle Miocene kangaroos 
(Macropodoidea: Marsupialia) from three locali- 
ties in northern Australia, with adcscription of two 
new subfamilies. Bureau of Mineral Resources. 
Geology & Geophysics, Bulletin 7: 287-302. 

FLOWER, W.H. 1867. On the development and suc- 
cession of lecth in the Marsupialia. Philosophical 
Transactions oflhe Royal Society of London 157: 
631-641. 

GODTHELP, H., ARCHER. M.. HAND. S.J. & 
PLANE. M.D. 1989. New poloroine from Tertiary 
Kangaroo Well Local Fauna, Northern Territory 
and description of upper dentition of potoroinc 
Wakiewakif lawsoni from Upper Site Local 
Fauna, Riversleigh. 5th Conference on Australian 
Vcncbratc Evolution, Palaeontology and System- 
atics. Abstracts: p. 6. 

HAND, S.J., ARCHER, M.. GODTHELP, H., RICH. 
T.H. & PLEDGE, N.S. 1993. Nimhadon, a new 
genus and three new species of Tertiary 
zygomaturines (Marsupialia: Diproiodontidae) 
from northern Australia, with a reassessment of 
Neoheios. Memoirs of the Queensland Museum 
33: 193-210. 

LUCKEIT, W.P, 1993. An ontogenetic assessment of 
dental homologies in iherian mammals. Pp. 182- 
204. In Szalay, FS., Novacek. M.J. & McKenna. 
M.C. (eds), MamiTial phylogeny: Mesozoic differ- 
entiation, multitubercuiates, monolremes, early 
therians and marsupials. {Springer-Verlag; New 
York). 

MARSHALL, L.G., CASE, J.A. & WOODBURNE, 
M.O. 1989. Phylogenetic relationships of the fam- 
ilies of marsupials. Current Mammalogy 2: 433- 
502. 

MEGIRIAN. D. 1994. Approaches to marsupial 
biochronology in Australia and New Guinea. Al- 
chcrmga IS: 259-74. 

MUIRHEAD, J. & ARCHER. M. 1990. Nimhacinus 
dick.soni, aplesiomorphic thylacine (Miirsupialia: 
Thylacinidae) from Tertiary deposits of Queens- 
land and the Northern Territory, Memoirs <){' the 
Queensland Mu.seum 28: 203-221. 

MURRAY, P. 1990a. Aikwcnatherium wehhi, a new 
/ygomaturine genus and species from the late 
Miocene Alcoota Local Fauna, Northern Territory 
(Marsupialia: Diproiodontidae). The Beagle. Re- 
cords o\' the Northern Terrilorv Museum oi Arts 
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1990h. Primitive marsupial tapirs {Propalorclwsies 
iwvacidacephahis Murray and P. ponticidiis sp. 
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(Marsupialia: Paiorchestidae). The Beagle, Re- 
cords oi the Northern Territory Museum of Arts 
and Sciences 7: 39-5 1 . 

MURRAY, P. & MEGIRIAN, D. 1990. Further obser- 
vations on the morphology oi Wakaleo 
vcmderleuen (Marsupialia: Thylacoleonidae) 
from the mid-Miocene Camfield Beds, Northern 
Territory. The Beagle, Records oi ihe Northern 
Territory Museum of Arts and Sciences 7: 9 i - 1 02. 

MURRAY, P.. MEGIRIAN, D. & WELLS, R. 1993. 
Kolopsis yperits sp. nov. (Zygomaturinae, 
Marsupialia) from the Ongeva Local Fauna: new 
evidence for the age oi the Alcoota fossil beds of 
central Australia. The Beagle, Records of the 
Northern Territory Museum of Arts and Sciences 
10: 155-172. 

MYERS, T.J. & ARCHER. M. 1997. Kuierintjangania 
(Marsupialia, Ihiriidae): A revised and extended 
systematic analysis based on fossil material from 
ihe late Oligocene oi Riversleigh. northwestern 
Queensland, Australia. Memoirs of the Queens- 
land Museum 41 : 379-392. 

RICH, T.H., ARCHER, M. & TEDFORD, R.H. 1978. 
Raemeotheriwn yatkolai gen. et sp. nov.. a primi- 
tive diprotodontid from the medial Miocene of 
South Australia. Memoirs of the National Mu- 
seum of Victoria 39: 85-91. 

STIRTON, R.A., WOODBURNE. M.O. & PLANE. 
M.D., 1967. A phylogeny of the Tertiary 
Diproiodontidae and its significance in correla- 
tion. Bulletin of the Bureau of Mineral Resources. 
Geology and Geophysics. Australia 85: 149-160. 

TEDFORD. R.H. 1967. Fossil mammal remains from 
the Carl Creek Limestone, northwestern Queens- 
land. Bulletin oflhe Bureau of Mineral Resources, 
Geology and Geophysics, Australia 92: 2 1 7- 237. 

WOODBURNE. M.O., TEDFORD, R.H., ARCHER, 
M., TURNBULL, W.D., PLANE, M.D. & 
LUNDELIUS, E.L. 1985. Biochronology oflhe 
continental mammal record of Australia and New 
Guinea. Special Publications. St)ulh Australian 
Department ol' Mines and Ener^v 5: 347-363. 

WOODBURNE, M.O., MACFADDEN. B.J.. CASE, 
J. A., SPRINGER, M.S., PLEDGE, N.S., 
POWER, J.D., WOODBURNE, J.M. & 
SPRINGER. K.B, 1993. Land mamma! 
biostraiigraphy and magnetosiraligraphy oi the 
Eladunna Formation (late Oligocene) oi South 
Australia. Journal ofVertebrate Palaeontology 13: 
483-515. 

WROE, S. 1996. An investigation ol phylogeny in the 
giant Rat- kangaroo Ekahadeia (Propleopinae, 
Poloroidae, Marsupialia). Journal of Paleontology 
70: 677-686. 
1997. Mayigriplnts orhus gen. el sp. nov., a new 
Miocene dasyuromorphian from Riversleigh. 
northwestern Queensland. Memoirs oi ihe 
Queensland Museum 41 : 439-448. 



SILVABESTIUS GEN. NOV., A PRIMITIVE ZYGOMATURINE (MARSUPIALIA, 
DIPROTODONTIDAE) FROM RIVERSLEIGH, NORTHWESTERN QUEENSLAND 

K. BLACK AND M. ARCHER 

Black, K. & Archer, M., 1997:06:30. Silvabestius gen. nov. a primitive zygomaturine 
(Marsupialia, Diproiodoniidae) from Rivcrsleigh, northwesiem Queensland. Memoirs of the 
Queensland Museum 4\{2)\ 193-208. Brisbane. ISSN 0079-8835. 

A new genus and two new species of primitive Oligo-Miocene zygomaturincs are described 

from Riversleigh, northwestem Queensland. A maxillary fragment described by Tedford as 
palorchestine (Marsupialia: Palorchestidac) is referred to Silvabestius, and is intermediate 
between 5. michaelbirti sp. nov. and S.jolmnikmdi sp. nov. Trends in premolar morphology 
within the genus support Slirton ct al.s* proposal that zygomaturincs arose from primitive 
diprotodontine-like forms in which the parastyle on P-^ was less developed. The tricuspid 
P- of 5. michaelbirti is intermediate between the bicuspid P-^ of primitive diprotodontincs and 
the more typical quadricuspid P-^ o{ S.johnnilandi. Cladislic analysis of the Zygomaturinae, 
based largely on the upper third premolar, is compared with previous analyses. 
n Zygomaturinae. Silvabestius, Riversleigh, Oligocene, Miocene. 

Karen Black & Michael Archer, School of Biological Science, University of New South 
Wales. New South Wales 2052, Australia: 4 November 1996. 



Tedford (1967) described Palorchesiinae gen. 
and sp. indet., a right maxillary fragment contain- 
ing P^-M' and the anterior half o{ M- from Site 
D, Riversleigh, northwestern Queensland. This 
assignment was based on plesiomorphic features 
that were at the lime otherwise only known in 
primitive palorchestids such as Ni^apakaldia and 
Pitikcintia. Tedford (1967) did, however, note 
enlargement of the parastylar region, a low 
parastyle and a more extensive posterolingual 
cingular shelf in P-^ that appeared to anticipate 
development of the quadricuspid premolars of 
primitive zygomaturincs. These features were 
also noted by Hand et al. ( 1 993b) who, in light of 
piimilive zygomaturincs from northern Australia 
[Nimbadon Hand ct al., 1993a; Alkwertatheriian 
Murray, 1 990b), Hand et al. ( 1 993b), referred the 
D-site specimen to the Zygomaturinae. 

Two perfectly preserved crania of the primitive 
zygomaturine diprotodontid, Silvabestius 
johniukmdi sp. nov. were collected at VIP Site in 
1989 and a complete skull o{ Silvabestius mic- 
haelhif'tisp. nov. was recovered from Hiatus Site 
in 1992. Both species show striking similarities 
in dental morphology to Tedford' s Site D maxil- 
lary fragment. 

This paper describes these new species and 
provides a phylogenetic analysis of the 
Zygomaturinae based on dental features. Partic- 
ular attention focuses on P\ which tooth appears 
most useful in diprotodonloid phylogenetic anal- 
yses (Stirton et al, 1967). 

Material is deposited in the palaeonlological 
collection of the Queensland Museum (QMF) 



and the palaeontological collection of the Bureau 
of Mineral Resources, Canberra (CPC). Cusp 
nomenclature follows Archer (1984) and Rich el 
al. (1978) except that what was then understood 
to be the hypocone of upper molars is now ac- 
cepted to be the metaconule following Tedford & 
Woodburne (1987). Molar homology follows 
Luckett (1993). Premolar homology follows 
Flower (1867). Higher level systematic nomen- 
clature follows Aplin & Archer (1987). 

SYSTEMATICS 

Superorder MARSUPIALIA Illiger, 1811 

Order DIPROTODONTIA Owen, 1866 

Family DIPROTODONTIDAE Gill, 1872 

Subfamily ZYGOMATURINAE Stirlon, 

Woodburne & Plane, 1967 

Silvabestius gen. nov. 

TYPE SPECIES. Silvabestius johnnilandi sp. nt)v. 

OTHER SPECIES. Silvabestius michaelbirti sp. nov., 
Palorchesiinae gen. and sp. indct. of Tedford (1967) 
{^Silvabestius sp. herein). 

DIAGNOSIS. Silvabestius differs from other 
zygomaturines in the following combination of 
features: small parastyle on P-^; absence of a deep 
trench separating parastyle from parametacone 
ba.se on P^; absence of a well-developed lingual 
margin on upper molars; presence of upper ca- 
nines (except Neohelos)\ absent or small 



94 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 1 . SilvabesthisjohnnUandi gen. et sp. nov.. holo(ype, QMF30504. A, righi lateral view. B, ventral view, C, 
dorsal view. Bar = 20mm. 



hypocone on P-^ (except Alkwertatheriiim wehbi); 
more anteriorly convex upper molar lophs (ex- 
cept kn-Nimbadon). It isdistinguishcd from other 
zygomaiurines except Raemotherium yatkolai 



and Nimbadon by: its small size; molar gradient 
that docs not appreciably increase posteriorly. It 
is distinguished from other zygomaturines except 
Nimbadon and Neohelos by: ihc antcrobuccal 



PRIMITIVE ZYGOMATURINE SILVABESTIUS GEN. NOV. 



195 




FIG. 2. Silvabestiusjohnnilandi gen. et sp. nov., hololype, QMF30504. Occlusal stereopair of upper cheektooth 
deniition. Bar= 10mm. 



blade from the parametacone on P^; absence of a 
well-developed buccal cingulum or metastyle on 
P^. It can be distinguished from other 
zygomaturines except Neohelos, Nimbadon and 
A. webbi by an undivided parametacone on P3. 

ETYMOLOGY. Latin silva, forest and bestia, beast; 
for its inferred habitat; masculine. 



Silvabestiusjohnnilandi sp. nov. 
(Figs 1-6) 

MATERIAL. Hololype QMF30504, a juvenile skull 
and associated mandible with completely unworn left 
and right cheek tooth rows. The basicranium, palate 
and nasals are incomplete. Both dentaries are missing 
the coronoid process. I-'^ is missing on each side. 
P^and M-^*^ on each side were unerupied at the time of 



196 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 3. Silvabesttus johnnilandi gen. el sp. nov., right 
type QMF30504. Occlusal stereopair. Bar= 10mm. 

death. Paratype QMF30505, a virtually complete adult 
skull; its fronlals are broken and slightly depressed; a 
large fracture runs diagonally IVom the left orbit to the 
dorsal surface of the premaxilla and continues ventrally 
through the right canine alveolus and onto the palate. 
The right palatine bone is fractured and depressed. The 
left mastoid/paroccipital process is broken. The supra- 
occipital is incomplete. Both types from VIP Site 
which is 3m below D Site; the latter contains 
diprotodontoids some of which belong to genera that 
occur in the late Oiigocene in the Etadunna Foration 
(Woodbumc et al., 1994) and thus we assign VIP Site 
to the late Oiigocene. 

Although a juvenile the holotype has perfectly unworn 
dentition which maximises information about crown 
morphology and both the skull and lower jaws, whereas 
the paratype is only a skull. 

ETYMOLOGY. For Professor John Niland, Vice 
Chancellor of the University o{ New South Wales who 



has been a strong supporter of the 
Riversleigh Project and helped collect 
at Riversleigh. 

RELATIONSHIP OF THE HOLO- 
TYPE TO THE PARATYPE. We 

conclude that QMF30504 was a de- 
pendent pouch-young living on its 
mother's (QMF30505) milk be- 
cause; 1) both specimens were in 
the same deposit, within I m of each 
other; 2) the lack of wear on the 
teeth of QMF30504 most of which 
were still erupting and the lack of 
fusion of any of its cranial bones; 3 ) 
when the upper and lower cheek 
teeth of the juvenile are brought into 
occlusion, the upper and lower inci- 
sors do not meet, but arc separated 
by a gap of approximately 6 mm, a 
gap appropriate to suit a mother's 
nipple. 4) LP of the adult skull, 
which was thought to be missing, 
was found adjacent to the nose of 
the juvenile skull suggesting that 
the adult and juvenile died nose to 
nose, suggesting an emotional rela- 
tionship. 

DIAGNOSIS. This species differs 
from S. michaelbirti in the follow- 
ing combination of features: it is 
much larger; it has an expanded 
paraslylar region on P-^ and more 
dentary of holo- distinct parastyle; it has a more pos- 
terior parametaconc on P^ it has a 
larger protocone on P-^ that is also 
more distinctly separated from the base of the 
parametacone by a deep fissure; it has a small but 
distinct hypocone on P' and it has less steeply 
sloping surfaces on the protolophs and metalophs 
ofM'-'*. 



DESCRIPTION. Upper incisors. L and Rl' in the 
Holotype; L, Rl'- and LL^ in the Paratype. l' 
large, curved, with a thick, rounded base tapering 
to its tip. Enamel confined to the anterolateral 
surfaces and the upper third of the distal surface. 
L and RI^ convergent at their lips. I- small, sub- 
ovate, tapering anteriorly, with narrow anterior 
tip contacting the posterolateral face of I^ with 
crown dominated by a large, ovate wear facet. I^ 
posterior and slightly lateral to I-, smaller than I-, 
with a triangular occlusal surface, with apex ori- 
ented anteromedially, with a wear facet on most 
of the occlusal surface. 



PRIMITIVE ZYGOMATURINE SILVABESTIUS GEN. NOV. 



197 




FIG. 4. Silvabestiusjohnnilandi gen. et sp. nov., right dentary of holotype QMF30504. A, lingual view. B, buccal 
view. Bar = 10mm. 



Upper canines. Adult skull with small canine 
alveolus just posterior to the premaxillary/maxil- 
lary suture; obscured in the juvenile skull. 

P\ Four cusps: a large parametaconc; a well- 
developed protocone; a small distinct parastyle; 
and a poorly-developed hypocone. Widest across 
the protocone. Emargination between the bases 
of the protocone and lobate parastylar region 
defining the anterior and posterior moieties. Para- 
type premolar proportionately longer than that of 
the Holotype and the anterior and posterior moi- 
eties are more distinct. Large, undivided para- 
metaconc the tallest cusp on the premolar, 
situated along the midline of the tooth, slightly 
posterior to its transverse axis. Protocone large, 
lingual ly opposite the parametaconc. Hypocone 
small but distinct, on the lingual cmgulum at the 
posterolingual base of the protocone. Paratype P-^ 
with extremely reduced and non-cuspate 
hypocone. 

Parametaconc pyramidal in occlusal view with 
sleep buccal, anterolingual and posterolingual 
faces. Each face defined by a distinct blade: an 



anterior preparametacrista, a posterior 
postparametacrista and an antcrolingually di- 
rected blade. Preparametacrista linear, extending 
anteriorly (and slightly buccally) from the 
parametaconc apex, continuous with a short, ar- 
cuate, posterobuccal blade from the parastyle. 
The anterolingually-oriented blade descending 
the anterolingual face of the parametaconc, ter- 
minating in the fissure that separates the bases of 
the protocone and parametaconc, just prior to 
meeting its counterpart, a short, linear, an- 
lerobuccally-dirccted protocone crest. A similar 
condition is found in Nimbadon lavarackorum 
(Hand et al., 1993a) and some Neohelos speci- 
mens from the Oligocene-Mioccne of Rivers- 
leigh. In the paratype, the anterolingual 
parametaconc blade is continuous with the an- 
terolingual cingulum, a feature previously re- 
garded as an aulapomorphy of Nimbadon (Hand 
et al., 1993a). A slight swelling at the junction of 
the anterolingual blade and the lingual cingulum 
may represent a small protostyle. 



198 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE L Measurcmcnis (mm) o\' Silvabestius. 
H=Hcighl of crown; A/P=anlcrior-posterior length; 
Th=Thickncss; W=widih; IMP'' = implantation 
angle; #=broken specimen; *=dimensions of alveoli. 



A. l' MEASUREMENTS | 


Species 


No 


Side 


H 


A/P 


W 


jolmnilandi 


QMF 

30504 


L 


7.1 


- 


6.5 


R 


8.3 


- 


6.7 


QMF 

30505 


L 


I6.0# 


8.7 


6.9 


R 


20. 1# 


8.3 


6.9 


michaelhirii 


QMF 

20493 


L 


18.9# 


7.2 


7.0 


R 


18.3# 


8.1 


6.8 


B. r MEASUREMENTS 


johnniliituii 


QMF 

30505 


L 


5.4 


6.4 


5.0 


R 


5.4 


6.3 


5.1 


niichaelbini 


QMF 

20493 


L 


. 


5.9* 


4.5* 


R 


5.7 


6.0 


4.7 


C. v'' MEASUREMENTS 


johnnilancii 


QMF 

30505 


L 


4.7 


5.3 


4.1 


R 


- 


- 


" 


michaelhirii 


QMF 

20493 


L 




5.3* 


4.8* 


R 


- 


5.2* 


4.3* 1 


D, UPPER CANINE MEASUREMENTS 


j oh tint land i 


QMF 


L 


3.0* 


2.2* 


30505 


R 




3.2* 


2.1* 


inichaelhirii 


QMF 

20493 


L 


6.3 


5.1 


3.5 


R 


6.2 


5.2 


3.5 


E. LOWER INCISOR MEASUREMENTS 






i__ 


Th 


W 


IMP^ 


johnnilandi 


QMF 

30504 


L 


7.0 


4.7 


30° 


R 


8.1 


4.6 


30° 



Postparamelacrisia wcll-devcU)pcd, blade-like, 
continuous with the poslerolingual cingulum. 

Small, erect paraslyle al the anterolingual cor- 
ner, slightly buccal to the tooth margin, domi- 
nated by a distinct, arcuate blade continuous with 
the prcparametacrista. Paraslyle in paralype re- 
duced to a slight swelling at the anterior lip of the 
preparamctacrista. Short, ill-defined, an- 
terolingual parastyle crest continuous with the 
anterolingual cingulum. Shallow anterolingual 
basin between the bases oi' the parastyle, 
parametacone and protocone, belter detlned in 
holotype RP^, but extremely shallow on the 
crown of the paratype. 

Short, posteriorly-directed, apical prolocone 
crest lerminaling 1/3 the way down ils posterior 
surface. Two short, linear crests extending from 
the apex oi' the hypocone; one extending anteri- 
orly and terminating at the posterior base o\' the 
prolocone; other extending posteriorly, continu- 



ous with the poslerolingual cingulum. Buccal 
cingulum poorly-developed in the Holotype. 
Buccal surface of the parametacone with uni- 
form, steep gradient to the base of Ihe looth 
crown. Well-developed buccal cingular shelf in 
the Paratype. 

Upper molars. Proportionately similar to 
Nimbadon whitelawi from the mid-Miocene Bul- 
lock Creek Local Fauna (Hand el al., 1993a). 

M'. Sub-rectangular, low-crowned, trans- 
versely-bilophodont, with an anterior proiok)ph 
and posterior metaloph. Prololoph and meialoph 
short, anteriorly-convex crests, former slightly 
more arcuate than the latter; prololoph consisting 
of a buccal paracone and a lingual protocone; 
metaloph consisting of a buccal metacone and a 
lingual meiaconule; metaconule the tallest cusp, 
followed by the paracone, metacone, then pro- 
tocone. Parastyle small, at the anterobuccal end 
oflheanteriorcingulum, connected to an anterior 
paraconal crest by a short, posteriorly-oriented 
blade, Paratype paraslyle poorly developed, 
slight swelling at the junction of the anterior 
paraconal crest and the anterior cingulum. 
Postmetacrisla distinct, extending posteriorly 
from the metacone apex, continuous with the 
posterior cingulum. Meiastyle a slight swelling al 
the posterobuccal margin, most reduced in the 
Paratype. Posterior cingulum terminating al the 
posterolingual base of the metaconule. Lingual 
cingulum in Holotype poorly developed, in Para- 
type short, crescentic. joining bases o'i the pro- 
tocone and metaconule, blocking the lingual exit 
of the interloph valley. Short, indistinct, 
postparaconal crest descending the posterior face 
of the paracone, terminating in Ihc interloph val- 
ley, before meeting ils counterpart, an indistinct 
prcmetacrista. Short, linear, apical blade on the 
metaconule. Paratype with facets extending 
posterobuccally from the apices of the protocone 
and metaconule, fading down the posterior Hanks 
o^ their respective cusps. 

M""*. Similar to M'exccpt: Larger, proportion- 
ally wider anteriorly, with molars trapezoidal in 
(Kxiusal view; paracone tallest cusp, prololoph 
higher than metaloph; parastyle and metastyle 
and their associated crests extremely reduced, as 
are the postparaconal crest and the prcmetacrista; 
lophs relatively longer, with prololoph longer 
than the metaloph. becoming more anteriorly 
convex in ihe more posterior molars; metaloph 
severely reduced, more obliquely oriented rela- 
tive to the prololoph. with posterior moiety se- 
verely reduced. Molars increasing in si/e through 
M''-. decreasing through M-^"^. M** reduced, lack- 



PRIMITIVE ZYGOMATURINE5//.V/\/?£5r/f/5 GEN. NOV, 



199 




FIG.5. Silvobesiitis johnnilandi gen. el sp. nov., paratype QMF30505. A, right lateral view. B. ventral view. C. 
dorsal view. Bar= 20nim. 



200 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 6. Silvabestiusjohnnilandi gen. et sp. nov., paratype, QMF30505. Occlusal slereopair of upper cheektooth 
dentition. Bar= 10mm. 



ing enamel. Adult cheektooth row with an in- 
creasing molar gradient from M' to M"*, 

Lower dentition. A single pair of lanceolate, 
procumbent lower incisors in the Holotype. 
Enamel only on the lateral and ventral surfaces of 
the crown. A longitudinal groove occupying the 
internal dorsal surface, extending from the base 
of the incisor to the tip, approximately 2mm 
below and lingual to the dorsolateral tooth mar- 
gin. 

P3. Short, subovate, longer than wide, tapering 
anteriorly, dominated by a high, steep protoconid, 
with a short, steep, arcuate blade extending pos- 



teriorly from the protoconid apex, terminating in 
a small cuspid on the posterior cingulid. A poorly 
defined crest descending the anterior face of the 
protoconid, terminating in a slight swelling at the 
anterior tooth margin. Posterobuccal cingulum 
poorly developed. Lingual cingulum curving 
around the posterolingual tooth margin, from the 
apex of the posteromedial cuspid to a point mid- 
way between the protoconid apex and posterior 
tooth margin. A vertical flanking crest descend- 
ing the steep buccal face of the protoconid to a 
point just anterior to the terminus of the lingual 
cingulum. 



TABLE 2. Measurements {mxw)o{SilvabestHis. L= length; AW=anterior width; PW=posterior width; ^estimates. 



A. UPPER CHEEK DENTITION 


Species 


No. 


Side 


P3 


Ml 


M2 


M3 


M4 


L 


W 


L 


AW 


PW 


L 


AW 


PW 


L 


AW 


PW 


L 


AW 


PW 


johnnilandi 


QMF 

30504 


L 


12.2 


10.5 


13.5 


10.4 


10.2 


14.8 


12.4 


11.2 


14.5 


11.9 


- 


12.0 


11.4 


9.0 


R 


11.7 


10.7 


13.6 


10.6 


10.2 


14.5 


12.5 


11.4 


14.3 


12.9 


10.8 


11.5 


11.8 


9.1 


QMF 

30505 


L 


13.1 


U.O 


14.4 


11.9 


11.5 


14.7 


13.2 


11.8 


15.2 


13.2 


11.4 


15.3 


12.8 


9.8 


R 


12.6 


10.9 


13.6 


11.7 


11.4 


15.0 


12.3 


11.2 


15.3 


12.8 


10.8 


15.5 


14.1 


10.7 


michaelbirti 


QMF 

20493 


L 


9.6 


7.5* 


10.4* 


9.4* 


9.0* 


12.9 


10.1* 


9.2* 


12.0 


11.4* 


9.8* 


12.0 


10.2 


8.2 


R 


9.6 


8.3 


11.3 


9.7 


9.3 


ll.O 


10.4 


9.6 


11.9 


10.3 


8.7 


11.3 


10.2 


8.1 


sp. 


CPC 

7337 


R 


12.5 


9.3 


13.5 


10.3 


10.7 


- 


12.4 


" 














B. LOWER CHEEK DENTITION 


johnnilandi 


QMF 

30504 


L 


10.8 


7.! 


13.5 


8.1 


8.9 


14.6 


9.6 


10.0 


14.6 


11.3 


9.6 


12.7 


- 


9.2 


R 


10.3 


6.6 


13.7 


8.1 


8.7 


15.3 


9^6 


9.9 


13.7 


U.O 


10.2 


12.9 


10.8 


9.3 



PRIMITIVE ZYGOMATURlNE5/LVv4B£5r/^/.S' GEN. NOV. 



201 




FIG. 7. Silvabestiusmichaelbirti sp. nov., hololype, QMF20493. A, right lateral view. B, ventral view. C, dorsal 
view. Bar = 20mm. 



202 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 8. Silvabestius michaelbirti sp. nov., holotype, QMF20493. Occlusal stereopair of upper cheektooth 
dentition. Bar= 10mm. 



Ml. Subrectangular, anteriorly-tapering, trans- 
versely bilophodont. Anterior protolophid short, 
steep, high, running parallel to a steep, low, pos- 
terior hypolophid. Protoconid higher than 
metaconid; entoconid higher than the hypoconid. 
Paracristid anteriorly directed, blade-like, linking 
the protoconid to the anterior cingulum, slightly 
buccally convex. Anterior cingulum curving lin- 
gually and downward, to the base of the crown, 
meeting a relatively indistinct preprotocristid. 
Deep anterior basin defined by the junction of 
these crests and the anterior base of the pro- 
toconid. A short, vague crest fading down the 
posterobuccal base of the protoconid. Similar, 
better-defined crest fading down the posterior 
face of the metaconid. A well-developed, an- 
terobuccally oriented prehypocristid terminating 
approximately 1 13 the way down the anterior face 
of the hypolophid. Lingual cingulum discontinu- 
ous, comprised of 2 ridges extending 
posterobuccally and postcrolingually from the 
bases of the protoconid and hypoconid, respec- 
tively, with both ridges terminating just short of 
meeting each other in the interlophid valley. Sim- 
ilar short ridge at the posterobuccal base of the 
metaconid. Posterior cingulum short, not extend- 
ing around the bases of the entoconid and 
hypoconid. Transverse valley U-shaped in lin- 
gual view. 



M2. Similar to Mi in all aspects except: slightly 
larger; protolophid subequal to its hypolophid; 
protolophid and hypolophid transversely wider; 
paracristid and prehypocristid extremely re- 
duced; all ridges associated with the main cuspids 
reduced; anterior face of the protolophid steeper; 
anterior cingulum better-developed; anterior 
basin absent. 

M3-4. Only partially erupted, similar to M2 in all 
aspects except: M3 less elongate, but wider than 
M2; M4 reduced relative to M2-3; hypolophid re- 
duced relative to the protolophid, slightly offset 
relative to the protolophid, more posteriorly con- 
vex; interlophid valley more open and broadly 
U-shaped. 

REMARKS. Variation in the hypocone is com- 
mon among Oligo-Miocene zygomaturines. In 
fact, hypocone-less variants have been recorded 
for species of Neohelos (Peter Murray pers. 
comm.) and Nimbadon (Hand et al., 1993a). The 
development of a hypocone has, in the past, 
served as a synapomorphy of the Zygomaturinae. 
We have used this feature in the phylogenetic 
systematic analysis presented below because, al- 
though variable, it is generally a good indicator 
of phylogenetic relationships. 



PRIMITIVE ZYGOMATVRINESILVABESTIUSGEN. NOV. 



203 




FIG. 9. Silvabesiius sp., CPC7337. Occlusal view of 
right maxillary fragmenl with P-^-M- from Site D 
Locality, Riversleigh. (After Tedford, 1967). 

Silvabestius michaelbirti sp. nov. 

(Figs 7-8) 

MATERIAL. Hololype QMF20493, a relatively com- 
plete cranium from late Oligocene Hiatus (South) Site 
(Creaser, 1997) with left and right check tooth rows; a 
longitudinal fracture runs through LM''-^; RM' is frac- 
tured diagonally from the anierolingual tooth comer to 
the posterobuccal tooth corner: L and RI* are present 



TABLE 3. Character state polarities for selected 
zygomaturine genera using species of Ngapakaldia 
(Palorchestidae) as an outgroup. A=absent; P=pres- 
em; S=small; D=distinct; W=weak; ST=strong; 
SQ=square; R=reduced; ==equal or sub- 
equal. PLESIO=plesiomorphic;APO=apomorphic. 



CHARACTER 


PLESIO- 


APO- 


1 . Parastyle development on P 


A 


P 


2. Hypocone development on P 


A/S 


D 


3. Division of parapietacone into two 
distinct cusps on P 


A 


P 


4. Well-developedjdiagonal crest on 
parametacone of v 


A 


P 


5. Link between pr9tocone and 
parametacone on P 


AAV 


ST 


6. Mesostyle retracted towards 
cingulum 


A 


P 


7. Size of P relative to equal molars 


S 




8. Elongate P" 


A 


P 


9. Buccal cingulum on P 


ST 


W 


1 0. Loss, incorporation or suppression 
of stylar cusps C and D with respect to 
lophs 


A 


P 


1 1 . Small parastyle on M 


P 


A 


12. Occlusal shape of m' 


E 


SO 


13. Short posterior moiety on Pi 


P 


A 


14. Paracnstidon Mi 


P 


R 



but incomplcle; left I- and L and Rl-' arc missing; the 
cranium is in two parts, the division occurring approx- 
imately 15mm posterior to the molar rows; the nasals, 
frontals and palate arc incomplete; the basicranium is 
relatively intact; the posterior region of the neurocran- 
ium is missing on each side. 

ETYMOLOGY. For the former Vice Chancellor of the 
University o\' New South Wales, Professor Michael 
Birt, who assisted in the collection of specimens and 
helped meet the cost of preparation.. 

DIAGNOSIS. Differs from S. johnnilandi by: 
being smaller; having canines; having reduced 
parastyle and parastylar region on P-^ and less 
elongate P-^; having a more central parametacone 
on P-^; having a smaller protocone on P-^ that is 
less distinctly separated from the base of the 
parametacone; lacking a hypocone on P-*; and 
having steeper protoloph and metaloph faces on 
M''"*. Differs from Silvabestius sp. in: being 
higher crowned; lacking a well-developed 
parastylar region on P^; and having a reduced 
posterolingual cingular shelf on P-^. 

DESCRIPTION. This species is described in so 
far as it differs from other species of the genus. 
Incisor arcade poorly-preserved with only the L 
and R I' (both of which arc incomplete) and the 
right I^ remaining. Incisors as in S, johnnilandi. 

Canines small, pointed, on the premaxillary- 
maxillary suture whereas in 5. johnnilandi the 
tiny canine alveoli lieapproximately 2mm behind 
the premaxillary-maxillary suture. 

Left cheektooth row badly fractured with a 
large fissure extending across the buccal margin 
of M', diagonally through M- and across the 
lingual region of M^, terminating at the anterior 
base of the hypolophid of M^. Right M^ fractured 
diagonally from the anterolingual tooth comer to 
the posterobuccal tooth comer. 

P-^ smaller, lacking a distinct cuspate parastyle, 
with parastylar region reduced; vague pre- 
paramelacrista continuous with the anterior cin- 
gulum; parametacone more posteriorly along the 
longitudinal axis, with postparametacrista more 
steeply inclined; protocone smaller, less dis- 
tinctly separated from the base of the 
parametacone; hypocone absent; protocone with- 
out crests. 

Upper molars. Lower-crowned, with more ex- 
tensive wear facets, with more steeply inclined 
posterior surfaces of the protoloph and metaloph; 
with a molar gradient not increasing posteriorly. 



204 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 10. Comparison of left P^ of Silvabesii us: A-A', 
occlusal stereopair of 5. johnnilandi, holotype, 
QMF30504. B-B', occlusal steropair of 5. 
johnnilandi paralype QMF30505. C-C, occlusal ste- 
reopair of 5. michaelbirti holotype QMF20493. Bar 

Silvabestius sp. 

(Fig. 9) 

MATERIAL. CPC7337, a right maxillary fragment 
with P-^, M^ and the anterior half of M- from late 
Oligocene Site D Locality, Riversleigh. 



DESCRIPTION. See Tedford (1967). 

REMARKS. The teeth are missing most of their 
enamel so our understanding is based on dentine 
core morphology. Compared to Silvabestius 
johnnilandi this specimen is lower-crowned 
(though the extent to which this is an artefact of 
enamel loss is not known); the protocone on P-"^ is 
smaller and is less distinctly separated from the 
base of the parametacone; and a hypocone is 
(most probably) absent on P-^. 

This specimen is not designated a new species 
because of its poor preservation. The lack of 
enamel on the crown makes it difficult to interpret 
the stage of development of the parastyle and 
hypocone. Variation between adult and juvenile 
dentitions of S. johnnilandi, suggests the above 
mentioned differences in dental moi^^hology may 
not warrant specific distinction. 

PHYLOGENETIC ANALYSIS 

Aplin & Archer (1987) and Marshall et al. 
(1989) recognised the Palorchestidae, divided 
into Palorchestinae and Ngapakaldinae, and the 
Diprolodontidae, divided into Diprotodontinae 
(sensu Archer, 1 977 ) and Zygomaturinae. Archer 
& Bartholomai (1978) united these two families 
in the Diprotodontoidea. The monophyly of these 
groups is under question as well as the basis for 
their higher level association (Archer, 1984; 
Aplin & Archer, 1987; Murray, 1990a). Conse- 
quently, it is difficult to select an appropriate 
outgroup for phylogenetic analysis of the 
Diprolodontidae and to determine character state 
polarities within the group (Murray, 1 990b; Hand 
et al., 1993a). We choose the palorchestids 



TABLE 4. Distribution of character stales in representative diprotodontoids. 0= plesiomorphic stale; 
l=apomorphic state; 2=most apomorphic stale; a=multistate character; ?=missing data. 



Taxon 


1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


11 


12 


13 


14 


Ngapakaldia 












































Diprolodontines 








































1 


Z\gomoturus 


2 


1 


1 





1 


1 


a 








, 


2 


1 







Ko I ops IS 


2 


I 


1 





1 


I 












2 


1 







Neohelos 


2 


1 








a 


1 












2 


1 







Kolopsoides 


2 


2 


1 





1 





1 


1 


1 




1 


1 







Nimbadon 


2 


1 





1 


















1 


a 







Plaisiodon 


2 


I 








1 





1 


1 


1 




a 


1 







Alkwertatherium 


2 






























I 




1 


Silvabestius 


1 








1 


















a 










Raemeotherium 


7 


? 


7 


7 


9 


7 


7 


7 


7 


9 


9 


9 


9 






PRIMITiVEZYGOMATURINE5//.VM5£57/L^5GEN. NOV 



205 



Ngapakaldia and Pitikantia as outgroup for anal- 
ysis of the Zygomalurinae due to their 
plesiomorphic position in the Diprotodontoidea. 

CHARACTER ANALYSIS. Fourteen dental 
characters considered taxonomically useful in 
zygomaturine intergeneric relationships are em- 
ployed in a cladistic analysis (Table 4). Selection 
of characters follows previous phylogenetic anal- 
yses by Murray ( 1 990b) and Hand et al. ( 1 993a). 
Character state polarities (Table 3) were deter- 
mined by outgroup analysis using Ngapakaldia 
and Pitikantia. Character state polarities for 
Nimbadon were scored only for Ni. lavarackoriim 
andM'. whitelawi, for reasons given in the discus- 
sion. /?«e/7U'o//zp/7"f//7rvar^o/r// was excluded from 
the phylogenetic analysis because 13 of the 14 
characters are unknown for this species. 

A Wagner analysis was performed using the 
PAUP (version 3.1.1) computer program 
(Swofford, 1993). Multistate characters were 
treated as polymorphisms. Character optimisa- 
tion was performed using both the accelerated 
(ACCTRAN) and delayed (DELTRAN) transfor- 
mation algorithms. 

CHARACTERS EXCLUDED FROM THE 
ANALYSIS. For many characters examined 
from previous phylogenetic analyses, intraspe- 
cific and interspecific variation was found to be 
high. Similarly high variation is characteristic of 
the Oligo-Miocene zygomaturines Neohelos and 
Alkwertatherium (Murray, 1990b). Characters 
with a high degree of intraspccific variation ex- 
cluded from the analysis include: the degree to 
which the parastylc of P-^ is posteriorly inclined 
or 'hooked'; shape of the lingual cingulum on P3; 
presence or absence of a proloslyle on M ' ; extent 
of metaloph reduction on M-^"*; extent of reduc- 
tion of M"^. 'Proportional similarity' of P-^ as a 
synapomorphy for a Plaisiodon/ Nimbadon/ 
Neohelos/ Kolopsis/ Zygomaturus clade, used by 
Murray (1990b) and Hand et al. (1993a), is a 
character complex relating to the development of 
the parastyle, protocone and hypocone. Accord- 
ingly, it is not included as a discrete character. 

RESULTS. Wagner analysis using the branch 
and bound algorithm generated a single most 
parsimonious tree (Fig. II) involving 23 steps 
with aconsislency index (CI) of 0.880. a retention 
index (Rl) of 0.880 and a rescaled consistency 
index (RC) of 0.774. The high consistency index 
probably retlects the limited number of charac- 
ters used in the analysis. Tree topology was iden- 



I I 


3 


: : 
5 J 


■ 2 


i 1 1 




5 
^ 


1 


1 














Qo 




[I|o->i 

[q]o->i 




^ l->2 


[T|i»->i 




T|o->i 






[j]c>->i 
[IT]o->i 






[i]o->i 




[T|v>; 

[l2ll)->1 






[4]0->l 




[T|()->i 

[7]0->l 
0O->1 










^fr>l 







FIG. II. Hypothesis of zygomaturine relationships, 
based on dental characters (Table 3) used by Murray 
(1990b) and Hand et al. (1993) in cladistic analyses 
of the Diprotodontidae. Apomorphies are represented 
by boxed numbers. Arrows indicate character state 
transformations. Symbols: 0, plesiomorphic character 
state; 1 , derived character state; 2, most derived char- 
acter slate. 

tical irrespective of whether ACCTRAN or 
DELTRAN character optimisation was em- 
ployed. 

DISCUSSION 

Silvabestius is the most plesiomorphic 
zygomaturine known with the possible exception 
of Raemeothehum yatkolai. Silvabestius is basal 
to the diprotodonloid radiation. Although similar 
to the primitive palorchestid Ngapakaldia in den- 
tition (Tedford. 1967) and the middle ear, 
Silvabestius is assigned to the Zygomaturinae 
because il possesses a number of zygomaturine 



206 



MEMOIRS OF THE QUEENSLAND MUSEUM 



synapomorphies including a parastyle on P^ and 
a ventral alisphenoid tympanic process. 

Stirton et al. (1967) and Murray (1990b) sug- 
gested that zygomaturines evolved from primi- 
tive diprotodontine-likc forms in which the 
parastyle on P^ was small. Trends in premolar 
morphology in Silvabestius support this hypoth- 
esis. Transition from the simple, bicusped 
diprolodontine P-^ to the plesiomorphic, 3-cusped 
zygomaturine premolar of 5. michaelbirti is 
achieved through expansion of the parastylar re- 
gion (resulting in a more elongate premolar) and 
the posterolingual cingular shelf. Subsequent 
transition to the more typical 4-cusped 
zygomaturine premolar of S. johnnilandi is 
achieved through development of a distinct 
parastyle on the enlarged parastylar region and 
development of a distinct hypocone on the en- 
larged posterolingual cingular shelf. 

Silvabestius \s the plesiomorphic sister group to 
all other zygomaturines (Fig. 9). Plesiomorphic 
features of the dentition include a poorly-devel- 
oped parastyle, a poorly-developed hypocone, an 
undivided parametacone on P'\ and a well-devei- 
oped paracristid on M|. 

Monophyly of Zygomalurinae is supported by: 
a parastyle on P\ P^ equal or subequal to the 
molars in length; and an enlarged posterior moi- 
ety on P3. The zygomaturine sister-group of 
Silvabestius is united by synapomorphic a mod- 
erate to large parastyle on P^ and square vs elon- 
gate molars (in occlusal view). A distinct 
hypocone on P-^ and a well-developed parastyle 
on M' are shared by Nitnbadon, Kolopsoides. 
Plaisiodon, Neohelos, Kolopsis and Zygo- 
maturus. The latter genera, with the exclusion of 
Nitnbadon. is united by a linking crest between 
the apices of the protocone and parametacone on 
P-*. Kolopsoides and Plaisiodon are united by an 
elongate P- and a weak buccal cingulum on P"*. 
Neohelos, Kolopsis and Zygomaturus form a 
clade rationalised by amesostyle that is retracted 
towards the cingulum on P^ and by a large 
parastyle on M' . Kolopsis and Zygomaturus are 
united by a parametacone on P^ that is divided 
into two distinct cusps. 

Topology of the cladogram (Fig. 9) is consis- 
tent with that found in Hand et al. (1993a) and 
Murray (1990b). However, it differs in the rela- 
tive positions of species of Nimbadon, Plaisiodon 
and Kolopsoides. These differences result from 
the exclusion of certain characters used in previ- 
ous analyses which are of low taxonomic value. 
A hooked or posteriorly inclined parastyle on P"* 
has been used by Murray (1990) and Hand et al. 



(1993a) as a synapomorphy uniting Plaisiodon 
and Nimbadon. In this analysis the character was 
found to be highly variable and hence of low 
taxonomic value. No special affinity was found 
between Nimbadon and the Kolopsoides/ 
Plaisiodon clade. Further, Hand et al. (1993a) 
united Nimbadon, Plaisiodon, Neohelos, Kolop- 
sis and Zygomaturus to the exclusion of 
Kolopsoides by the similarly proportioned P-^. 
This character was excluded from the current 
analysis because it was found to be dependent on 
the degree of development of the parastyle, pro- 
tocone and melacone on P-^. 

The position of Kolopsoides cultridens within 
the Zygomaturinae is difficult to determine, 
partly because of the paucity of knowledge about 
late Miocene diprotodontoids. Stirton et al. 
(1967) suggested that K. cultridetis is most 
closely related to Kolopsis on the basis of a di- 
vided parametacone on P\ Flannery (1994) sug- 
gested that all New Guinea diprotodontids, 
including K. cultridens, are closely related and 
probably stemmed from a Kolopsis-Wke ancestor. 
K. cultridens is here linked with the late Miocene 
Plaisiodon centralis, from the Alcoola Local 
Fauna, a relationship first proposed by Archer 
(1984). If this relationship is accepted a divided 
parametacone has developed within the Zygo- 
maturinae in the Kolopsoides/ Plaisiodon lineage 
and again, convergently, in the Neohelos/ A.'o/o/?- 
5/5/Zv^(9//i«r//n/A lineage. Homoplasious division 
of the parametacone into 2 distinct cusps would 
then have developed during the late Miocene 
perhaps in response to increasing dryness. With 
plant materials becoming less succulent and more 
abrasive, there may have been strong selective 
pressure on all lineages to increase molariform 
features as well as the surface area of premolars. 

We suggest that Nimbadon scottorrorutn is 
more appropriately assigned to Neohelos. Fea- 
tures of the dentition that align Ni. scottorrorum 
with Neohelos are; larger, squarer molars; more 
robust cingula on the cheekteeth; greater 
parastyle development on M^-; a relatively erect 
parastyle on P-^; and an anterolingual crest ex- 
tending from the parametacone apex that meets a 
buccal crest from the apex of the protocone (a 
feature common in species of Neohelos) rather 
than continuing on to meet the anterolingual cin- 
gulum as in Ni. lavarackorum and Ni. whitelawi. 
Material referrable to a small species oi Neohelos 
from Riversleigh's System A Local Faunas, and 
forms intermediate between this small Neohelos 
species and N. tirarensis from Riversleigh's Sys- 
tem B Local Faunas, are relatively indistinguish- 



PRlNfmVEZYGOMATURlNE StLYAliESTWS GBi. IsOV- 



2W 



able in terms of morphology and dimensions from 
ihc llololype and only specimen of Ni. 
icattmronuiu QMF23I57. Consequently, char- 
acter polarilies for species ol Nimbadon in Ihis 
analysis were determined using A'(. la\'- 
antikomm and Ni. whitt'lawi ou\y. 

Similarities in dentitions of Silvubcsrius- 
johnniUmdi and Nimbadon lavamckorion, both 
friHn Rivcrslcigh. and Ni whitelawi. from the 
Bulltx;k Creek Local Fauna include: molar di- 
mensions; .structure of the parastylar region; pro- 
loconc; hypocone; buccal cingulum; and 
anierolinguai parametacone blade on P*. The lasl 
fe^iiure was isuggeMed (Hand el al.. I^*^3a) as an 
autapomorphy for Nimhadon but it is variably 
expressed in Neolielos and well-developed m 
SihahesTitts. Dentitions of 5. jofutnilandi and 
Nimbadon differ mainly m the degree of devel- 
opment of the parastyle on P^ and the extent to 
which it is separated from the base of tlie para- 
melacofie. Plesiomorphic features of Nimhadon 
include a small parastyle and undivided para- 
iuelacone on P\ elongate vn relatively squarish 
mvilars, and a well-developed paracrisiid on Mi. 

Murray ( 1 990b) suggested that Nimbadon may 
be u basal zygtimalurine with affinity to Nvohvlos 
and Plaisiodon; he suggested 2 minor /ygomalur- 
ine lineages. Nimbadon, Neohelos and Kolopsis, 
united Jargcly through plesiomorphic features 
and Alkwertafherium, Plaisiodon and Kolops- 
aides, united through phenelie similarities of ihe 
dcniary and lower incisors. He also suggests that 
these lineages may be united through common 
ancestry in Nimbadon. In our analysts Nimbadon 
occupies a more picsit amorphic position than in 
o^hcr analyses. 

Some doubt rem^tins about whether 
Hacmerodwnnm vatkolal is appropriately in- 
cluded within Diproiodontidac. Richetal.l 1978) 
noted that it exhibited no apomorphic chantcier 
scales that precluded the possibility that it was a 
primitive macropodoid hut, equally, neither does 
it exhibit any character stales that arc undoubted 
nuicropodoid synapt>mo»phies. The relative phy- 
logcnetic positions of Siivabesiins and Raemco- 
;/u'r/wmcannol be resolved until upper dentitions 
ar« discovercil for the laltei. Differences in the 
lower dentition o1\^. johnnikmdi compared with 
R. yatkolai include: il is larger; lophid^ on ilw 
lower molars arc less anteriorly convex; the antc- 
liof mniely of Mi js broader; ihe prernelacrislid 
of Mi is bewcr developed as is the unterolingual 
basin M Uic anterior base of the pioiolophid. the 
pmioslyhd I. Rich cl al.. 1978) al Ihe buccal Kise 
ul the [Hvtoconid wi M| h ahMint: and ihcpru- 



tolophid on Mi ts more pc»slenuriy inclined. B*>lfi 
species exhibit an Mi: My raiio (equivalent to M?; 
M4 in Rich ct al.. 1978) approaching 1 , a condi- 
lion regarded as primitive among diprolo- 
dontoids. There are no apomorphic Jeaiures o( the 
lower dentitions that enable either to be regarded 
as unambigviously more plesiomorphic than the 
other. 

Sil\*<ibcstii4sjohnnilandi is derived relative loS, 
mkhsxelbirii as evidenced by the following fea- 
tures of its P-^: a more distinct parastyle; a larycTw 
lobaic parastylar region; a better defined prepara- 
metacrisla; dv\\\ a &jn-iill but distinct hypocone. 
The relative |xisjtion of Sihalyesiius sp. is more 
diftlcult to determine because of poor preserva- 
tion. However, the following features of the den- 
tine core suggest that Silvubestins sj), is 
strut-turally inteTmedi;iic between S. michaelbini 
and 5. jtfhnnilamii: parastylar region more exten- 
sive than S. michavlbirti but reduced in compar- 
ison to S. johnnilandi\ hypocone absent in S. 
michaelbini, a more extensive posten)lingiial 
cingular shelf in .S. sp., a feature that anticipates 
the development t»f a hypocone in S. jolmndandi. 
Silvalyestius sp. has no aulapomorphic features m 
die dentition that would preclude it being the 
ancestor of 5. johnnilondi. Allernalivcly. 
Silvabestius sp. may represent an eMreme end of 
an SJohnniianJi morphocline, If iliis is the case. 
The simple denial structure of 5. michaelbini -dJid 
the absence of any autapomor[»hic features of the 
dentition, suggest this species may represent the 
direct ancestor ofS.johnnilandi and the slruclurul 
antecedent of all other /ygomaturincs. 

ACKNOWLbDGEMFNTS 

We thank Peter Murray and Richard Tcdford 
for critically reading a draft of this manuscript. 
We acknowledge support from: Auslralum Re- 
search Grant Scheme. The University of New 
South Wales, National Rstate Grants Scheme 
(Queensland), World Heritage Unit of Ihe De- 
partment of Environment. Spotis and Territories, 
Queensland National Parks and WiJdIile Servic<r 
(particularly Paul Sheehy), Waanyi People and 
Carpentarian Land Councd, ICI Australia, Aus- 
tralian Geographic Society, Queensland Mu- 
seum; Australian Museum. Century Zinc 
(particularly D<iug Fisliburni, Mount Isa Mines. 
Mount I.sa City Council. Surrey Beatiy & Sons. 
Riversleigh Society Inc., blaineClark. Sue & Jim 
Lavarack. Sue A: Don ScoU-Orr^ Margaret Beavii 
and Martin r)icV;5on; tvscaa-h colk^igues iiolahly 
Hcnk Godlhclp. Sii/annc Hand. Bemic Cooke. 



im 



MEMOIRS OF THE QUEENSLAND MUSEIFM 



Alan Barlholomai, Phil Creaser, Peicr Murray, 
David Ride. Anna Gillespie, Virginia 
O'Donoghue, Cathy Nock, Syp Praseuthsouk, 
Sicphan Williams and Gill GcJode, and numerous 
postgraduate students working on Rivcrsleigh 
fossil materials. Particular thanks are due to Alan 
Rackhain for assistance m the field. 

LITERATURE cm* D 

APLIN. K. &. ARCHER. M. 1987, Rcccni advances in 
ncarsupial sysicmaiics Willi a new synciviic das- 
sificalion, Pp. xv-lxxii In M Archer (ed.). Pos- 
sums and opossums: studies in evolution. (Surrey 
Bcaliy <& Sons and Royal Zoological St»cicly o( 
NSW: Sydney). 

ARCHER, M. 1977. Ongins and suhfamilial relation- 
ships of Dipwtodtm ( Diprotodoniidae^ 
Alarsupialia). Memoirs of the Queensland Mu- 
seum 18:37-39. 
1984. Tl;e Australian marsupial radiation. Pp. 633- 
8().S In Archer, M. Si. Chiyion, G (eds), Verte- 
brate zoogeography vatid evolution in Australasia 
(Hesperian Pa^ss: Perth). 

ARCHER. M. ik BARTHOLOMAi. A. 1978. Tertiary 
tnnmmals of Australin: a synoptic review W- 
clieringa2: 1-19. 

ARCHER. M. & BLACK. K. 1995. A moment oi 
motherhood* a reconstruclion of an ancient drama, 
Riverslcigh Note^ 25: 4-5- 

ARCHER. M.. GODTHELP. H.. HAND. S.J. ik 
MEGIRiAN, D. 19S9. Fossil mammals ol 
Riverslcigh, northwesiem Queensland: prelimi 
nar)' overview of biosiratigraphy. correlation and 
environmental change. The Australian Zoologist 
25: 35-69. 

ARCHfiR. M , HAND, S J. & GODTHELP, H. 1994. 
Riversleigh, 2nd Ed (Reed Books: Sydnev). 

FLOWER. W H. IS67. On the devcjopmem and suc- 
cession of teeth in the Mursupiatia. Philosophical 
Transactions of the RoyalSocieiy of London 157: 
631-641. 

HAND. S.J.. ARCHER. M.. GODTHEt.P. If. RICH, 
Til ^. PLEDGE, N.S. 19933, f^imhadon.-dnew 
genus and three new species of Tertiary 
/ygtHnaliirincs (Marsupialia; Diprotodontidac) 
I'rorn norilicm Ausiralid, with a reas-sessmcn* of 
Ni'olwhs. Mcmoir>. of ihc Queenskmd Museum 
33: 193-210. 

HAND. S.J.. ARCHER. M., GODTHELP, H. I993l.r 
■^Ruemeoiherium whitwonhi, a new primitive 
zygomalunne (.Marsupialia, Diproiodontidae) 
from Tertiary hmeslones on Rivcrsleigh Station* 
northwestern Queensland (unpublished). 



LUCKETT, VV P 1993 An ontogenedc assessment of 
dental homologies nitherian m:immals Pp. 182- 
2(H. In F.S. S/ala>, M.J. Novacek Sl MC Me- 
Kenna (cds). Miimmal phylogeny: Mcsozoic 
differentiation, multituberculates. monoiremes, 
early iherians and marsupials. (Springer-Verlag: 
New York) 

MARSHALL. L.G., CASE. J.A & WOODBURNE. 
MO. 19S9 PhyUigeneticrciationshipsorihc lam- 
ilics of marsupials. Current Mammalogy 2' 433- 
502 

MURRAY. P. I99(Ja. Primitive marsupial lapirs Pw- 
pahrvheslcs rwvatuiavephahts Murray and P 
ponticuhis sp. nov) from the mid-Miocene Kyi 
noith Australia (Marsupialia- Palorcheslidae), 
The Beagle, Records i^i the Northern Territory 
Museum of Arts and Sciences 7: 39-51. 
1^9()h- Alkwertatherium wehbi.. a new zygomaiur- 
inc genus and species from Ihc late Miocene 
Aleooln Local Fiiuna, Northern Territory 
(Marsupialia: Diproli>dontidae).The Beagle, Re- 
cords of (he Northern Territory Museum of AiU 
and Sciences 7: 53-80. 

MYERS.T& ARCHER, M. 1994. A caaiiouseorrcla- 
lion. Rivcrsleigh Notes 23: I L 

RICH.TH.. ARCHER. M.&TEDFORD, R.H. 1978. 
Haemeothcriitm yatkoliii gen, et sp nov a priini- 
live diprotodonlid from the medial Miocene oi 
Souih Australia. Memoirs of the National Mu- 
seum of Victoria 39: S5-9I, 

STIRTON. R A., WOODBtJRNE, MO. & PLANE. 
M.D. 1967. A phyiugeny of ihe Tertiary 
Diproiodontidae and its significance in corrcla- 
lion. Bullcrin of the Bureau i»f Miticml Resources. 
Geologv and Geophvsics. AusUalia S5: 149- 160. 

TEDFORD. R. H. 1967. Fossil mammal rem;uns fmm 
ihc Carl Creek Limestone, norihwesiem Queens- 
land. Bulletinofihe Bureau of Mineral Resources. 
Geology and Geriphysics. Australia 92: 217-237. 

TEDFORD. R H. & WOODBURNE, M.O. I9S7. The 
llariidae, a new family of vombaliform niarsupiiils 
from Miocene strata of South Ausiralia and an 
evaluation of (he homology of molar cusps in Ihc 
Diprotodontidac. Pp. 41)1-41 8. In M. Archcrfed.), 
Possums and opossums: studies in evolution. 
(Surrcv Bcalty & Sons and Royal Zoological So- 
ciciv of NSW: Sydney). 

WOODBURNti MO,, MCFADDEN, B.J . CASE, 
J. A.. SPRINGER- M.S., PLEDGE. N S-, 
POWER, J.D., WOODBURNE. J.M. & 
SPRINGER, K.B. 1994. Land mammal 
biostratigraphy and magnclostratigntphy of ihe 
Etadunna Formation (Late Oligocene) of South 
Australia. Journal of Vcrtcbrale Paleontology 13: 
483-515. 



NIMIOKOALAGEN. NOV. (MARSUPIALIA, PHASCOLARCTIDAE) FROM 
RIVERSLEIGH, NORTHWESTERN QUEENSLAND, WITH A REVISION OF 

LlTOKOAi^ 

K BLACK AND M ARCHER 

Black, K.. & Archer, M,, 1^97:0(>:30 NimioU'aht gen. nov. (Marsupiaiia. Phascolaraidacl 
from Riversleigh, norihwcsicm Quccnslaad, with a revision ol LiiofsOtila- Memoir^ offhf 
Queensland Museum 41(2): 209-228 Brisbane, ISSN 0079-8835. 

The early lo middle Mincene phascolarciit! Nimtokoala gfeysianesi gen. et sp. nov. Is 
described Irom Rivcrslcigh. northwestern Queensland. Niwiokoala sp, is recognised from a 
iaic Oligocene deposi? in South Australia. The complex molar morphology and 
plcsiomorphic hasicranial morphology of Nimiokoala are indicati\e of a more diverse 
infraorderof phascolarciomorphums. Similariiieb in molar morphology between Niminkoala 
and eklopodon(ids arc noted and may rcflecl similar ecological niche. Litokoala konunkaen- 
six \t described from the Miocene of Riversleigh extending its range and distribution. 
C!;uJistic analyses of dentiil characters of all living and exiinci ia'Sd support the inirafamilial 
relalion^hips proposed by Woodbu me etal.(l*^)87ai.LjYo/cpa/tvi&idemillcda^ the sister group 
of the modern genus andNimioktwla is most closely related lo iht Ldokoala/PhoscohvTtt?s 
clade. D Phase okua'uUe. Nmiokouki. Litokoala. Oltgocem; Mufcene. Hiveniei^h. 

A*, Black & M, An:hen School of Biological ScknCe. University of New South Wales 205Z 
Aiwralia: received 4 November 1996. 



A parlitil skull, representing a small, highly 
specialised phascolarctid was collected fi'om 
Boid Site East, System B deposits (Archer el al., 
1989; 1991 ). Riversleigh. northwestern Queens- 
land- Based on a superficial analysis of dental 
morphologyv Uie skull was incorrectly figured by 
Archer et al (19911 as a new species of the 
Miocene phascolarctid Litokoala, Two species of 
Litokoala were known: L. kutjamarpensis 
(Stinonct al.. 1967) from the Kuijamarpu Local 
Fauna. Wipajiri Formation. Lake Ngapakaldi. 
South Australia (known from a single upper 
molar) and L kanimkaensis (Springer, 1987) 
from the Kanunka North Local Fauna, Eladunna 
Fomiaiion, South Australia (known from 2 iso- 
lated teeth and some rnolar fragments). 

N. grcystanesi gen. el sp. nov. is similar lo 
Liiakoala but its generic separation is substanti- 
ated hy comparison with new material (an M-^ and 
a lower dentition ) of L ka}innkaensis from Sys- 
tem C (Archer el al..l989: 1991) al Riversleigh. 
A further new species of Nimiokoala is 
rccogni.scd from South Prospect B Locality. Lake 
Namba. Frome Downs Station. South Australia. 

The motlcraie abundance oi N. greysfanesi in 
Riversleigh deposits contrasts with the paucity of^ 
material of other phascolarctids which are known 
from isolated leeth or, at best, dentitions. This 
paper describes dentition of the new genus and 
analyses phascolarctid phylogeny based on den- 
tal characters. 

Suprageneric nomenclature follows Aplin & 



Archer (1987). Dental terminology follows 
Archer ( 1 978b) except thai what was then under- 
stood lo be the hypocone of upper molars is now 
accepted lo be the metaconule (Tedford ^ 
Woodburne, 1 987 ). Cheek tooih homology is ihal 
proposed by Luckelt (1993). Biostraligraphic no- 
menclature follows Woodburne el ai. (1993), 
Archer et al. (1989) and Archer et al. (1991). 
Specimens referred 10 are housed in the following 
repositories: Queensland Museum (QMF), South 
Australian Museum (SAMP), University of Cal- 
ifornia al Riverside (UCR), Measurements were 
made using a Wild MS microscope with digital 
length measuring sei. 

SYSTEMATICS 

Order DIPROTODONTIA Owen, 1 866 

Suborder VOMBATIFORMES Woodburne, 

1984 

Infraordcr PHASCOLARCTOMORPHIA 

Aplin & Archer, 1987 

Family PHASCOLARCTIDAE Owen, 1839 

Nimiokoaia gen. nov. 

TYPE SPECIES. Nimiukaalagrcxstivtesi sp. nov. 

ADDITION.AL SPECIES- Nimiokoaia sp. 

DIAGNOSIS. Nimiokoaia differ from all other 
phascolarctids in: being smaller (except species 



2!Q 



MEMOIRS OF THE QIEHNSLAND MUSEUM 



of Liiokoaia): having ;\ large. Citspaie lUrasiylc 
•which is pyramidal in occlusal view (cxicpt 
Litokoaki kiHiamorpensisy, a well-developed, 
cresccniic paraconulc and ncomciaconulcnn M'' 
\ the latter of which is dtiubic cusped in the rnurc 
postefior iiiolats. a marc reduced mciaconc and 
mctaconulc on M^ and a more rounded posterior 
margin ol' that luolh. 

Rohust, ridgc-like crcnulalhins; strong 
posierohuccal ridges irnm the apices of the pro- 
toconc and t^ietaconule; a discontinuous 
nci)meiaconuJe subdivided into two or more parts 
;ind a (variably) discontinuous^ paraconulc; a 
posierolingual cusp on P\, a well-developed 
pnslerolingual crest iroin the apex oi" the pni- 
tosiylid on Mi; a weaker meiaslyiid; a well-de- 
veloped neomorphic cuspid occupying, the 
Irigonid basin between the meliiconid and pro- 
toconid on M: -c, an anierobuccally dirci'icd prc- 
eniocrisiid (as opposed to anierolingually 
diiccred in other phascnlarctids); a wcU-dcvcl- 
opcd cuspate cniostylid ridge; and a more 
postcrohuccally dirccied postproiostylid cristid. 

Nimiokoaia ^reysmnesi dilTcrs I'rom Koohor 
in: being higher crowned; lacking ihe bicusped 
P'; having strong posierolingual paracristac and 
pONlerolingual melacri&tae: having a pi'oiostylc, 
and in having reduced slylar cusps and from 
Madakoala in: being higher crowned; more cren- 
ulaied, in having a more cuspatc, isolaied 
postcrolingual cusp on P^ and in lacking the lin- 
gual cingular ridge of that t(x>th (in M. sp. cf. M. 
wclhi)\ liaving wider, less elongate u[>per molars; 
a more buccal junction of the prcmctacristae and 
postparacristae; a well-developed |5roloslyie; rel- 
atively smaller buccal surfaces of die pafacone 
and metaconc; lacking the transverse connection 
between the metaconc and melaconule on M"*; 
having a less elongate P> thai consists of three 
main apices (as opposed to lour in Motlakcmki). 
a stnall posteiolingual cuspule ond a pioponum- 
atcly larger, more isolated postcrnbuccal cusp on 
that tooth; lacking the well-developed buccal and 
lingual crests from the main apices of P^, having 
a larger, more cuspaie protostylid on Mi; a more 
lingually situated pnoloconid and a more lingual 
junction of the crisiid obi i qua and 
posiproiocrisiid in the Mj , a well-devcloticd an- 
terobuccal ridge off the enioconid aficx; and in 
lacking the hypoconid-cnloconid cre^st on M>-a, 

Nimiokoaia differs from Perikoola in being less 

crenulated; lacking the lingual shelf basal to the 

mclaconid and cntoconid on Ihc lower molars; 

and in those features listed ioz Madakoala, 

Nimiokoaia differs t"iom Litokoala, 



Phascolarctos and Cundokoala vorkemis in: 
having a bulbous, less trenchant P^; lacking the 
bulbous cuspule at U)e anicrolingual base of the 
metaconule of M* (M* unknown for /-. 
koftt4nk<ii'nsfs) and the resultant jiockel developed 
between the prcmciaconuk' crisiii. 
postprotocrista and the lingual margin ; lacking 
the anicrohngual prolocnne crest on M', lackmg 
a mctastylid fold; lacking buccal ribs on llw pro- 
toconid. mclaconid and cntoconid; having a dis- 
continuous lingual ectolophid wherein the 
postmeiacnstid and precntt>cristid do not mc\ji: 
having a weaker entostylid: and in having less 
separation between the protoconid and paraeonid 
*>n Mi(Mi unknown for /.. kutjamurpensis). 

Nintiohtalft ^n^Ysrant^si differs Irom L 
kt4tjttfuari)e/isis in lacking the buccal extension 
connecting the paraconulc to the buccal margin. 
Nimiokoaia differs frotii L kafumkat^nsis in: hav- 
ing a poslerolingua! cuspid on P^; a greater sepa- 
ration of the anterior and posterior apices ol P? 
and a more cuspaie anterior apex of thai looih; 
lackmg die lingual and buccal ribs t'lom the main 
apices on P3; Utcking an entoconid lingual shelf 
(posto.niostylid crislid) and mclaconid lingual 
shelf; and in lacking a postcrolingual paiUvonid 
ridge on M^, 

Nimiokoaia differs irom Phascolarctos and 
Cundokoala yorkensis in: having a large 
postcrolingual cusp on P^ and strong antcritn. 
buccal and lingual crests that extend fronrt Ihc 
antcrioi apex of ihat tcniih; lacking the lingual 
cingular ridge of P\ having proportionately 
wider upper molars; lacking the pocket al the 
anterior base of the protocone on M'; having a 
prt>porlionaiely less elongate P3; having a large 
postcrohuccal cusp on P^, and strong anterior and 
buccal crests extending from the most anterior 
apex of that tooth; having a prepaHosiylld cristid 
on Ml that is noi continuous with the anterior 
cingulum; lacking the transverse connection be- 
tween the apices of the mclaconid and protoconid 
on M:: lacking the columnar Mylids of the 
metaconid and entoconid on M1.4; and in lacking 
the lingual ridge connecting the bases of the 
protoconid and hypoconid on M2 4. 

REMARKS- Nimiokoaia sp. is known only from 
a dentary tVagmcnt with M2-4 Comparisons in- 
volving the upper teeth of this genus are iherefcre 
confined to features of jV. ^^reysrar^esi sp. nov. 

ETYMOLOGY Latin nimio, excessive: refers w the 
complex molar morphology relative to utber 

phascolarctids. 



mMiOKOAUiGEK NOV.. NEW KOALA FROM RIVERSLEIGH 



^1 



Nimiokouia gn?v»ilantsi ^p. rtov. 
(Figs I -3; Tabic I) 

ETYMOLOGY. For Grcystiincs High School, winner 
Dplhc Sydney Muining Uer;ilJ/ Riverslcigh compcii- 
lion 

MATERIAL Hnlolype 0MF3f»4S2. cruniiil fnigmenl 

with pans of Ihc Icfi and right mayillHe> iuguls; and 
palaimcpurioricriprcmuxiliaandpurlurrigliifronial. 
■|'hc lefi ;iIvcoli ol l'"*. P\ M^'^ and righi P-\ M*'-. 
from Neville's Uardea Site. Sysien-i B (Archer cl al . 
l9Sy), early lo middle Miocene. Olher maierial; boid 
site cast - OMF.^r>483 partial skull wiih ihe lefi and right 
pa*ma\illae, nas;iK, palatine, pari oflhe leU and nghl 
jugals. right Ironial, pan of n^hl parietal, pan of 
basi.sphennid and basioccipiial. pari ofrighl ivmpaiiie 
lmlla.lcfi l'-^ C'. P-^ and anterior hal f of M^'righl i*. 
alveoli ol I^'-^. alveolus ul C\ P^ Iniissnig buccal 
m(irgin)iind m' (which is missing the enamel from the 
parasiylMf corner); OMF3G484. righl M-"^: 
QMF304S.^. IcM M-; QMF30486. left M- tragmeni 
with p<i^tc^rior part of met;iconule and Ihe 
nciimciacontile prci^crved; OMF304S7. light deniary 
f'ra^mcnl with alveolus of Ii, brt»ken P'^, Mi.4- 
Neville s Garden Site - QMF30512. right M^; 
QlVlF23(»26.1efi M-;QMF24232. lell M-,QMF24267. 
right M-, OMF24233. left M^: QMF2090L IcO M"^ 
and nghl M^: QMF23()27, left M"* and right M*^; 
OMF29624. right dcntary with P;. M i-i. QVIF304SH, 
lei! M I ; QMF30489, riglit M i . QMF24266. right M | : 
QMF24265. right Mr QMF20901 left M2. Camel 
Sputum Site - 0MF3{I49{). Icfl P-*. QMF3049K left 
deniary with h. alveolus o) P^ and alveolus of M1-2; 
QMF'^()492, broken righi M3; QMF24351, rigJK P3. 
Inabeyaiiee Site - QMF30493. left deniary fragment 
with 1 1, P3, M |.2, Ml missing the anlcrobuccal comer, 
M4 missing the buecal margin of the proioconid and 
ihe poslerobuccal tooth comer. Dirk's Towers Site - 
QMF245 1 6, molnr fragment ; QMF2429 1 . le 11 M^ Irag- 
jncnt; Rat Vomit Site - OMF30494, Icfi deniary iVag- 
Ttienl with P3, M1.3 (all mi.ssing the lingual margin), 
and alvciMus of M4. Lpper Site - QMF3n495, ieli P^; 
QMF3fJ496, rii!hl P-\QMF30497, rieht M\ RSO Site 
QMF3()498. left M-\ OMF30499. lalonid orM3. 

DIAGNOSIS. N. gfe\smnesi diticrs from 
Nimiokoala sp. in having: less rounded lower 
molars; less lingually sloping surfaces of the 
mciacumd and entoconid; a slightly Uirger en- 
loslylid ridge; a Uirger tieomDrphic ciispid in the 
irigonid basin ii> M2-4, and a lesser reduction of 
theialoniiHtiM4. 

DESCRIPTION. Incisors. Left and tight 1* of 
QMF30483 shtm.puinied. with convergent tips, 
with enailvcl rosirictcd 10 ihc anterior tooih lace, 
wiih postcrolingual iaccs dominated by large, 
itiiingular wear f jicets. Left 1^ small, subovatc in 



occlusal view, titperii>g;<nieroIingually. without 
enamel on the occlusal surface. Lcli 1^ stnaJI, 
pointed, peg-likc, conuctrng postcrobuccal cor- 
ner ol the left I-, wilh enumel preserved On Ihe 
btdcal fuce. 

Canincs. Left canine short, peg-like, approxi- 
nialely 4mm behind the upper incisor arcade. 

P-. LF' relatively robust, bulbous, wider posle- 
ritvly ihitn iinteriorly, with 5 majors cusps, 3 of 
which lie ^loflg a .slightly crcsccnlic longitudiniil 
crcsl wilh an additional poslerobuccal and 
posierolingual cusp. The anterior most cusp 
1.6 1 mm posterior lo the anterior looih margin. 
Prominent crcsls extending imleriurly, posted- 
oHy and buccally from its apex, anterior crcsi 
bifurcating into antcriorand postcrolingual spurs; 
spurs extending towards the bitsc of the crown. 
Medial cusp separated from Ihe anterior cusp by 
a deep crevice, connected to the posterior cusp by 
a short longitudincil crest. Apex of the medial 
cusp 2.32mm posterior m the anterior tooth mar- 
gin. Well-developed crest e.\tendif»g anicrohucc- 
ally faiw ihe medial apex, fading into the base of 
the crown. Posterior cusp 3.19mm behind artJe- 
riur looth margin, with w«li-dcvcIopcd posieritir 
crcsl extending from its apex: bifurcating ai ll>e 
t>ase of the cusp, with one arm continuing poster 
riorly and fading into ihc base of the crown; other 
cinguluin-likc ami cuning around ihe lingual 
looth margin, connecting lo the posterior crest of 
ihc well-developed posierolingual cusp. 
Posierolingual cusp iipex opposite and slightly 
poslcrior to the medial cusp, with an ao- 
icrolingual ridge fading mto ihe lingual base of 
ihe crown from the apex. Poslerobuccal cusp 
npp<xsilc, but slightly anlenor to the posterior 
apex, with *1 weH-dcveloped crest cxlending 
postcrobuccally from the apex, and fading into 
the base of the crown. An additional small 
cuspule ai the anterior ha.sc of the posicnilinguiU 
cusp. 

RP-^ similar to LP-\ but wilh less worn, medial 
cusp, smaller posierolingual cusp having wraker 
posterior crest extending from below the cusp 
apex and more distinct postcrobuccal cusp, 

Upper premolars (OMF3f)490, QMF30J95, 
QMF304%) resemble P' of the hololype cxc<:pl 
for: buccal and lingual crcsU of the anlenor ciKp, 
ihe anlerobutvul crest of iIk medial cusp and the 
postcrobuccal crcsl of the poslerobuccal cusp are 
reduced in QMF30495; undivided posterii\r crcsl 
of the posterior cusp fading into the anterior base 
of the crown in QMF30495, QMF3()49l) and 
QMF30496; small cuspule at the anlei ior base of 
the posierolingual cusp absent in QMF3()495 and 



212 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 1. Partial skull of Nimiokoala greystanesi gen. et sp. nov., QMF30483; A, ventral view; B, right lateral 
view; C, left lateral view. Bar = 1 0mm. 



QMF30496; enamel missing from most of the duced buccal crest of the anterior cusp, terminat- 
occlusal surface of larger QMF30490 with re- ing before reaching the base of the crown. 

M^ Buccal margin of left M' slightly concave* 



NMiOKOALA GEN. NOV.. NEW KOALA FROM Rl VERSl^IGH 



113 



sloping anicroiingually opposite ihc inciaconc; 
iipices* of ihc mctacH>nc and parutonc slighil> 
overhanging iheir lin^uaj bases: iinguiiJ biiNt;s of 
paraconc and metaconc more lincually sloping in 
gMF3()483 and QMF3()5 1 2 ihan in the holotypc; 
metaconc lallcrr itian paracunc; niclaconulc lallcr 
than proioconc; parasiylar curncT, posierioi and 
anterior base of metaconc, posterior base of 
paraconc, buccal bases of proioconc and 
nR'taconule, lingual bases ul" paratonule and 
neometaLonule and uiinsvci'se vaJIey ercnulaicd. 
I\iriR'onule well-dcvclopcd, crcscenlic. ciispalc, 
occupying the longitudinal valley between 
patacone and proiocone, bifurcaic anieriorly, 
main arm cxicnding antcrobuccallv and connect- 
in*; 10 Ihc antcrolingunl base ol the parastylar 
comer, with two shorter. anteri<irly directed spurs 
terminating at the base of the anieiior cin^Hilum 
and meeting the anicrior cingulum in 
QMF30483- Posteriorly paraconule eonnecting 
to a strong posterolingual paracrista and a well- 
developed crest extending postcrobuccally irom 
ihe prordcone apcv; this connection more ad- 
vanced in LM- \ Paraconule not bifurcaic poste- 
riorly m RM* of holotypc (or in QN4F3()483) but 
connecting to the (>oslcro lingual paracrista. Pos- 
teriorly bifurcate paraconule in QMF305 1 2 with- 
out connection to the poslerobuccal protocone 
ca'st. l^arpc, crcsceniic, hieusped neometaconule 
occupymg longitudinal valley between meiacone 
and metaconule in M'. NeoitJeiaci)nule in LM^'^ 
divided into two dish net cuspale V-shape-d siruc- 
lures. Neometaconule highly variable. In LM* ol' 
hololype connecteil to premelacvmiile crista by 
.short anterior spur (not in RM'). with second spur 
extending anierobuccally from the 
neometaconule and terminating in the iransversc 
valley. In QMI"305 1 2 anierobuccai arm conlmu- 
ous with wcll-dcvclopcd iransversc ridges occu- 
pying Ihc transverse valley of ihai tooth. 
Neometaconule connecting to anlcrulingual 
mciacrisia in RM' of QMF304S3. Bifurcating 
posteriorly in LM* of the hololype: Short 
posierolingually directed spur meeting a well-de- 
veloped crest extending pc»stcrobuccaity from the 
metaconule apex (these structures not meeting in 
KM' ol the holotypc); main arm of the 
neometaconule extending posterobuceally, mcei- 
in>^ both the posterolingunl mctacrista and the 
pi)sIeriorcmgulum. Neometaconule not meeting 
posterolingual mctacrista or posterior cingulum 
in QMF30483 and QMF3()5 12. Slightly crcsccn- 
Oc preparacrisla extending anierobuccally from 
Ihc paraconc apex U^ the buccal margin, bilurcal- 
ing, Willi one iirm continuing anierobuccally lo 



meet the iK»sipar«siylarcnsta; second arm exiwKl* 
ing postcrobuccally. connecting with siylar cu.sp 
B. Slightly longer poslpuracrislu bifurcating at 
the buccal margin; wiih one arm continuing 
postcrobuccally to mcci the prcmciacrisia at the 
buccal margin; with second iirm extending un- 
tctol>uccally lo meet siylar cusp C at buccal ini>ih 
margin. Paraconc buccal basin closed on M' of 
holcrype. variably closed in other specimens. 
Metaconc with siylar cusps D and li reduced 
(stronger on QMF305 12); with buccal basin shal- 
Jdw amJ opcu. Hcighi of ihe siylarcusps decrcas- 
mg progPL'SSivciy Irom the parastyle to E. Su^mg 
posterolingual paracristae. anierolingual 
mciacriMae and p<')sterolingual mctacrista in ihc 
upper molars of ihc holotypc. Anlerolinguul 
paracristae in QMF305I2. CiMI'20yoi and 
QMF23027. Proioconc apex opposite but slightJy 
posterior to the paraconc apex, slightly lingual 
relative lo the metaconule apex. Crcsceniic 
posiprotcxrrisia and prcmctaconule crista mccling 
in Ihe Iransversc valley well buccal lo the lingual 
looih margin. A series of short spurs extending 
biiccally from Ihis junction, terniinatc in the 
transverse median valley. PrcUoslyle well- tlevei- 
oped, ai the anxcrohuccal base of the protocnnc. 
extending posteriorly from Uie preprotocrisia. let- 
minating at the buccal base of the proioconc 
opposite the apex of the paractmule. Surlace 
enamel crcnulations in the transverse valley ul' 
the LM' aligned into iwo p;irallel (discontinuous) 
iransversc crests, cnnirastine with the rcticulaic 
pattern in RM' and more poslenor molars of 
hololype. Distinct transverse crests occupying 
Ihc transverse valley in QMr3tJ483 and 
OMF30498. 

M- K Similarinmost aspects. loLM' except for. 
teeth tapering more posleric^lly^ triangular buccal 
surfaces of the paraconc and meiacone sloping 
more steeply to ihc buccal margin; buccal nuirgin 
more concave, buccal miirgin of the melacoiitf 
sloping more posierolingually. paiasiyle absent; 
more reticulate cieitulations; reduced stylar 
cusps; a deep pocket ibrrncd by Ihe junction of 
the preineiaconulecrisia, postprotocrista, the 
postcrobuccal crest of ihe protocone and the 
posterolingual arm of iheparaconule; proto.siyle 
reduced, connecting to the parucunulc; 
neometaconule reduced, divided into 2 disiinci 
V-shaped structures; neometaconule meeting an- 
icrolingual mciacrisia; a vague anlerolingual 
paracrista in LM- '. 

M"* (Fig. 2B). Paraconc. paraconulc and pf<v 
tocone at similar stage of development lo M^of 
the holotypc. Meiacone and mciaconulc ex- 



214 



MEMOIRS OF THE QUEENSLAND MUSEUM 



tremely reduced to small cuspules on the 
posterohuccal and postcrolingual margins, re- 
spectively. Posterior margin crescenlic, consist- 
ing of a series of ridges extending anteriorly into 
the transverse valley. Premctacrista forming part 
of the posterior margin, curving anterobuccally 
around the tooth margin to meet the 
postparacrista at the buccal exit of the transverse 
valley. Neometaconule reduced, indistinguisha- 
ble from the enamel crenulalions. 

Right dentition of the hololype similar to the 
left except for: smaller RM'; less convex buccal 
margin; paracone and metacone sloping more 
gently to the buccal margin; deeper and less cren- 
ulated transverse valley lacking parallel trans- 
verse ridges; protocone higher than the 
metaconule; paraconule and neometaconule 
taller and less crescentic; paraconule apex closer 
to the protocone, not bifurcate anteriorly, without 
connection to the posterohuccal crest of the pro- 
tocone; neometaconule not bifurcate anteriorly, 
protostyle weaker; anlerolingual paracrisla 
vaguely developed in the right M'. 

LOWER DENTITION (Figs 2C, 3A-C). Ii . Inci- 
sor narrow, lanceolate, with enamel covering 
ventral and buccal surfaces of the anterior half, 
with line shallow longitudinal groove lineariy 
along the buccal margin terminating approxi- 
mately 1 mm posterior to anterior tip of tooth. 

P3. LP3 small, short longitudinally, of 3 aligned 
main cusps (anterior, medial and posterior), a 
large posterohuccal cusp and a small 
postcrolingual cusp; lingual surface sloping stee- 
ply to the lingual tooth margin; shallow crevice 
separating the anterior cusp from the medial cusp; 
medial apex connected to the posterii)r cusp by a 
short longitudinal crest; prominent anterolingu- 
ally directed crest and posterobuccally directed 
crest extending from the anterior apex and fading 
into the base of the crown; prominent crest con- 
necting posterior cusp to posterior cingulum; 
apex of the posterohuccal cusp opposite and 
slightly posterior to the posterior cusp. Short an- 
terobuccally and posterolingually directed crests 
extending from the apex of the posterohuccal 
cusp. 

Lower molars distorted, wuh metaconid and 
entoconid anterioriy displaced relative to the pro- 
toconid and hypoconid, respectively. 

Ml. Subrectangular, anteriorly attenuated; 
hypoconid largest cusp on the lalonid; protoconid 
largest cusp on the trigonid; well-developed pro- 
losiylid at the buccal margin, opposite and 
slightly posterior to the protoconid apex. Pro- 



tostylid and hypoconid pyramidal in occlusal out- 
line in QMF30493 but more rounded in 
QMF30494. Preprotocristid extending an- 
terolingually from the protoconid apex to a rela- 
tively small paraconid at the anterolingual corner 
(in (3MF3()488 paraconid connected to the pre- 
metacristid by a short longitudinal crest). Pre- 
metacristid poorly developed. Linear 
postprotocristid extending posteriorly into the 
central bcisin. meeting the crenulated crisiid ob- 
liqua slightly lingual to the longitudinal tooth 
axis; postmetacristid extending posterolingually 
towards the weakly developed mciasiylid at the 
lingual tooth margin, with a well-developed 
posterolingually directed ridge branching off and 
extending into the transverse valley separating 
the talonid and trigonid. Crescentic pre- 
protostylid crisiid curving anterolingually along 
the anterior tooth margin, connecting to the an- 
terobuccal base ofthe protoconid. ihis connection 
slightly more lingual in QMF3()48H and slightly 
more buccal in QMF30489. Postproioslylid 
cristid extending towards the lingual margin, ter- 
minating at the anterior base of the hypoconid; 
less developed in QMF30488 and QMF3()494, 
fading into the posterior base of the protostylid. 
Well-developed posterolingually developed crest 
extending from the protostylid apex, terminating 
at the posterior end of the longitudinal crevice 
separating the protostylid and protoconid, more 
crescentic and with an additional short lingual 
ridge extending from the protostylid apex in 
QMF30488. Preentocrislid extending an- 
terobuccally (as opposed to anterolingually in 
other phascolarclids) into the transverse valley 
separating talonid from trigonid. A second crest 
extending anterobuccally from the entoconid 
apex, running parallel to the prcenlocristid. An 
additional buccally directed crest extending from 
the entoconid apex into the valley separating the 
bases of the entoconid and entostylid ridge. 
Postentocristid extending postcrioriy along the 
lingual margin to a pooriy developed entostylid 
at the postcrolingual corner o( the tooth. Large 
cuspatc ridge in the longitudinal valley between 
the entoconid and hypoconid analogous to, al- 
though more strongly developed than, the en- 
tostylid ridge in Litokoala and Phascolarctos. 
Entostylid ridge bifurcating anterioriy; two an- 
terobuccally directed ridges extending from its 
apex, meeting the crisiid obliqua. Bifurcating 
postcrioriy: two posterolingually directed ridges 
terminating at the base ofthe posterior cingulum. 
Entostylid not bifurcate in QMF3{)494. 
M2-3. M: and M3 similar to Mi except for: 



NIMIOKOAIA GEN. NOV., NEW KOALA FROM RIVERSLEIGH 



215 




F\G. 2. Nimiokoalagreysmnesi gen. asp. now. A-A\QMF30482,hololype, occlusal slereopairofroslral region. 
B-B\ QMF23027, right M"^, occlusal stereopair. C-C\ Q1VIF30487, right deniary fragment, occlusal stereopair. 
Bar = 5mm. 



protostylid absent; paraconid absent; preprolo- 
cristid continuous with anterior cingulum; 
postprotocristid linear, extending posterolingu- 
ally towards its junction with the crislid obliqua; 
premetacrislid me^re prominent, continuous with 
the anterior cmgulum; entoslylid reduced result- 
ing in a more rounded postcrolingual tooth cor- 
ner; eniostylid ridge reduced, not bifurcate; 
cntoconid reduced in M^; hypoconid reduced; 
preentocristid and posterobuccal metacristid 
joining, closing the transverse valley well buccal 



to the lingual tooth margin. A neomorphic cuspid 
in the longitudinal valley between the metaconid 
and protoconid in M2-4, with a well-develc^ped 
posterolingually directed ridge and a weaker an- 
terobuccally directed ridge extending from its 
apex, more strongly developed and more sepa- 
rated from the metaconid in M2-4 of QMF30487 
thanofQMF30493. 

M4. M4 similar to M2-? except for; entoslylid 
ridge further reduced, connecting posteriorly to 
the posterior cingulum; neomorphic cuspid re- 



216 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 3. Nimiokoala greysianesi gen. el sp. nov. A-C, QMF30493. A-A\ occlusal slereopair of left dentary. B, 
buccal view. C, lingual view. Bar= 10mm. 



duced; postprotocristid and cristid obliqua not (relative to the hypoconid and protoconid,respec- 

meeting end to end but parallel each other (as in lively) (as in L kanunkaensis and some 

L kanunkaensis and some P. cinereus): talonid pseudocheirids). 

severely reduced resulting in a greater anterior 

displacement of the cntoconid and metaconid REMARKS. QMF30482 (Fig. 2A) was chosen as 



NIM/OKOALA GEN. NOV., NEW KOALA FROM RIVERSLEIGH 



2!7 



TABLE 1. Denlilion measuremenls (mm) for Nimiokoala greystanesi sp. nov. and Nimiokoala sp. (last line 
SAMP19952 only). L= length; W=width; AW, anterior width; PW, posterior width; *=estimate. 



A. UPPER CHEEK DENTITION 1 


Spec. 


Side 


P3 


Ml 


M2 


M3 


M4 1 


L 


W 


L 


AW 


PW 


L 


AW 


PW 


L 


AW 


PW 


L 


AW 


PW 1 


QMF30482 
Holotype 


L 


4..^0 


3.98 


6.45 


5.94 


5.66 


6.05 


6.18 


5.52 


5.70 


5.67 


4.63 








R 


4.48 


3.51 


5.99 


5.74 


5.35 


5.92 


5.90 


5.54 














QMF30483 


L 


4.51 


3.95 


- 


5.39 






















R 


4.63 


3.83 


6.30 


5.47 


5.68 




















QMF30484 


R 












5.53* 


5.63 


5.22 


5.16 


5.15 


4.46 


4.60 


4.48 


2.12 


OMF30490 


L 


4.87 


4.15 


























QMF30495 


L 


4.52 


3.68 


























QMF30496 


L 


4.46 


3.53 


























QMF305I2 


L 






6.81 


5.99 


5.93 




















QMF24267 


R 












6.12 


6.14 


5.41 














QMF24232 


L 












6.16 


6.01 


5.42 














QMF30485 


L 












- 


- 


5.23 














QMF30498 


L~^ 












5.18 


5.27 


4.85 














QMF23026 


R 












5.67 


5.88 


5.52 














QMF30497 


R 


















5.28 


5.28 


4.72 








QMF24233 


L 
























4.12* 3.96 


1.91 


QMF23027 


L 
























4.47 4.24 


2.65 


R 
























4.67 


4.32 


2.62 


QMF20901 


L 
























4.68 


4.69 


2.81 


R 
























4.29 


4.16 


2.30 


B. LOWER CHEEK DENTITION 


OMF30487 


R 


4.10 


3.00 


5.70 


3.15 


3.69 


5.77 


3.28 


3.62 


5.56 


3.19 


3.28 


5.07 


3.16 


3.00 


QMF30493 


L 


3.35 


2.44 


5.95 


3.54 


3.40 


5.85 


3.40 


3.43 


5.50 


_ 


3.04 


5.10 




. 


QMF30494 


L 


3.60 


2.46 


5.37 


- 


3.41 


5.86 


3.22 


3.28 


5.57 


3.17 


3.05 








QMF29624 


L 


3.86 


2.54 


5.91 


3.07 


3.48 


5.75 


3.26 


3.34 


5.53 


3.30 


3.12 








QMF2435I 


R 


3.21 


2.25 


























QMF24266 


R 






6.22 


3.29 


3.66 




















QMF30488 


R 






6.19 


3.42 


3.66 




















QMF30489 


R 






5.96 


3.45 


4.04 




















QMF20903 


L 












6.31 


3.55 


3.92 














QMF24265 


R 












5.84 


3.27 


3.30 














SAMP19952 


L 












5.04 


2.77 3.00 


5.18 


2.95 


2.88 


4.32 


2.53 


2.19 



holotype because it contains most of the upper 
dentition well-preserved and relatively unworn. 
The holotype is thought to represent a juvenile 
because of the shorter premaxilla in comparison 
with the adult skull, QMF30483. 

Nimiokoala sp. 

(Fig. 4, Table 1) 

MATERIAL. SAMP19952 left dentary with alveoli of 
P3 and Mi, M2-4 intact from late Oligocene to middle 



Miocene South Prospect B locality, Lake Namba, 
Frome Downs Station, South Australia. 

REMARKS. Enamel is missing from the lingual 
margins of the melaconids and entoconids and the 
buccal margins of the protoconids and 
hypoconids. Features of the M2 are difficult to 
discern because of extreme wear in that tooth. 
Only points of difference from N. greystanesi are 
noted here. 

Lower molars are proportionately smaller (by 
approximately 15%) and more rounded, espc- 



218 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 4. Nuniokoala sp.. SAMP 19952, from South Prospect B Locality. Lake Namba, South Australia. Occlusal 
slcreopair of right (Jentary. Bar= lOintn. 



daily the talonid. The neomorphic cuspid be- 
tween ihe mctaconid and proioconid in M:.^ is 
weaker, as is the entosiylid ridge. The lingual 
surfaces of the proioconid and nielaconid slope 
more linguaily. The lalonid o'^ M4 is further re- 
duced. 

This laxon is left in open nomenclature in the 
absence of the upper dentition. The features men- 
tioned above as differences between the central 
Australian and Riversleigh materials vary signif- 
icantly between individuals of /V. greysianesi and 
may not be reliable species indicators. 

Litokoala Slirton, 1967 

TYPE SPEClES.Litokoala kutjamarpensis Stirton, 
1967. 

OTHER SPECIES. Litokoala kanunkaensis Springer, 
1987. 



DIAGNOSIS. Litokoala differs from other 
phascolarctids in having a posterobuccal crest 
extending from the apex of the mctaconid on 
M2-4 (although this crest is reduced on M4); a 
well-developed poslerolingually directed cresi 
from die proioconid apex of M4. L/rt>Acw/n differs 
from other phascolarctids except PhascoUirctos 
and Cundokoala yurkensis in: having a neomor- 
phic cuspuie at ihe anterior base o'i ihe mela- 
conuleofM';ananlerolingualprotoconecreston 
M' and consequently a much squarer an- 
terolingual margin of that tooth; a metastylid fold 
wherein ihe poslmeiastylid cristid is continuous 
with the preentocrislid; an anteriorly bifurcate 
preentocristid on the lower molars; and internal 
ribs on the mctaconid and hypoconid of Mi. 

Litokoala differs from all other phascolarctids 
except Niniiokoala in having: a well-developed 
crescenlic paraconule and neometaconule on 
upper molars; an anteriorly displaced entoconid 



NIMIOKOALA GEN. NOV., NEW KOALA FROM RIVERSLEIGH 



219 




FIG. 5. Litokoala kanunkaensis, Henk's Hollow Site, Riverslcigh Station. A-A', QMF30502, RM-^ occlusal 
stereopair. B-D, QMF30500, RP3. B-B\ occlusal siereopair. C, buccal view. D, lingual view. Bar = 5mm. 



(relative to the hypoconid) on M4; an anteriorly 
displaced meiaconid (relative to the proloconid) 
on M4; and a parallel arrangement of the 
postprotocristid and cristid obliqua on M4 
wherein these cristids do not meet end to end. 

Litokoala differs from Koobor in: being 
smaller; having less rounded bases of the 
paracone, metacone, protocone and metaconule; 
having less elongate molar proportions; reduced 
stylar cusps; a paraconule on M-^; and a 
neometaconule on M'-^. 

Litokoala differs from Madakoala and Peri- 
koala in: being smaller; higher crowned; more 
selenodont; having a larger neometaconule; a 
more crescentic, less linear paraconule; an en- 
tostyHd ridge in the lower molars; a more lingual 
junction of the cristid obliqua and 
postprotocristid; a larger developed protostylid 
and a more lingual protoconid on Mi; a less 
strongly developed, more buccally positioned 
paraconid on Mi; and a more strongly developed 
entostylid. 

Litokoala differs from Perikoala in: lacking the 
well-developed stylar border of the paracone and 
metacone; lacking the additional stylar cusp an- 
terior to stylar cusp E; having a weaker entoconid 
lingual shelf; and a weaker premetacristid. 

Litokoala kutjatnarpensis differs from 
Madakoala and Perikoala in having a buccal 
extension of the paraconule which connects to the 
buccal margin of M'. Litokoala kanunkaensis 
differs from Madakoala and Perikoala in: lack- 
ing the M2 protostylid ridge; having a less elon- 
gate P3; a more prominent posterobuccal cusp; 



and having 3 apices on the longitudinal crest (4 
in Madakoala devisi: 5 in M. wellsi) on P3. 

Litokoala differs from Nimiokoala in: having 
less steeply (more buccally) sloping buccal sur- 
faces of the paracone and metacone; lacking the 
division of the neometaconule in the more poste- 
rior molars; lacking the well-developed 
posterobuccal crest from the apex of the 
metaconule; having a weaker posterobuccal crest 
from the protocone apex; lacking the 
posterolingual ridge off the apex of the pro- 
tostylid on Mi; having a less cuspate, more pos- 
teriorly positioned entostylid ridge; having an 
anterolingually (as opposed to anterobuccally) 
directed preentocristid; a more posterolingually 
directed postprotostylid cristid; a lingual en- 
toconid shelf; and a more strongly developed 
entostylid. Litokoala kanunkaensis differs from 
species of Nimiokoala in: lacking the 
posterolingual cusp on P3; in having a weaker 
anterior cusp on P3; buccal and lingual ribs which 
extend from the main apices of the longitudinal 
crest in the P3; and in lacking the neomorphic 
cuspid between the metaconid and protoconid of 

M2-4. 

Litokoala differs from Phascolarctos and 
Cimdokoala yorkensis in: being smaller; having 
well-developed anterolingual metacristae; hav- 
ing proportionately less elongate lower molars; a 
less pronounced entoconid basal shelf; and in 
lacking the ridge connecting the buccal bases of 
the protoconid and hypoconid of M2-4. 



220 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Litokoala kanunkaensis Springer, 1987 
(Figs 5-6, Table 2) 

MATERIAL. Holoiype SAMP32397 (=UCR21926) a 
right MiiVom latest Oligocene UCR Locality RV-8453 
Kanunka North Local Fauna, Etadunna Formation, 
west side of Lake Kanunka, South Australia. Other 
material: from KanunkaNorth Site: UCR2I945, aright 
M4: UCR2 1 980, a melacone of a right M^ UCR2 1 979. 
a metacone of LM'; late Oligocene. From Henk's 
Hollow Local Fauna: QMF30500, right P3; 
QMF 13079, right dentary fragment with posterior half 
of P-i, M|.7; QMF30502, right M^; from Gag Site: 
QMF3050 i , right Mi ; from Gotham Site: QMF30503, 
M'* fragment containing paraconc and buccal half of 
protocone. From JC9 Site: QMF20809, right M3; Sys- 
tem C, Riversleigh, middle Miocene (Archer et al.. 
1989; 1991). 

DIAGNOSIS. Litokoala kcmiinkaensis differs 
from L kutjamarpensis in: having a short longi- 
tudinal spur connecting the paraconule to the 
anterior cingulum; lacking the bulbous cuspule al 
the anlcrolingual base of the metaconule; lacking 
the prolostyle (the latter two features may be 
result of differences along the tooth row rather 
than interspecific differences); lacking ihe dis- 
tinct postcrolingual and anlerolingual paracrista; 
having a strong anterobuccal crest from the 
metaconule apex. 

COMPARISON WITH THE HOLOTYPE. Prior 
to this study L kanimkaensis was known from 
two isolated lower molars, an M2 and M4. Refer- 
ral of the Riversleigh material to L kannnkaensis 
is based on the similarity of the M2 of QMF 1 3079 
to the holoiype. Some small differences are: M2 
of QMF 1 3079 is slightly larger: poslcrobuccal 
meiaconid strut is poorly defined and bifurcates 
in the trigonid longitudinal valley, one spur fad- 

TABLE 2. Measurements (mm) for right lower denti- 
tion of Litokoala kanimkaensis from Rivcrsleigh's 
System C deposits. Abbreviations as for Table 1 . 





P^ 


M, 


M-) 


M^ 1 


QMF 

No. 


L 


w 


L 


AW 


PW 


L 


AW 


PW 


L 


AW 


PW 


30500 


4.12 


3.10 




















30501 






4.72 


2.63 


3.04 














13079 


- 


2.62 


4.89 


3.14 


3.58 


5.15 


3.43 


3.57 








20809 


















5.48 


3.39 


3.40 



ing buccal ly into the protoconid base and the 
other spur continuing posteriorly then turning 
sharply lingually before becoming part of the 
molar crenulalion pattern; the entoslyiid ridge is 
less cuspate and arises from the postcrobuccal 
base of the enioconid (unlike the holotype in 
which it arises from the postenlocristid); and the 
transverse median valley, the posterior base of the 
protoconid, the cristid obliqua and the transverse 
valley separating the bases of the protoconid and 
hypoconid are more crenulated than in the holo- 
iype. 

QMF 13079 M2 is significantly more worn than 
the holotype which may account for the poorer 
crest definition. Analysis of intraspecific varia- 
tion in Phascolarctos cinereus and Nimiokoala 
greystanesi indicate that these differences do noi 
warrant specific distinction. Most variable fea- 
tures in ihe modern species include: molar size; 
pattern and degree of molar crenulations; crislid 
obliqua; entoslyiid ridge. Although the entoslyiid 
ridge may be phylogenelically significant in 
phascolarctids, it is highly variable. It may be a 
well-developed cuspate structure, relatively in- 
distinguishable as a series of discontinuous cren- 
ulale ridges, completely isolated at the base of the 
enioconid or variably connected to the 
posientocrisiid, the base of the enioconid or the 
posterior cingulum. Hence differences between 
the holotype and QMFI3079 are within intraspe- 
cific variation. 

DESCRIPTION. P3 (Fig. 5B-D). Short, semi-rec- 
tangular, of 4 major cusps anteriorly, medially, 
posteriorly and posterobuccally on the crown. 
Anterior, medial and posterior cusps on a longi- 
tudinal cresi extending the length of the crown. 
Anterior cusp tallest, 1/3 of way along longitudi- 
nal crest, above the anterior root of P3. Anterior, 
buccal and lingual crests extending from its apex; 
anterior crest fading into the anlerolingual base 
of the crown; lingual crest fading down the lin- 
gual tooth margin, curving posteroiingually at its 
tip; buccal crest short, fading into the buccal tooth 
margin. Anterobuccal. anlerolingual and 
posterolingual crests extending from the poste- 
rior cusp apex; anterobuccal crest poorly devel- 
oped, extending into the valley between the 
posterior and posterobuccal cusps, meeting a 
short lingual ridge from the posterobuccal cusp 
apex; anlerolingual crest belter developed, fading 
down the lingual margin; posterolingual crest 
curving anlerolingually along the base of the 
crown, becoming cingulum-like, fading into the 
base of the posterior cusp. Posterobuccal cusp 



NIMIOKOALA GEN. NOV., NEW KOALA FROM RIVERSLEIGH 



221 




FIG. 6. Litokoala kanunkaensis. A-A\ QMF13079, right dentary fragment with posterior half of P3, Mi. 2, 
occlusal stercopair. B-B', QMF30501, RM|, occlusal stereopair. C-C', QMF20809, LM3 occlusal stereopair. 
Bar = 5mm. 



opposite but slightly posterior to the posterior 
cusp, with a short posterolingual ridge extending 
from its apex and terminating at the posterior end 
of the valley between the posterior and 
posterobuccal cusps, with short, anterobuccal 
crest extending from its apex and fading towards 
the base of the crown. 

Ml (Fig. 6B). Scmi-rcctangular, tapering 
slightly anteriorly. Protoconid most worn of the 
major cusps, with well-developed protostylid op- 
posite but slightly posterior to apex. Short, linear 
cristid extending posteriorly from the apex of the 
protostylid into the transverse valley, bifurcating 
into posterolingual and posterobuccal arms, both 
arms terminating in the transverse valley between 
the talonid and irigonid. (In QMF13079 
postprolostylid cristid more crenulaled, bifurca- 
tion less distinct and more a part of the crenula- 
tion pattern). Anterolingual cristid from 
protostylid apex crescentic, well defined, contin- 
uous with the anterior cingulum extending to the 
anterolingual corner of the tooth. Lingual rib 
from the protostylid apex short, fading into its 
base. Crescentic, cingulum-like ridge extending 
posterobuccally from the posterobuccal base of 
the protostylid, fading into the anterior base of the 



hypoconid. Preprolocristid extending an- 
lerolingually to the paraconid, a moderately de- 
veloped, slightly cuspate structure at the 
anterolingual tooth corner, with a posterolingu- 
ally directed ridge fading down the lingual mar- 
gin towards the base of the crown. Anterobuccal 
crest continuous with the anterior cingulum/pre- 
protosiylid cristid. Postprotocristid extending 
posteriorly meeting cristid obliqua slightly lin- 
gual to the longitudinal axis of the tooth. 
Postmetacristid extending posierolingually from 
the apex of the mctaconid to the lingual margin, 
swelling at this point to form the metastylid. 
Posterolingual ridge from the metastylid short, 
fading down the lingual tooth margin, belter de- 
veloped in QMFI3079. Postmetastylid cristid a 
short, buccal ridge, continuous with the pre- 
entocristid, resulting in the metastylid fold. En- 
toconid sub-pyramidal. Preentocristid extending 
anteriorly towards the transverse axis, curving 
lingually, bifurcating with one spur extending 
lingually to meet the postmetastylid cristid and 
other extending anterobuccally into the trans- 
verse valley to lingual base of the junction of the 
postprotocristid and cristid obliqua. QMF13079 
(Fig, 6A-A') with a short anterolingual spur ex- 



in 



MUMOIRS OF THE QUEENSLAND MUSEUM 



lending fmm the junction of the poMproiocrislid 
and crisiiil t>hliqiia lo the anienibiR-fal spiiroFihe 
prccnlocrislid. A wcll-dcvcIopcd antcrohuccal 
ridge extending From the cnlocunid apcK, lernii- 
niiling at the anlerohiiccal base of the cntucunid, 
just Hngual to the anterior arm of the enlostylid 
ridge. Fosleniueristid curving poslen»hngualiy 
from the cnloconid apex to the postcrolinguai 
tool htorner,. swelling at ihispt>inl to form a small 
enlostylid. A short anierolingual ndgc lading 
down the lingual tooth margin from thcentostylid 
apex, better developed in QMri3()7*>. A short 
poslerobuceal ridge extending from the en- 
(osiylid apoK, continuous with the posteiior cin- 
gulum. Wcll-dcvclopcd cuspatc enlostylid ndge 
at the poslcrobuceal base of the proloeontd. foul- 
ing ameriorly along the longitudinal looih axis, 
with a shorter posterior spur terminating before 
meeting the posterior cingulunr QMF13()7y with 
enlostylid ridge extending huceally from the 
postentcKTistid. similar in this respect to the ho- 
lotype. Hypoeonid largest of the main cusps. 
Poslliypocrislid extending posterolingually along 
ll\c posterior tooth margin, becoming continuous 
with the posleriorcingulum.Crisiidobliqua long, 
well-developed, crenulatcd, relatively linear, ex- 
lending anterolingually from the hypoeonid apex, 
meeting the posii>nMoci*isiid on the nansversc 
valley shghlly lingual to the longitudinal axis. 
Talonid basin, transverse valley between the 
lalonid and trigonid and buccal base of the en- 
loconid and paUoconid crenulatcd. Lingual base 
of the hypoeonid and protosiylid lightly crenu- 
laied. Protostylid occtipying ihe rintei'obuceal 
corner ol the tooth, lying well buccal to the lon- 
gitudinal axis Short lingual ribscKiending fron\ 
apices of the entoconid, hypoeonid, protoconid 
and jiroioslyHd. Short buccal ribs off the pro- 
toconid antl mctaconid apices, less prominent in 
the holoiype. 

Ml (Fig 6C-C' ) IS sinnlar lo the holotype ex- 
cept for: hcmg slightly larger, having buccal 
bases of the protoconid and hypoeonid more 
ir>unded. the lingual shelf of the ntetaconid better 
dt-'veloptrtl, the crislid obliqua and 
postprotocrislid meeting at a more lingual posi- 
tii^. the anierobuccal entoconid ridge and the 
poslcrobuceal mctaconid ridge pooriy developed 
(possibly due to wear) and the enlostylid ridge 
extending buccally from iheposlentostylid cristid 
ralher than Irorii the postcntocnstid. 

M' (Fig. 5A A) Length :S I6mmi anterior 
width 5.22mm: posterior width 4.2Smtii. Sub- 
t|uadra!e, tapering slightly posteriorly, with shal- 
low surface enamel crenululions in the transverse 



valley and posterior base of paracone. Buccal 
bases of the protocone and metaconc large, 
rounded, with lingual bases of all major cusps 
sloping gently lingually. Triangular buccal sur- 
t jc<r of paracone redticed relative to the metaconc 
(less marked in QMF30503). Buccal margin of 
theparacone sloping anierolingually; buccal mar- 
ginof the metaconc slopingslightlyposterolingu- 
ally in occlusal view. Buccal basins of the 
paracone and meiacone shallow and open. Buccal 
ba.sin of the paracone deeper ihan that of die 
metaconc. Slylar cusps B andC wcll-dcvclopcd 
on the paracone. Stylar cusps on ihe mciaci>ne 
l>x>rly developed, stylar decrease proga^ssively 
from B to E. In QMF303U3 stylar cusp C more 
strongly developed, almost closir»g the pafacone 
buccal basin. Prcparacrisia relatively short cresl, 
extending anierobuccally to the anterobuccal 
tooth comer, bifurcating, with one arm curving 
buccally and becoming continuous with the ante- 
rior cmgulum. the other short ridge extending 
postcrobuccally, fading along the paracone buc- 
cal OKirgin to u slight swelling representing .stylar 
cusp B Longer, relatively linear postparacrista 
extending postcrobuccally to the buccal margin, 
meeting premctacrista, closing Ihe buccal exit of 
the transverse valley. Stylar cusp O a slight swell- 
ing at the buccal tip of the premctacrista. 
Posimelacrisla linear, extending postcrobuccally 
from Ihe metaconc apex lo the poslcrobuceal 
tooth corner, meeting the posterior cingulum. 
Piwaconule at the anierolingual base of the 
piiracone. slightly cuspaie ai this |>oint, bifurcaF 
ing antcrioriy, with main linear arm extending 
anierobuccally. terminating at tlie anierolingual 
base of the paracone before tneeiing the anterior 
cingulum. with a minor ami connecting the ante- 
lior cingulum Linear posterior arm of the 
paraconulc extending posteriorly along the longi- 
tudinal valley between the paracone and pro- 
tocone. bilurcating at a point jusi artieriiir to the 
transverse median valley: A shon poslcrobuceal 
arm meeting the posterolingual paracrisia; sec- 
ttml arm coniinumg posii^riorly into the trans 
verse median valley. A cuspate, anteriorly 
bifurcate ncnmctaconule at the anierolingual base 
oi the metaconc. with ihe main crenulatcd arm 
extending antetobuccally, meeting Ihe an- 
ierolingual melacrisia at ihc base of the metaconc. 
A shorter arm fading lingually from the 
neomelaconule apex mto the longitudinal valley 
at the buccal base of the bypoconc Postenorly, 
the linear arm of the oeometaconule fading into 
Ihe longitudinal valley bel ween the metaconc and 
melaa>nulc. Buccal surfaces of the protocone and 



NIMIOKOALA GEN. NOV., NEW KOALA FROM RIVERSLEiGH 



223 





CS 




Cl 


^ 


-^ 


o 


C2 


-^ 


"^ 


o 


ta 


^ 


S 



o 






o 
o 






[Llo->i 

[3]o->l 

0->l 



10 



13 



16 



-> 1 
(1->1 
0->l 



[i]o->i 
-> 1 
-> 1 

l->2 



11 



12 



14 



17 



l->2 
0->l 
-> 1 



□ o->i 
[T5I -> 1 



1] 



0->l 



[l]o->i 
[?]o->i 

0->l 



12 



FIG. 7. Phylogeny of the Phascolarctidae based on 
cladistic analysis of dental characters (Table 3). 
Apomorphies are represented by boxed numbers. Ar- 
rows indicate character state transformations. = 
plesiomorphic character state; 1 = derived character 
slate; 2 = most derived character state. 

melaconule sloping gently buccally. Pre- 
prolocrista long, crescentic crest, extending an- 
terobuccally, curving more buccally inlo the 
longitudinal valley, meeting the anterior spur off 
the paraconule, becoming continuous with the 



anterior cingulum. The slightly more crescentic 
postprotocrisla extending posterobuccally to- 
ward the transverse valley, turning sharply bucc- 
ally, meeting the short, slightly crescentic 
prehypocrista at the buccal end of the transverse 
valley between the melaconule and protocone. 
Prehypocrista bifurcating just posterior to the 
transverse valley, with one arm continuing an- 
terobuccally to the postprotocrisla, with the other 
crenulaled arm extending buccally, then an- 
terobuccally into the transverse median valley. 
Junction of the postprotocrisla and prehypocrista 
effectively sealing off the lingual exit of the trans- 
verse valley well-in from the lingual margin. Two 
short ridges at the posterolingual base of the 
protocone and the anterolingual base of the 
melaconule. Posterolingual base of the protocone 
slightly crenulaled and cingulum-like. The long, 
slightly crescentic posthypocrista extending 
posterobuccally, becoming continuous with the 
posterior cingulum. A relatively well-developed 
anterobuccal ridge from the melaconule apex, 
fading towards its base. Posterobuccally directed 
ridge from the protocone apex fading into the 
poslerobuccal base of the protocone. 
Posterolingual paracrista bifurcating at its base, 
one arm meeting the anterobuccal arm of the 
paraconule and continuing into the transverse 
median valley, with the other extending 
posterobuccally inlo the transverse median val- 
ley. Anterolingual metacrista bifurcating at the 
anterolingual base of the metacone, one arm con- 
tinuing anierolingually to meet the anterobuccal 
arm of the neomelaconule, the other extending 
anlerobuccally inlo the transverse valley between 
the paracone and metacone. 

PHYLOGENETIC ANALYSIS 

Seventeen dental characters which provide 
phylogenetically useful data on phascolarctid re- 
lationships are analysed (Tables 3, 4). Character 
polarities were determined using selenodont 
ilariids and subselenodont wynyardiids as out- 
groups. 

CHARACTER ANALYSIS 

Thirty potentially useful dental characters were 
reduced to 17 following analysis of variation in 
the living species. The Pleistocene Cundokoala 
yorkensis Pledge, 1 992 is not significantly differ- 
ent in dental morphology from Phascolarctos: 
generic distinction is not warranted. It is here 
regarded as a separate species of Phascolarctos. 



224 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 3. A list of character state polarities (PLES= 

plcsiomorphic state; APO= apomorphic state) for the 
Phascolarcticlae using ilariids and wynyardiids 
(Marsupialia: Vombatimorphia) as ouigroups.A=ab- 
seni; P=presenl; LI=linear; C=crcsccniic; LA=large; 
R=reduced; W=vveak; S=smaII; M=modcrate; 
B=bLiccal; LIN=lingual third of irigonid 



CHARACTER 


PLES 


APO 


KB laded premolar 


A 


P 


2Trenchanl premolar 


A 


P 


3. Posterobuccai cusp on P 


P 


A 


4. Structure of paraconule 


LI 


C 


3. Development of neomeiaconule 


A 


P 


6. Proiosiyle de\e!opmeni 


A 


P 


7. Slylar cusp development 


LA 


R 


8. Postcrolingual paracnstae 


A/W 


P 


9, Neomorphic cuspule ai base of 
metaconule 


A 


P 


10. Anlerolingual prolocone crest 


A 


P 


1 1 . Parastyle development on M 


S 


M/L 


12. Protosiylid development on Mi 


A 


P 


13. Mctaslylidloid 


A 


P 


14. Enloslylid ridge 


A 


P 


15. Position of protoconid on M| 


B 


LIN 


16. Buccal ribs on lower molars 


A 


P 


17. Anteriorly displaced cnloconid relative 
to hypoconid and anteriorly displaced 
inelaconid relative to protoconid in M,4 


A 


P 



L Premolar cusp pattern: P3 in most 
phascolarclids (e.g. Madakoala, Perikoala, 
Nimiokoaku Litokoala, Phascolarctos) has a ten- 
dency to be hladed. as opposed to bicuspcd and 
weakly bladed in Koohor. Woodburne el al. 
(1987a) regard the bicusped P3 as plcsiomorphic 
among phascolarctids but its occurrence in other 
vombaliform groups (e.g. ilariids and 
wynyardiids) suggests that it is apomorphic in 
phascolarclids. 

2. Trenchant P-^- Phascolarctos has a trenchant 



F^ in contrast to a bulbous P^ in Koohor, 
Madakoala, Perikoala and Nimiokoala. Ilariids 
and wynyardiids have a bulbous, non-trenchant 
P^ suggesting the trenchant P^ in Phascolarctos 
is apomorphic. 

3. Posterobuccai cusp of P-^'- The small 
posterobuccai cusp on P-^ in Koobor, Madakoala, 
Perikoala and Nimiokoala is lacking in 
Phascolarctos. It is absent in wynyardiids and 
variable in ilariids. Its occurrence in most 
phascolarctids and ilariids suggests that it is 
plcsiomorphic among phascolarctids. 

4. Crescentic paraconule: The paraconule var- 
ies among phascolarclids from a small, linear 
structure in Madakoala and Perikoala to moder- 
ately developed in Phascolarctos (although not 
uniform), to a large crescentic paraconule in 
Litokoala and Nimiokoala occupying the longitu- 
dinal valley between the paracone and protocone. 
It is a moderate swelling at the anterolingual base 
of the paracone in M' of Koohor, however, it is 
suppressed on M--^ as in ilariids. A paraconule is 
ab.seni in wynyardiids but a small swelling at the 
anterolingual base of the paracone in ilariids. 
Hence, absence of a crescentic paraconule is re- 
garded as plcsiomorphic among phascolarclids. 

5. Neometaconide: Absent in Koobor , weakly 
developed (or absent) in Madakoala and Peri- 
koala, variable in Phascolarctos, large and cres- 
centic in Litokoala and Nimiokoala (although 
reduced in the more posterior molars) and double 
cusped in the latter. Its absence is regarded as 
plcsiomorphic in koalas because of its absence in 
ilariids and wynyardiids. 

6. Protostyle: Present in Phascolarctos, 
Nimiokoala and Litokoala although it generally 
diminishes in size from M' to M"*. It is absent in 
Madakoala, Perikoala, Koobor, ilariids and 
wynyardiids which suggests its presence is 
apomorphic. 

7. Stylar cusp development: Large slylar cusps 
arc thought to be primitive among 



TABLE 4. Character slate distribution within Phascolarctidac. Abbreviations: 0, plcsiomorphic stale; 
apomorphic siate; 2, more derived apomorphic stale; ?, signifies missing data. 







-) 


3 


4 


.^ 


6 


7 


8 


9 


to 


11 


12 


13 


14 


15 


16 


17 


Phascolarctos 




i 


1 







1 


I 


1 


1 


1 


1 


2 


1 


1 


1 


1 





Litokoala 




1 


1 


1 




1 


1 


1 


1 


1 


1 

- 


-) 


1 


1 


1 


1 


1 


Nimtokoalo 










1 




1 


1 


1 








'y 


o 





1 


I 





1 


Perikoala 


















1 














1 








1 








Madakoala 

































1 

















Koobor 


\ 


























1 


— ^ — 


9 

■ 


'J 

■ 


■> 


9 


■) 



NIMIOKOALA GEN. NOV.. NEW KOALA FROM RIVERSLEIGH 



225 



diproiodonlians (Rich & Archer, ! 979), retlected 
in the enlarged stylar cusps of ilariids and 
wynyardiids. Large stylar cusps occur in Koobor 
and Madakoala. Ahhough Pehkoala has reduced 
stylar cusps, the stylar region is represented by a 
longitudinal crest that appears lo subsume rela- 
tively large stylar cusps. Those oi Nimiokoaku 
Litokoalo and Phascolarctos (excluding the 
paraslyle of M') are suppres.sed relative to all 
other phascolarctids. Because of the large stylar 
cusps in ilariids and wynyardiids, reduction in 
some phascolarctids is regarded as apomorphic. 

8. Posterolinguolparacristae: Well-developed 
and aligned with the preparacristae in 
Nimiokoala, Utokoala and Phascolarctos but ab- 
sent (or weakly expressed) in Koobor, 
Madakoala, Perikoala, ilariids and wynyardiids. 
Absence is regarded as plesiomorphic. 

9. Cuspule at the anterolingual metaconular 
base of M^: A bulbous cuspule lies at the an- 
terolingual base of the nietaconule on M' of L. 
kittjamarpensis (M-'' unknown). It is variable in 
Phascolarctos but is absent in all other 
phascolarctids and ilariids and wynyardiids sug- 
gesting that absence is plesiomorphic. 

10. Anterolingual protocone crest of M^' An 
anterolingual crest extends from the apex of the 
protocone towards the base of the crown on M' 
of L. kiiijatnarpensis and Phascolarctos resulting 
in a relatively .square protocone base. This crest 
is absent in all other phascolarctids, in 
wynyardiids and in ilariids. Absence is regarded 
as plesiomorphic among phascolarctids. 

1 I . Parastylar development on M'- The 
parastyle ('stylar cusp A') is regarded as a sepa- 
rate character (distinct from character 7) because 
it undergoes independent evolutionary change 
relative to .stylar cusps B through E. Small on M' 
of Madakoala and Perikoala\ moderately devel- 
oped in Koobor, Utokoala and Phascolarctos; 
large in Niniiokoala. The small parastyle of 
ilariids and wynyardiids suggests that this is the 
plesiomorphic condition. 

12. Protostylid: The protostylid on M| has de- 
veloped independently in pseudochcirids 
(Woodburne ct al., 1987b). macropodoids 
(Archer, 1978a) and ilariids (Tedford & 
Woodburne, 1987). In koalas, the protostylid 
ranges from weak {Madakoala. Perikoala) to 
very large {Ni/niokoala, Utokoala. Phascol- 
arctos). Lower molars of Koobor are unknown. 
Because a protostylid is absent in wynyardiids 
(Pledge, 1987a) and probably ilariids (Tedford & 
Woodburne, 1987 consider both alternatives), a 
protostylid is interpreted here as derived. 



13. Metasryiid fold: An autapomorphy of the 
Phascolarctidae. Phascolarctos and Utokoala 
have a melastylid fold in which the posi- 
metastylid cristid is continuous with the pre- 
entocristid. Perikoala and Madakoala lack the 
metastylid fold which is represented by a swell- 
ing at the posterior tip of the postmetacrislid. 
Niniiokoala lacks a metastylid fold and the 
melastylid is reduced relative to other 
phascolarctids. Because it is absent in ilariids and 
wynyardiids it is difficult to determine polarity. 
However, considering that discrete melaconids 
and cntoconids is the plesiomorphic condition in 
all marsupial groups, the condition within the 
Phascolarctidae in which these cusps are linked 
by blades, in this case via a melastylid. is inter- 
preted as derived. 

14. Entostylid ridge: Present in L kannnkaensis 
and Phascolarctos and a well-developed cuspid 
in an homologous position in Niniiokoala. Us 
absence in the lower molars o( ilariids and wyn- 
yardiids suggests that absence is plesiomorphic. 

15. Position of the protoconid on Mi: 
Woodburne et al. (1987a) used this to detemiinc 
koala intrafamilial relationships, with the more 
plesiomorphic koalas having a less lingual pro- 
toconid because of weaker protostylid develop- 
ment. The protoconid is within the lingual third 
of the trigonid of Mi in Perikoaku Niniiokoala, 
Utokoala and Phascolarctos. Considering the 
buccal protoconid in wynyardiids and ilariids, the 
more lingual position in some phascolarctids is 
considered to be derived. 

1 6. Ribs on the conids of lower molars: Internal 
ribs on the protostylid, metaconid, protoconid, 
entoconid and hypoconid of lower molars is 
apomorphic and shared by Utokoala and 
Phascolarctos. These ribs are absent in all other 
koalas. Presence is considered derived. 

1 7. Anteriorly displaced entoconid and anteri- 
orly displaced metaconid of M4: Torsion of M4 
such that the entoconid is displaced anteriorly 
relative to the hypoconid and the metaconid is 
displaced anteriorly relative to the protoconid 
occurs in Utokoala and Niniiokoala but no other 
phascolarctids. This condition is absent in ilariids 
and wynyardiids; absence is interpreted 
plesiomorphic. 

RESULTS 

Wagner analysis using the branch and bound 
algorithm produced a single most parsimonious 
tree involving 22 steps with a CI of 0.864 and a 
RCof0.746(FIG.7). 



^ 



MEMOIRS OF THE QUEENSLAND MUfihrUM 



DISCUSSION 

KcccTilclassjficaiionsic.g. WotKlburnc. 1984; 
Apliii & Archer. 1987) ^oup \hv 6 fo&sil ami 
living genera (14 species) ol koalas in Uie 
PhascoUirclidac; however, ihcy also predict a 
iargc morphologic al range of koalu-like animals. 
WuiKUninie (1984) erected Ihe Superraiiiily 
Phascolarctoidca and Aplin A Arclicr ( 1987) Ihc 
Infraordcr Fhascxilarciomorphia. each containing 
only the PhaNcolarclidae. Complex molar mor- 
pholagy and plestoniorphic hnsicranial morphol- 
ogy of Nimiokoulo indicate a more diverse 
inlraorder ol phascoliirclumorphiar»s. 

In some aspects of dental morphohigy, 
Nimokoala appears to have converged on 
pseudocheirids anil ekte»podunliLis, Slight torsion 
is evident in lower molars oi'Nintiokoata and the 
eonset|Uenl arrangement oi" the rneiaconid and 
cntoconid is similar to some psciidocheirids (e.g. 
Hsemiorhmfps archvri) Springer (1987) also 
noted more similarities in the M4 of L 
kantfnkaensisio pscudocheirids Ihiin lo any other 
phjiscoUirctid. This is also Ihe case wilh the Ma of 
Nimiokouki. 

Wcll-devclopcd parastylc. large paracomilc 
and large ncomclaeonulc of the upper molars of 
NitniokiXiUi and the enlosiylid ridge and 
neomorphic cuspid on ihe lower molars are rem- 
iniscent ol (he cuspalc ioph(id)s oleklopodontid 
molars; in parlicular tltc lower moliirs of D^vrt /h.v 
(hi^^uni Rich. 1986 from the Pliocene Hamilton 
Local Fautu, Vicioria. Archer {1976) suggested 
that Ihe ektopodontid molar pattern may have 
evolved through an alijjnmcni of the wrinkles, 
conulesand civnulations in the molars of seleno- 
donl forms and in particular, the koalas. The high 
crowned, highly crenulatc, complex molars oi 
Niniiohmht h.u\y\M.n[ this view. 

Recent clasiitkalions (Archer &, Aplin 1987; 
Marshall et al.. 1989) support an 
ektopodontid/phalangend affinity and place the 
Hktopodontidac within the Phaiangcnndea based 
on the shared strongly ungulate cnstid obliqua 
and reduced melaconid on the lower molars and 
the loss ot PI. Consequently, similarities in 
phascolarciid and ektopodontid molars suggest 
similar ecological niches. Pledge (1982) >iUg- 
gested several possible dietary preferences of 
ekiopodontids, ranging Jrom browsing and fru- 
givonuis to gninivorous iind inseclnorous Sim- 
ilar dietary specialisations could explain the 
complex dentitions oiNimiokoala. 

Since the first fossil koala was described in 
1957 therx? have been two different attempts to 



analyse intrttjamilial relationships. Archer 

1 1 978a) separated phascolnrcrids into Ltiokoala^ 
Ktfohor and Ferikualal Hiasvolatcws lineages. 
Liwkoaki kuTJanmrpensis was considered lo be 
the most plesiomorphic phascolaiciid based on 
the metucunule and lingual buttresses on the 
melacone of the upper molar (Archer. 1978a). 
Pefikoala and Phascolarctos were interpreted to 
he more closely related lo each other than eiUier 
was 10 L kutjamarpensis. However. Atcher 
(1978a) also noted a large structural distance 
separating these groups, ferikoalu exhibited a 
numt>er i)f plcsionK>rphic features relative to 
Phascolarctos and several autapomorphic I'ea- 
ttircs that precluded it being antecedent to 
Phascolarctos. Molar morphology o\ Koobor led 
to its interpretation by Archer (1978a) as the 
sister-group of a combined Phascolarctos! Prri 
kf>alacUdC- 

Woodburne el al.'s (1987a) cladislic analysis 
followed a reassessment of character state 
morphoclines for phascolarciids. Relative posi- 
liiins of PhascolafCfos. Utokoala, Pcrikoala and 
Madakoaln arc confirmed herein (Fig. 7). Rela- 
tionships oi LItokoala have been unclear due to 
its poor fossil record. Archer ( 1 978a) regarded it 
as rhe plesiomorphic sister group of all 
phascolarciids based on a single upper molar. 
Re-analysis of that tooth by Woodburne ei al. 
(1987a) and Sponger's (1987) 2 isolated lower 
molars and upper molar fragments of L. 
kurumkaetisis suggested Litokoala was more 
closely related to Phascolarctos. Litokoala miUe- 
rial from Riversleigh supports Woodburne et al.*5 
(1987a) hypothesis. 

Features now found to support \\^e, Lifokoola 
and Phascolarctos relationship include the 
neomorphic euspuic ai the anlcrolingual base of 
the mciaconule on M' (and M*^"* in 
Phascolarctos). the metastylid fold in which (he 
postmeiastylid cristid is a continuous fold with 
the preeniocristid and the internal ribs on the 
cuspids of lower molars. Features, regarded by 
Springer (1987) as aulapomorphies of L 
LuutnkavnsishiW which are in fuel variably pres- 
ent in Phascolarctos include anteriorly bifurcate 
pre-enlocristid. reduced talonid on M^ und paral- 
lel arrangement of the cristid obliqua and 
poslproiocrislid on Ma wherein ihese crislids do 
not meet end to end. The latter two fearurcs are 
also variably present in Nimiokoala. Litokoala is 
a derived relative to Phascolarctos in having a 
more crcscentic paraconule and neometaconule 
in the upper molars, a well-developed melaconid 



MM/OKOAL\ GEN. NOV^NKW KOALA FROM RIVHRSLUKHI 



m 



pos(crobuccal crcsl on M^.aand in having a wcll- 
devclopcd poNlentlingual prulomslid un M^. 

\iiHit)k(fciIay LitokiHiid and Fhasntkirctos tofm 
a cbdc by shjiring i\ proH>siylc in (he upper nio- 
iars, strong pt)slerolinguaI paracristae. a large 
pruloslylid on Mi and a well-developed en- 
losiylid ridge. Liiokoakt and Nuniokaalti arc de- 
rived relative to Phosi'olarcros in the shared 
trcsccnlic paruconulcand neomelaconutc m M'"^ 
aiui a well-developed patasiyle on M'- iMokonia 
and Nimiokoala synapt)morphies, interpreted as 
convergences (Fig. 7) are: well-developed, highly 
Cfcscenlic paraeonule and neomeiaeonutc, large, 
pyramidal paraslyle and extreme torsion on M; 
such thai the entoeonid and meiaconid are dis- 
placed anteriorly relative to the hypoeonid and 
proioconid. respectively. Nimiokoala is derived 
relative lo other phascolarctids in having strong 
posterobuccal crests Irom the apices of Uie pR>- 
locone and metaconule. a disconiinlious 
neometaconiile which is subdivided iniotwocus- 
palc pans, a posicrolingual cuspulc onP^. a well- 
developed posterolingual prolostylkl crislid on 
M). weak nictasiylid in ihe lower molars, an 
anicrobuccally directed precntocrislid (an- 
icrolingually directed tn other phascolaieiids) 
and a neonuuphic cuspid occupying the irigonid 
ha.sin between mctaconid and protoconid on M:.a 

There are doubts about die i!\cKision t)\'K(n}hor 
in Phascolarctidae. Pledge (1987b) suggested 
lliat Koohori^ allied lo ilariids such as Ktttcrtntju 
Hgiwia, Figure 7 may support the hypothesis thai 
it lies outside the Phascolarctidae. Clarificaiion 
of Koobor's relationships requires discovery of 
lower molars because these difter significantly in 
ilariids and phascolarciids. 

l-xcept hir 2 partial skulls irom Riversleigh, 
extinct phascolarctids are represented by isolated 
teeth 01 dcniiiions. As a result, current under- 
standing of the basicranial region is based <in the 
modern species- Pltoscalonuts cine feus has an 
autapomorphic bilaminar bulla wlierein the lym- 
j>;»flic cavity is roofed by both an alisphenoid 
tympanic process and the squamosal cpilympanic 
wing(Aplin. 1987). The partially pteserved basi- 
cranial region of Niinutkoala gieystancsi (Fig. 
I A-C) exhibits the plesiomorphic diproioduniian 
condition wherein the alisphenoid forms the roof 
of the lytnpanic cavity, suggesting a large struc- 
tural distance between these koalas. The 
plesiomorphic basicraniumoI^V. greystanvsi sup- 
potls Ihe hypothesis thai phascolarciids diverged 
from near the base of the diprotodontian tree 
(Archer J 976). 

Intraijcncric relationships of LitokihUa ait: Uif 



ficuli lo inlcrprct due to a lack of eotnpafiibk 

material between the species, Ihokoalo kiU^- 
///iA</^/7\7.v appears to be plesiomorphic reialivcto 
L kuijinruirpensis because it lacks a proto style 
and the bulbous cuspulc at the anterolingual b^sc 
of Ihe metaconule on M^ However, these features 
may be a rcsuh of changes along ihe tooih ru'w 
(ie. an artefact of comparing an M^ with an M') 
raihcnhan interspecific differences. 

Litokoaki kanunkaemis J rum carlv-HliMillc- 
Miocene System C( Archer ct at., IS)S9; 1991 Its 
the same ago nr slightly younger than, tbc- 
Kuljamarpu Local Fauna. 

ACKNOWLEDGFMFNTS 

Weihank Neviile Pledge and Mike W<H)dbumc 
who read a draft of this paper. Linda Gibson. 
Ausiralian Museum for access U» collections aivd 
Ross Arneti for assistance v\ith ph<nography. 
Riversleigh research is supported by the Auslnv- 
lian Research Grant Scheme; Naitunal Esialt 
Gnmis Scheme (Queensland); University of 
N.S.W.;CommonweaithDepartmenU)fLnviron- 
ment, Sports and Terriiories; Queensland Na- 
tional Parks and Wildliie Service; 
Commonwealth World Henlitge Unii> ICI Aus- 
tralia; Australian Geographic Society; RoyaJ 
Zoological Society of N.S.W.; Linnean Society 
of N.S.W.; Century ZifK*; Riversleigh Socidy; 
Elaine Clark; Miurgarci Bcavis; Martm Dickson; 
Sue & Jim Lavaxack; and Sue & Don Scoit-Onf. 
Field suppoii canie from hundreds of volunteers 
and staffand students ofthc Uni versify of NS.W 
Skilled preparaljon of Riversleigh maicriaJ has 
k-ieit cwnicd out by Anna Gillwpic. 

LITERATURE CITPJ) 

APUN, K. 1987. Ba5icr<Knial nnaioniy ot ihe caily 
Miocene dtpruuxlonuan >Vv»v/7r////7 hassf/tfM 
(Mnrsupiaha: Wynyardiitliie) and Us unplicalions 
io( wynyardiid phylogcny md classificaiion I'jj. 
36*^*>I In M Archer (cJ.). Possums and rpas- 
Hums: studies in evolution. (Siirrcy Bcaiiy Sl Soiis 
and Ri>yal /.oologicjl Socicl> of NSW: S)dnc> ). 

APLIN, K. & ARCHER. M, 1*>S7 Recent advances ui 
marsupial sysiomuiics wiih uiicw synciviic clan- 
Kilkaiion. Pp. xv-lxxii. In M Archer (v-d.), Ptw- 
sums and opossums; studies in evci)ution.{Sum:y 
Bcauy & Sons and Royal Zoological Society of 
NSW: Sydney). 

ARCHER. M ]9lh. Rhascobrclid ongitis and the |xi- 

lential oi ihc sclcnodont molar in the cvoluiiunof 

diproiixJonl m.irsnpial.s. Mcimjirs v>t the Queens- 

biiUMuM:ui)i 17; 367-37 L 

I97JU Kt»;*las (phu.sci*liiieudst and their signiff- 



228 



MEMOIRS OFTHEOL'EENSLAND MUSEUM 



cunfc in marNiipial trvohUion. Pp. 70 H In T. .1. 
Bcrgiiv. (cU.). llic Koala. (Zuulogicai Packs^ 
Board N.S.W: Sydney). 
t978b. The natui-e of ihc molar-preniolar houndary 
in marsupials and a reinterprcliti ion oCilK'hunniI- 
ogy 1)1 marsupial check tccllv Mt-muirs ot the 
Queensland Museum 18: 157-164. 
1087. fosunis andi>possunis; studies incvoluuon. 
(Surrey Bcnity & Sons and Royal /.oolugical 
■Scxicty of NSW. Sydney). 

AKCHER, M..GODTIIELP. H.J I AND, S.J. & MEG- 
IRIAN. D I98'J. Fossil mammals of Rivcrslcigh. 
fionhwcsiem OucetiHland. prclmnnai^ overview 
orhinsiratigraphy. cnrrclalion and cnviammental 
chance, Australian Zt»ologis*i 25 27-ft5. 

AKCMhR. M.. HAND. S.J. & GODTHIil.P, H. ]99\. 
Rivcrslcigh. {Rccd: Sydney) 

LUCKETT. W.P lOO}. An oniogcnciic asscssmcnl of 
denial lujmologie.s in Uierian mammaK. Pp- IK2- 
204 In S/al.ty. FS,. Novacck, M J, Si McKcnna. 
M.C (edsK Manunal ptiylugcny ML'S(»/oicdifler- 
cntialum, rnullilnbercuhuos. momtircmes> early 
dierians and marsupial. (Sprm^LT-Veriai^. New 
Ytirk). 

MARSHALL. L.G.. CASE. J.A. & WOODBLIRNE. 
M.0. 1*JS9. Phy|(»genelicrelalionship.sorihe (am- 
ilirs of mitrsupials. Current Mammoiogy 2: 433- 
502. 

PLEDGE. N.S. ltJ82, Eniy malic Ekiuiuuhn: A cttsc 
history ol palafoniv>li)git.al iiUerprclaliim, Pp. 
477-4Si:i In P V Rich & E.M. Thompson icds). 
Ihe (ossil vencbrale record ol Auslrata.sia. (Mni> 
ash University OfTsci Priming Unit: Clayumi 
iyX7a. Muramura wHliamsi, a new genus and ni>c 
cies of'wynyardiid (Marsupialia: Vomhatoide.i) 
Irom ihe middle Miocent' CiadunnaFormali(»nMr 
S(MUhAusiralia Pp V) 3-401 > InM Archcr(edl.. 
Pojs.sums and upo.sMims. sludich in evolulmn. 
(Surrey Beany A: Stvns and RoyaJ Zoological 
Society of NSW Sydney). 
l9X7h. hnrt-nnrjtt n^atnu . a new genus and .species 
i.}\ pinniiive vombaloid marsupial \'rom Ihc me* 
dial Miocene Ngtima Local Fiiijnaof Souih Aus- 
tralia. Pp. 419-22. In M. Archer (ed.). Possums 
and opossums: sludics in evolution. (Surrey 
Bcaliv &. .S*>ns jind Roynl Zoologivul Sociciy i>f 
NSW^: Sydney). 
1992. The Cuiran^ulka Local Fauna: a new Late 
Tertiary (bssil asse?Tihlajie from Yorkc Peninsula. 
South Ausifalta. the Beagle: Records of ihe 
Niirihcrn Terrilory Museum of Arts and Sciences 
9: ll.S-l'J2. 

'RICH.T. H. 1982. Monotrcmcs. placemais and marsu- 
pials: llicM ivcord in Auslralia and its biases. J'p. 
3K5-477 In P V. Rich & EM. Thompson iah). 
The fossil venebiaie recufd ot Ausifala.sia. 
(Monash University Ollsei Prinling L'nil: Play- 
lon). 

i9"8(>. Darcius dug^ani. a new eklopodi>ntid 
(Marsupialia. Phalangeroidea) from (he early 
Pliocene Hamilton Local Fauna, Auslralia- Uni- 



■versily ol* California Publications m the Geologi- 
cal Seicnees LM.(i«-74. 

RICH. T.H.V. S!. ARCHER, M. 1979. Nmiilamadtta 
stiidcri, a new diproiodonlan fmiu ihe medial 
Mioceneof South Australia. Alchcringa3: 197- 
208. 

SPRINGER, M.S. 1987. Lower molars of Litokoala 
(Miu-supialia;Phasco!arctidac) and their hc;iring 
on pha-scolaalid evolution. Pp. .^19-25. hi M. 
Archer (ed.). Possums and opossums: studies in 
evolution. (Surrey Bealty & Sons and Koyal Ztio- 
logicul Society of NSW: Sydney). 

STIRTON. R.A, 1957- A new koala (nxn the Pliocene 
Pjilankarinna Fauna ol South Auslralia. Record* 
of ilie South Ausiralian Museum 13: 7| 7S. 

STIRTON. R. A, TEDFORD. R.H. & WOODBURNE, 
MO. 1967. .A new Tertiary formaiion and fauna 
from the Tirari Desert, South Ausiiulia. Records 
ol the South Australian Mu.seuin 1 5: 427-462. 

TEDFURD. RM..^ WGGDBliRNL:. M.O. I9H7. Tlic 
llariidae. anew family ol vombalitonn marsupials 
from Miocene strata of South Australia and an 
evaluation oTlhe homolgy of inolat cusps in the 
DiprotodiMUidae. Pji 401-418 InM Archcr(etf). 
possums and opossvmis; studies in evolution. 
(Suni'y Beatiy & Suns and Royal Zctological So- 
cictvol NSW': Sydney). 

WOODBURNE, MO. 1984. Families of marsupials: 
jvlaiion.ships. evolution and biogcograptiy. Pp. 
48-7 1. In T, W. Broadhead (ed ). Mammals: notes 
for a ■'thorl course. (PnIeonioU>gical Society: Boul- 
der). 

WOODBURNE. M.O.. TEDFORD. R H.. ARCHER. 
M. TURNBULL, W.D , PLANE. M.D. &. 
LUNDELIUS. ED. 1985. Biochronology of the 
continental mammal record of Auslralia and New 
Ciuinca. Special Publication, Stmili .Australian De- 
paitmeni of Mines and Energy 5: 347-363. 

W(X)DBURNE. M t), Tr'.DFORD. R.H., ARCHER, 
M- A PLEDGE, N.S 1987a. Mudukaaht, a new 
genus and iwo species of Miocene koalas 
{Marsupialia:PhHstolarclidac) Irom South Aus- 
lralia. and a new species of Perikinikt. Pp. 293- 
317 In M Aa'her (ed.). Possums and opcts.surns: 
.studies in evolution. tSurrev Beatty & Sons and 
Rovnl Zoological Societv of NSW: Sydney). 

WOODBURNE. M.O..TEDFbRD, R H & ARCHER, 
M 1987b. New Miocene ringtail pt>ssuins 
(Marsupialia: Pseudochciridae) Injin South Aus- 
tralia. Pp. 639-79. In M. Archerted.). Possums tind 
opossums studies in evolution. (Surrey Bealty & 
Sons and Royal Zoological Society of NSW. Syd- 
ney ). 

WOODBURNE, M.O,. MACFADDEN, B,J-. CASE, 
J. A.. SPRINGER, M.S.. PLEDGE. N.S., 
P(3WER, ID , WOODBURNE. J,M. & 
SPRINGER. K.B. 1993. Land mammal hiostrat- 
Igraphy and niagnetostraiigraphy of the Eitadunna 
Formaiion (late Oligocene) of South Australia. 
Journal ol' Vertebrate Paleontology 13; 483-515. 



A KINGFISHER (HALCYONIDAE) FROM THE MIOCENE OF RiVERSJ>ElGH, 

NORTHWESTERN QUEENSLAND. WITH COMMENTS ON THE EVOLUTION OF 

KINGFISHERS IN AUSTRALO-PAPUA 

WALTER E. BOLES 

Boles. WE. 1997 06 30: A kingfisher iHaloyonidaei Truni the Miocene of Rivcrslcigh, 
norlhwesicm Queensland, wiih commcnls on rhcevoluiion olkingfishcrs in Ausimlo-Pupua. 
Memoirsofihe Queensland Museum 41(2); 229-234. Brisbane. ISSN 0079-8835. 

A Miocene kingfisher from Riverslcigh, norlhwcstern QueensiantI, represented by a com- 
plete carpomelacarpus, is ihe cariicsl record of the Hakyonidac from Australasia, h shares 
similariiies with several modem genera, bui a positive generic identification cannot be made. 
Although it can be dlstingiiislied froin exiaiii species, this skeletal elemem is insufficient to 
creel a new genus. A processus dcntiformis in Tanysipwra and Mcluiam and its absence in 
Todimmphus and other genera suggest that the lormcr genera arc among the more pnmitixe 
of the Ausiralo-Papuan kingfishers. The less developed processus denliforniis in the 
Rivcrslcigh specimen is consistent with it being an earlier member of ihe Todiramphus 
lineage. Of living kingllshers examined, all that retain the processus denliformis are inhab- 
itants of rainforest, \2iKifi ^fisher. HakyofUdcH. Hivi-nletfi/h Miocene, evolution. 

Waiter E. Boles. Attsmdum Mustum, 6 C(dlege Street. Svdney N.S.W. 2000, Australki: 
received 4 November J 996. 



The kingfishers (Alcedinidae s.l.) are subdi* 
vicled into 3 suhfamilies. DNA-DNA hybridisa- 
tion studies (Sibley & Ahlquisi, 1 990). suggested 
that these should be recognised as families. 
Cerylidae do not occur in Australasia. Al- 
cedinidae {/river kingfishers*) and Halcyonidae 
i= Dacelonidae auct] ( 'tree kingfishers' ) arc rep- 
resented in Ausiralo-Papua by 5 species in I 
genus and 21 species in 5-6 genera, respectively 
(Bcehleret al.. 1986; Fry et a!.. 1992; Christidis 
& Boles. 1994). 

There are no named Tertiary forms from out- 
side Australasia (Olson, 19S5) Mourer- 
Chaurvire {\9^1) listed this family (Alcedinidae 
s.l.) from Eocene-Oligocene deposits at Quercy, 
France, and Olson ( 1985) noted thai he had ex- 
amined specimens close to this family originating 
from the lower Etx:ene of North America and the 
medial Eocene of Germany. All Australian Qua- 
ternary kingfisher material is referable to modem 
taxa: Alcedo cizurea, Dacelo novae guineae, 
Todiramphus pyrrhopygia and To, sanctus 
(Baird. 199 1 ). No Tertiary kingfishers are known 
from Australasia (Fordyce, 1991: Vickers-Rich, 
1991). 

Described herein is a Miocene kingfisher from 
Rivcrslcigh. northwestern Queensland. 

METHODS 

Measurements (Stcadman, 1980) were made 
with vernier calipers accurate to tll)5mTn and 
rounded to the nearest (11mm, Temiinolooy of 



bones largely follows Baumel & Wilmcr ( 19^>3). 

Todiramphus and Syma are considered distinct 
from Halcyon, following Christidis &. Boles 
( 1994). Institutional prefixes are AM (Australian 
Museimi). ANWC (Australian National Wildlife 
Collection), MV (Museum <y\ Victoria), QM 
(Queensland Museum) and USNM (United 
States National Museum). 

SYSTEMATIC PALAEONTOLOGY 

Family HALCYONIDAE 

Although the Halcyonidae includes some of the 
largest kingfishers in the world, si/e is not a valid 
character for family allocation o\ osieological 
■material. Australia's 2 Alcedinidae, Alcedo 
pusilla and .4, azuren. are the country's smallest 
kingfisher species (wing lengths 55mm and 
75mm. respectively), but the closely related A. 
websien of New Britain has a wing length of 
9()mm,ovcrlapping in si/e the smaller halcynnids 
(e.g., Todiramphus macleavii, wing length 
90mm). 

The carpometacarpus of the Halcyonidae can 
be distinguished from that of the Alcedinidae and 
Cerylidae (Table 1) and on this basis the 
Rivcrslcigh Ibssil is assigned lo the Halcyonidae. 

HaLcyonid gen. indei. 
Fig. IC 

MATERIAL QMF297 1 9. right carpometacarpus with 
only mmor abrasion to some surfaces from ?niiddic 



230 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE I . Characters for separating the carpometacarpus of the Alcedinidae, Cerylidae and Halcyonidae. 



Character 


Alcedinidae 


Cerylidae 


Halcyonidae 


proximal border of dorsal carpal trochlea 


more angular 


more rounded 


more angular 


development of os metacarpalis alulare 


more gracile 


more robust 


more robust 


orientation of os metacarpalis alulare 


more caudal 


more caudal 


more proxunal 


lip of processus extensorius 


rounded 


expanded, slightly rugose 


rounded 


position of processus intemieiacarpalis 


more proximal 


more proximal 


more distal 


width and distal extension of sulcus 
inierosseus 


narrower, not as 
extensive distad 


broad, extending almost 
to facies digitalis minor 


broad, extending almost 
to facies digitalis minor 


plane of synostosis metacarpalis distalis 

and distal ends of os metacarpalis major 

and OS metacarpalis minor 


OS metacarpalis minor 
depressed belovs plane 


OS metacarpalis minor 

depressed slightly below 

plane 


Hat, coplanar 



Miocene to early laic Miocene Last Minute Site from 
System C (Archer et al., 1989, 1994). This site is 
interpreted to represent shallow pools or even emer- 
gent accreting surfaces, and is dominated by terrestrial 
vertebrates including the possums Djilgaringa 
gillespieae Archer el al., 1987 and Sirigocuscus reidi 
Fhuinery & Archer, 1987. Other avian laxa from this 
site are a range of passerines, including the logrunner 
Orfhonyx kafdowinyeri Boles. 1993. 

DESCRIPTION. Length 15.8mm. Length of 
spalium intcrmelacarpale 61% of length of car- 
pometacarpus. Processus dcnliformis low and 
pointed, located about midway between the distal 
edge of facies arlicularis alularis and the 
cranialmost point of facies arlicularis digitalis 
major. Os metacarpale majus of about equal 
thickness for entire length, in ventral view. Spat- 
ium intcrmelacarpale gradually becoming wider 
distally. Processus intermetacarpalis far proxi- 
mally in spalium inlermelacarpale. 
REMARKS. Among extant Australo-Papuan 
Halcyonidae. Tanysiptera and McUdora have a 
low. Hat processus dentiformis, Syma has an al- 
most non-existent processus dentiformis as a 
barely raised roughened area, and Todiramphus 
and Dacelo, as well as Afro-Asian Halcyon, lack 
il (Boles unpubi. data). X-!"ay photographyof 
study skins showed Actenoides and Lcicedo have 
a low processus dentiformis. but Halcyon 
(Pelargopsis) and Clyrocey.x do not. The x-rays, 
while sufficient for determining this process, are 
not adequate for detailed comparisons of these 
laxa. Other than size, there are not substantive 
differences between the carpomclacarpi of 
Todiramphus, Dacelo (and presumably 
Clyioceyx)\ Syma differs only in its low processus 
dentiformis. Because Dacelo and Clxfoceyx are 
considerably larger (Z). novaeguineae 32.5- 
35.6mm), they are noi considered further. Subse- 



quent comparisons involve Todiramphus, Syma, 
Tanysiptera and Melidora. 

The fossil (length 15.8mm) is in the size range 
of Tanysiptera (sylvia 14. 0-15. 2mm; gaUitea 
15.2- 16.5mm) and Todiramphus [sanctus 14.8- 
15.1mm; macleayii 14.5- 15.7mm; pyrrhopygia 
15.7- 16.5mm; chloris 16.5- 1 7.8mm). It is larger 
than Syma (torotoro 13.5mm; megarhyncha 
\ A Axw\x\)dW(l^m'iX\\cx\\\ixn Melidora macrorrhina 
(18.9 mm). It differs from Tanysiptera and 
Melidora and resembles Syma and Todiramphus 
by being more slender and by having spalium 
intcrmelacarpale longer relative to the length of 
carpometacarpus and the dorsal rim of trochlea 
carpalis extending less dislally relative to the 
ventral rim. The fossil differs from Todiramphus 
and resembles Tanysiptera, Melidora and Syma 
by having processus dentiformis present. This 
process, however, is narrow and pointed, rather 
than broad and tlal as in these genera (and larger 
than in Syma), and is situated more proximally 
relative to the spalium inlermelacarpale and pro- 
cessus intermetacarpalis than in Tanysiptera and 
Melidora, and more distally than in Synui. From 
all 4 genera, the Riversleigh specimen differs by 
having the caudal edge of os metacarpale majus 
siraighler and less caudally concave, making os 
metacarpale majus thicker and spalium inter- 
metacarpale proportionally narrower relative lo 
the width of the carpometacarpus. 

The significance of these differences is uncer- 
tain. They are individually minor, yet within the 
Halcyonidae the amount of variation in this bone 
is little so that these differences may assume 
greater importance. Variation in the caqximcta- 
carpus, however, is not representative ol' that of 
the rcmainderof the skeleton or indeed the rest o\' 
the morphology (Boles unpubi. data). 

The fossil cannot be assigned to Tanysiptera, 
Melidora, Syma or Todiramphus and can be dif- 
frereniialed from extant species of these genera. 



A MIOCENE KINGFISHER FROM RIVERSLEIGH 



231 




FIG. 1 . Carpometacarpi of Auslralo-Papuan halcyonid kingfishers, in ventral view. All right side except for B. 
A, Melidora macrorrhina (ANWC CORS-53). B, Tanysiptera sylvia (AMO.60926). C, Rivcrsleigh 
Halcyonidae gen. indel. (QMF297I9). D, ToiUramphus sanctus {MA0.51\%2). E, Dacelo novaeguineae (AM 
0.59217). Scales = 5mm. 



Overall it has the greatest resemblance to species 
of Todiramphus in size and morphology, despite 
the presence of a small processus dcntiformis. 
Given the relatively limited importance of carpo- 
metacarpal variation in the halcyonids and the 
limited fossil material it is imprudent to recognise 
a new genus al this time. 

DISCUSSION 

To place the Rivcrsleigh fossil in perspective 
within halcyonid kingfisher evolution, the prim- 
itive members of the family must be identified. 
Fry ( 1 980a, b) employed 3 criteria for this. Prim- 
itive kingfishers have 1) generalised diets and 
relatively unspecialised modes of foraging (i.e.; 
sitting and pouncing, non-fishing); 2. stable hab- 
itats and not of recent origin (rainforest), within 
which they may be discontinuously or relictually 
dislributed;and3.oligotypic genera (i.e. with few 
species) without close relatives. Fry ( 1 980a) con- 
cluded that kingfishers (all families as recognised 
here) arose in Malesia, the area between Indo- 
china and the Coral Sea. 

Prominent among the primitive forms were 
halcyonid kingfishers, many o\ which live in 
Malesian rainforests. Fry (1980a) speculated that 
'the Daceloninae [= Halcyonidae] have a history 
of evolution in eastern equatorial rainforests al- 



most as ancient as the mid-Cenozoic origin of the 
Alcediniformes [sensu Feduccia( 1977)]'. By the 
early Miocene, the present geographical config- 
uration of Malesia had been reached. This, in 
Fry's opinion, provided 'ideal circumstances for 
the multiplication of species, resulting in a fauna 
of forest-dwelling, non-fishing kingfishers ... At 
some more recent time, perhaps about the mid- 
Pliocene, this fauna gave rise to a lineage, Hal- 
cyon [s.l.], adapting to more open habitats'. 

Under this suggested sequence of events, the 
rainforest-dwelling Tanysiptera, Melidora, Acten 
oides and Lacedo would be among the more 
primitive genera. Todiramphus, included by Fry 
(1980a) in his open habitat Halcyon, would be 
more derived. Presumably Synia (also included in 
Halcyon sensu Fry [1980a]j also was considered 
by Fry (1980a) to be a more derived taxon. Al- 
though entering open country adjacent to forest, 
the two species ofSyma are essentially rainforest 
inhabitants, particularly in New Guinea (Coates, 
1985). 

In addition to sharing the primitive characters 
proposed by Fry (1980a, b), species of Tan- 
ysiptera and Melidora (as well asActenoides and 
Lacedo) have a low but well-defined processus 
dentiformis on the carpometacarpus. This is ab- 
sent in the other halcyonid genera examined, the 
Alccdinidae and Cerylidae, and some other cora- 



2i2 



MEMOIRS OFTHEOUEENSUND MVSEUM 



ciitbrm families leg. Momotidac. Coraciidac). If 
any of ihese fyrtiilies is used for outpA^up com- 
parisons, the suggested polarity oi Ihc processus 
dcnliforrnis h thai its presence is dcnved. A sim- 
ilar comparison using olhcr coraciiform families 
(Tod i due. Pho^niculidue, Upupidac, 
Bucerotidae). in which the structure is present. 
^ivt*N the opposite conclusion: presence of the 
pmcessus dcnliforrnis is primitive, its absence 
derived. Within the Mcropidae, the structure is 
prcseni in some species {Merops ormtms) arni 
ahseni in others {M. optaster). The significance 
of Ihis variation is unknown, as arc the tunclumal 
aspects ol the processus denliformis. Thus itie 
polarity of this character's presence i.s not known. 
(This, oi course, assumes thai the processus 
deniilormis is homologous across ihe order. 
Whether this is so, and what relationship it has to 
tlie siiniliir process lound in the majority of the 
Passeriformes, is unknown.) Within the 
halcyonid kin^dshcrs the presence of the proces- 
sus denlifontiis exhibils a strong correlation with 
Fry's ( I980a,hj primitive criteria. 

Suporficially there seems to be lidle in conmion 
exieriially between Mciidora and lanystpwra 
beyond the hiisic kingfisher similarhics. Hachhas 
specialised generic characters: a hooked bill in 
Mvlidom (Hooked-hilled Kingfisher) and elon- 
gated, spalulaic central iccirices in Tan\sipiera 
(paradise kingfishers). Metiiioni macronhimi is 
a ralher drably coloured species, Other ihan blue 
scalloping on the crown, Ihc plumage is a combi- 
nation of bixjwns and white. The underside is 
white, while the hack, rump. tall, and wings arc 
dark brown with paler broAO Vv-alloping. This 
plumage is quite unlike that of the pariidise-king- 
lishcrs '/anysiptera, adults of which arc strikingly 
patterned in unmarked blues and blacks, and usu- 
ally either white or buff/rose. Thejuvenile plum- 
;ige of Tonysiptera, however, is brown with 
scalloping, and Fry ( l^SOb) was 'impressed by its 
{MvHdora''s\ plumage resemblance to the distinc- 
tive juvenile of Tanysiptera i^ahfea\ That these 
two genera mighi be closely related was sug- 
gested by Fry (1980b). who thought it 'possible 
that Melidont and Tunysipteru are of immediate 
common descent and the former is a specialised 
derivative that has retained, in the adult, the an- 
cesiral juvcnde plumage'. The presence ol sim- 
ilar plumages is also evident in female Lacedo 
and Nume species ol Actenoidcs. niMably A 
pHmeps and A. Undsayi of all ajtcs. This resem- 
blance beiwecni Actenoides and Ikmysipura and 
l»eiwe«n LmTdv and MvUdont w;js commented 
onhyFiry(1980h). 



Bell (1981 J also considered that MflUhnt was 
closest to Tanysiptera. He iK>ted thai the call 
notes of these two species were similar and the 
distress notes identical. They have simihir skele- 
tal proportions, particularly in the rehttivc length 
of Ihe legs wlien coniparcd to Todiramphus, Hid- 
cyan or Pa celo {Boles, unpuhl. data), These two 
genera thus have more similarities than might lie 
immediaiely obvious, They also share habitat 
prelercnces. Although Mvlidora martonhiua 
and Tanysipteni species will enter miingrovex, 
teak pl.iotations and drier adjacent country, they 
arc primarily occupants of rainforests. Lacedo 
puJchvlla und the 6 .species i)iAnefundt's inhabit 
rainforest, preferably in a primary, undisturbed 
slate 

Species oi' Todiramphus are unifonn in pUnri- 
age. Most have a variation of the basic pallcni of 
green or blue uppcrparts and while or light orange 
under parts and collar. Several subgroups can be 
discerned, but Ihey still show only small diver- 
gences from this general form. These species arc 
almost all sit and pounce feeders, Habiiai prefer- 
ences aniong species arc more varied, ranging 
Irom rainforest to ttpcn country. Williin Ausiralo- 
Papua. hi»wever, few occur in rainlorest and there 
is a decided bias towards open loreM. woodland. 
mangroves, clearings and open country. Most 
rainforest inhabiting forms are found on islands 
of the southwest Pacific. 

The processus denliformis in the Riversleigh 
kingfisher is smaller than thai in primitive mod- 
em torms. This could indicate thai it has been 
undergoing reduction since the split of its lineage 
from that of TanysipK^rn-Mfluiofn. In this re- 
spect it is consistent wiih what would be [>Tcdicted 
for a primitive species of Todiramphtis. The bone 
exhibils a hirgely lodirarnphtts chacacict while 
retaining this more primilive halcyonid lealure. 

The ideniilicalicvn of the Riversleigh fossil as a 
halcyonid is compatible with Fry's ( 1 980a) inter- 
pretation, as IS considering ihe piesence of the 
processus denliformis as primitive. According to 
Fry's scenario, un early Miocene kingfisher 
should be a pi'imitive form. Hiscriteria, however, 
are not useful in this silualion. The foraging meth- 
ods of the fossil cannot he determined, nor can its 
systematic isolation be ascertained. The 
Riversleigh habitat is considered (Archer cl al, 
1992) to have been rainlorest. but this ciinnot be 
used as a characior for making a taxonnmii. de- 
terrninaiion, 

Although this fossil permiis idenlilicadon as a 
halcyonid kingfisher, it is noi clear whether ihis 
species belongs loan existing genus or should be 



A MIOCENE KINGFISHER FROM RIVERSJL£1GH 



«3 



allocated m a new one. The presence oi a feature 
found in living primitive genera and ihe 
kinyt'isher's t;K.\*iirrencc in what is considered lo 
have been rainlbrcst suggest ihai ii. too. w;in u 
more primitive fomi. This \^ consisicnl with the 
sicqucnceof evolutionary cventssuggesltid by Fry 

ACKNOWLEDGFMFNTS 

For access to specimens I ihcink Wayne 
Longmore (Queensland Museum). Jerry vanTcts 
and Richard Schiulde (Australian National Wild- 
hlc Culleclion). Les Chrislidis and Rory O'Brien 
(Museum of Vieiotia). itnd Slorrs Olson (United 
Stales National Museum). The Auslralian Mu- 
seum provided a venue in which to work and 
funds 10 support this research. The Rivcrsleigh 
material was collected via an ARC Pnj^Tainme 
Gtant 10 M. Archer; support Irotii the University 
of New South Wales; a grant tiom the rvp;iri- 
ment of Atls, Sport, the r-lnvironmenl. Tounsm 
and Territories to M Archer. S. Hand and H. 
Godlhelp; a grunt from the National Estate Pro- 
jjratnme Grams Scheme to M. Archer and A. 
Bartholomai; and grants m iiid lo the Riversleigh 
Research Project from Wang Australia Ply Ltd, 
ICI Australia and the Auslralian Geographic So- 
ciety. 

REFERENCES 

ARCHKR, M.. GODTMULP, M, HAND, SJ. Si 

MEOIRIAN. D. I98S). Fossil manitnals of 
Hiversleigh. nunhweMcni Quccnslarul: prclitni- 
ntiry overview of biosiraiigraphy. conclalion and 
cnvironniemal change. Au.sLralian ZotKjgisi 25: 
29-65. 

ARCHER. M., HAND, S.J & GODTHCLP, II 1991. 
Rivcrsleigh. (Reed; Sydney). 
1994 Riversleigh. 2nd edition. (Reed: Sydney). 
1995. Terliarv environmental ;md hiotic change in 
Australia Pp. 77-90. lnVrba.E.S..Denion,G.H.. 
Panndge,T.C-& Burkle. L.H. (cds)Palcoclimaic 
and cvolulion, wiiti tmphnsis im human origins. 
(Yale University Press: New Haven). 

ARCMHR. M., TEDFORD. R H it RICH. T.H. 1987. 
Tliir Pilkpildrid.ic. n new family and four new 
species of ?peiaurid possums (Marsupialia: 
Phalungotida) from ihc Au'^tralian Miocene. Pp- 
607-627, In Aivher, M. (edt Possums and opos- 
sums: Studies m evolution. (Surrey Bcatiy. Chip- 
ping Norton) 

BAIRD, R.h*. 1991. Avian fossils from Ihc Quaicniarv 
of Ausu-alia. Pp. 809-870. In Vickers-Ridi. P.V'. 
Mtmaghaii. .I.M.. Baird. R.F. & Rich, T.H, (cds) 
Vertebrate palaconiology of Australasia. (Pioneer 
Design Studio: Melhoume). 



BAUMEL, J,J. & L.M. WITMER. 1993. Osteologia. 
Publications of ihe. Nuilal! Ornithological Club 
23:45-132, 

BEEHLER, B.M.. PRATT. TK & ZIMMERMAN. 
DA. 19S6. Birds of New Guinea. (Pfiaeeloit Um- 
versity Press: Princeton). 

BELL, H.L. 1981. Jnfurmation on New Guinea king- 
fishers. Aleedinidae. IIms 123; 51-61. 

BOLES. W.H. 1993. A iogrunncr Onhoffvx from iho 
Miocene o( Rtvcrsicigh. nonhwesiem Qucen*^ 
land. Emu 93. 44-49. 

CHRISTIDIS. L. & BOLES, W.E. 1994. laxunoiny 
and si>ecics of birds of Australia and its lerriltiriev 
RAOU Monograph 2 (Royal Australasian OnU- 
tJioIogists Union: Melbourne). 

COATES, B.J. 1985. The birds of Papua New GuiiitM. 
vol i . (Dove: Brisbane). 

FLANNERY.T F,& ARCHER,M. 1987 StrifioatsaiT 
tfidi and TrUhosurus dickMmi, two new li."*i.sil 
plialangerids (Marsnpialia: Phulangcn)idea) from 
the Miocene of noithwcsiern Quccnslajid. Pp. 
527-536. In Archer. M. (cd) Possums and op<j.s 
sums; .Snidics in evolution- (Surrey Bratly: Chip- 
ping Norton). 

FORDYCE. R.E. 1991 . A new look at the fossil veac- 
braie record of New Zealand. Pp. I I9i-I31(>. In 
Vickcrs-Rich. P.V.. Monaghun. J.M.. Baird. R.F, 
&. Rtch, T.H. (cds) Venebiaie pakieontology ol 
Anslralasia (Pioneer r)csign .Smditr Melk^ume) 

FEDUCCIA. A. 1977. A model for the evolution of 
pcreliing birds. Systematic Zoology 26: 19-31. 

FORSHAWJ.M. 1987. Kingfishers and related birds. 
vol. 2. Alcedinidae: Hakyon to Tfinyxipk^m, 
(Lansdowne: Sydnev). 

FRY. C.U- 1980a. The origin of Afroimpieal kingllsh- 
crs. Ibis 122:57-72. 
1 980b. The evolutionary biology Of ki nglishers < Al- 
cedinidae). Living Bird !8* 1 13-160 

FRYX-H. FRY, K <fe HARRIS, A, 1992. Kingllshcrj, 
l>cc-e3iers i.utd njlleis, tCrooin Hehn; London). 

MOURER-CHALIRVIRE. M 1982 Les ojscau.N 
fn^silcN des Phosphorites du Quercy (Eocene 
Supi^ricur a OligocL^nc Supericur): implications 
paleohiog<5ogrnphiques. Geobios, mem. sp6c 6; 
413-426. 

OLSON, S.L. 1985. The fossil record of birds. I»p. 
79-238. hi Fanicr, D.S., Kmg. J.R. ik f^arkes,K.C. 
(eds) Avian biology, vol 8. (New York: Aw 
dcniic EYcss). 

SIBLEY, C.G & AHLQUIST.J.E. 1990. Phylogcny 
and chissirication of birds: A Study In molecular 
evolution, (Yale llniverMly Pres.';: New Haveni. 

STEADMAN. D W. 1980- A review ot the oslcoUigy 
and paleontology of turkeys (Aves; 
Melcagridinae).Coniribut]ons to Science from the 
Nauiral History Museum of Lxts Angeles Ciiunty 
330: 131^207. 

JEDFORD. R.H. 1967. Fos.sil mammals from the Carl 
Creek limesione, northwestern Queensland. Bul- 
letin otWx Bureau of Mineral Resources Geology 
and Ocophy-sicN 92: 2l7-23ft. 



234 MEMOIRS OF THE QUEENSLAND MUSEUM 



VICKERS-RICH, P. 1991. The Mesozoic and Tertiary VICKERS-RICH, P.V., MONAGHAN, J.M., BAIRD, 

history of birds on the Australian plate. Pp. 721- R.F. & RICH, T.H. (eds) (1991). Vertebrate pal- 

808. In Vickers-Rich, P.V., Monaghan, J.M., aeontology of Australasia. (Pioneer Design Stu- 

Baird, R.F. & Rich, T.H. (eds). Vertebrate pal- dio: Melbourne), 
aeontology of Australasia. (Pioneer Design Stu- 
dio: Melbourne). 



HINDLIMB PROPORTIONS AND LOCOMOTION OF EMUARIUS GIDJU 
(PATTERSON & RICH, 1987) (AVES: CASUARIIDAE) 

WALTER E. BOLES 

Boles, W.E. 1 997 06 30: Hindlimb proportions and locomotion of Emitarius gidjii (Patterson 
& Rich, 1987) (Aves: Casuariidae). Memoirs of the Queensland Museum 41(2): 235-240. 
Brisbane. ISSN 0079-8835. 

Using proximal and distal fragments, the length of the tarsometatarsus of Emuarius gidju is 
estimated and compared lo that of other hindlimb elements. From these proportions and other 
hindlimb morphology, the inferred locomoiory mode off", gidju is compared with Recent 
casuariids. Emuarius gidju appears to have been more cursorially adapted than Casuarius 
and dwarf Z)romrt /Hi', suggesting at least some open habitat in the Riversleigh palaeoenviron- 
ment. Using the relationship between weight and least circumference of the femur and 
tibiotarsus in Recent birds, the weight of E. gidju is suggested to have been 19-2 1 kg. □ 
Emuanus. Aves. hindlimb, locomotion. 

Walter E. Boles, Australian Museum, 6 College Street, Sydney NSW 2000, Australia; 
received 4 November 1996. 



Emus, Dromaius (Dromaiinae), form a promi- 
nent element of Australia's avifauna. The closely 
related cassowaries, Ccisiiarius (Casuariinae), are 
more restricted in distribution. These two groups 
occupy very different modern habitats, and loco- 
motory adaptations correlated with these differ- 
ent habitats are obvious in the relative pro- 
portions of the lower limb bones. Because of the 
relationship between habitat and liinb propor- 
tions, fossil emus and cassowaries are potentially 
goodpalaeoenvironmental indicators. Emus have 
a belter fossil record (Patterson & Rich, 1987). 
than cassowaries (Vickers-Rich, 1991). Patter- 
son & Rich ( 1 987) described lower limb elements 
from the Miocene of central Australian as a small 
emu, Dromaius gidju. Boles (1991) erected 
Emuarius for this species and considered it closer 
to emus than cassowaries, but nearer their dichot- 
omy than any other described laxon. 

The type material of Emuarius gidju is part of 
the Kutjamarpu Local Fauna, recovered from the 
Leaf Locality (UCMP V-6213) on the E shore of 
Lake Ngapakaldi in the E Lake Eyre Sub-Basin, 
South Australia. Much E. gidju material occurs 
at Riversleigh NW Queensland. Archer el al. 
(1989, 1994) considered the Riversleigh pal- 
aeohabiial as rainforest, based on mammal re- 
mains. 

This paper is to 1) estimate the lower limb 
proportions ofE. gidju\ 2) compare these to mod- 
ern emus and cassowaries and, by implication, to 
their style of locomotion; 3) make an initial esti- 
mate of the weight of £". gidju\ and 4) interpret the 
possible palaeohabitat at Riversleigh where E. 
gidju occurs. 



MATERIALS AND METHODS 

Osteological terminology follows Baumel & 
Wilmer (1993). Measurements were made with 
vernier calipers accurate lo 0.05mm and rounded 
lo the nearest 0.1mm. Institutional acronyms are 
AM (Australian Museum), QM (Queensland Mu- 
seum) and SAM (South Australian Museum). 

The type specimen of £. gidju (SAMP26779) 
is an associated distal tibiolarsal fragment, prox- 
imal tarsometatarsus including much most of the 
shaft, and complete set of pedal phalanges 
(Patterson & Rich, 1987) and numerous speci- 
mens of Emuarius are known from Riversleigh 
(Table 1). 

To compare changes in relative proportions of 

the casuariid hindlimb, the following measures 

were calculated for the bone lengths of the 3 

extant species of Casuarius and the 1 living and 

2 recently extinct dwarf species of Drotnaius: 

T^o-r^ Tibiotarsus X 100 

TBX4?MR - - 



TMXtMR = 



TBT/TMT 



Femur 
Tarsometatarsus x 100 

Femur 
Tibiotarsus x 100 



Tarsometatarsus 

Because the purpose was to find general, rather 
than detailed, directions of change, measure- 
ments were taken from the literature (Table 2) and 
rounded to the nearest mm. Means were used 
where available. The sample sizes were often 
small, sometimes comprising single individuals. 

Predicted body weight of Emuarius gidju was 
calculated (Campbell & Marcus, 1991) from AM 
F78585 (near complete femur) and QMF 16827 



236 



MEMOIRS OF THE QUEENSLAND MUSEUM 



(distal libiolarsal fragment). Tape was wrapped 
around the bones at their least circumferences, 
marked at the point of overlap, straightened and 
measured with calipers. These results were used 
in the equation logio(weighl)=a*logio(circumfer- 
ence)+b. where the values of a (slope) and b 
(inierecept) were those determined by Campbell 
& Marcus (1991) for all birds for the respective 
elements (femur: a=0.41 l,b=-0.065; libiotarsus: 
a=2.424,b=0.076). 

RESULTS 



Increased cursoriality in these birds is associ- 
ated with an increase in the lengths of the tibio- 
larsus and tarsometalarsus relative to that of the 
femur (Howell, 1944). Relative proportions of 
the hindlimb contributions of the long bones in 
Casuariits, Dromaius and Emuahus (Fig, 2) 
show that thai of the libiotarsus remains more or 
less consistent in all taxa; that of the femur de- 
creases with increased cursoriality, whereas that 
of the tarsometalarsus increases. The relationship 
between these changes is consistent for Recent 
species: TBT/FMR = 0.45 [TMT/FMR] +113; 
r=0.84. 

These changes appear independent of overall 
size when compared between genera, but within 
genera the smaller members have the greater 
TBT/TMT and smaller TMT/FMR. The 
TBT/FMR in Casuarius remains constant. It sug- 
gests from these figures that the smaller species 
in each genus are the leasi cursorial members. 

The Kutjamarpu femur (AMF78585; Boles, 
1991 ) oi'E. gidju is 194mm long, which, because 
of abrasion, is a few mm less than its original 
length, of about 198mm and very likely not 
200mm. Complete libiolarsi are unknown for E. 
gidju (Table 1). Nevertheless, it is possible to 
predict the size and relative proportions of this 
bone from other hind limb elements. Tar- 
sometatarsi are represented by the holotypical 
proximal end and shaft, and several distal frag- 
ments. An estimate of the tarsometatarsal length 
was made by using the proximal end and shaft and 
a distal fragment. The proximal tarsometatarsal 
fragment is 276mm long; the longest edge of its 
shaft is straight and shows no evidence of flaring 
outward to trochlea metatarsi II. The distal piece 
is 64mm long; the small portion of shaft remain- 
ing is just proximal to the flaring of trochlea 
metatarsi 11. Because there is little, if any, overlap 
between these two pieces, minimum length of the 
tarsometalarsus is 340mm (Fig. 3). 



TABLE I. Specimens that have been referred to 
Emuarius gidju. *=speciniens described by Boles 
(1991). 



SITE 


REG. NO. 


ELEMENT 


Lake Ngapakaldi 


SAM 

P26779 


Holoiype: associated 

distal tibiolarsus, 

proximal 

tarsometarsus. 
complete pes 


AMF78585* 


Femur 


Riversleigh 1 


System A or B 
While Hunter 


QMF16827'*= 


Tibiotarsus 


S\stem B 
Camel Spuiuni 


QMF29720 


Vertebrae 


QMF29721 


Vertebrae 


QMFI6828* 


Femur 


QMFI6829* 


Femur 


AMF78586* 


Tibiotarsus 


QMF29722 


Tibiotarsus 


QMF29723 


Tcu-so metatarsus 


QMF29724 


Tarsometalarsus 


OMF29725 


Tarsometalarsus 


QMF29726 


Tarsometalarsus 


QMF29727 


Tarsometalarsus 


Upper 


QMF16830* 


Rostrum 


QMFt683l* 


Scapulocoracoid 


QMF29728 


Vertebrae 


S>stem 7B, Dirks 
Towers 


QMF29729 


Tiirsomeiaiarsus 


System C 
Gag 


QMF16832* 


Femur 


AMF78587* 


Tarsometalarsus 



Using a tarsometatarsal length of 340mm and a 
femoral length of 198mm, gives a TMT/FMR o{ 
172, greater than that of any Recent casuariid 
except Dromaius novaehollandiae. Using these 
values with the regression equation for the family 
(Fig. 4) gives a predicted TBT/FMR of 190, 
which corresponds to a tibiotarsal length of 
376mm. These 3 values give a combined length 
of 916mm, virtually the same as the hindlimb of 
Casuarius casuarius, although the proportional 
contribution of each bone to this length is differ- 
ent (Fig. 2). 

This figure must be used with caution. The 
fragments on which it is based represent different 
individuals from different localities. As such, 
there are several sources of potentially significant 
variation between components. Geographical 
variation may not have been of major importance; 
living D. novaehollandiae exhibits little differen- 
tiation across its range. There is much greater 
size variation across this species' chronological 



HINDLIMB PROPORTIONS OF EMUARIUS GIDJU 



131 




FIG. 1. Leaf Locality femur oi' Emuarius gidju compared wilh femora ol 
Recent casuariids, in cranial view. From left, Casuarius Cositariits, C. 
bennettii, Emuarius gidju, Dromaius novaehollandiae and D. aler. 



range, with mainland fossils usually smaller than 
modern birds. Patterson & Rich (1987) suggested 
that it 'may have been at any one lime in the 
Pleistocene both larger or smaller than at 
present'. Considerable intraspecifie variation in 
living D. novaehollandiae is probably related to 
age and sex. as well as individual differences 
(Marchant & Higgins. 1990). For example, 
among 22 modern specimens, Patterson & Rich 
(1987) found a range in tarsometatarsal length of 
332-422mm (mean 383mm: s.d. 18.0). 

Miller ( 1 963) described Dromaius ocypus from 
the middle lo late Pliocene Palankarinna Local 
Fauna from Lake Palankarinna, northern South 
Australia. This species was intermediate in si/.e 
between Enuiarius gidju and living Dromaius 
novaehollandiae. The tarsometatarsus of D. oc- 
ypus is markedly shorter relative lo its width than 
is that of D. novaehollandiae, but less so than in 
Casuarius. Vickers-Rich (1991) inlerpreled this 
as suggesting a less cursorial lifestyle for /). 
ocypus than for D. novaehollandiae. Because 
Patterson & Rich ( 1 987 ) did not give comparative 
figures ft>r relative widths, it is difficult to quan- 
titatively compare E. gidju with these species. 
Visually, Emuarius appears to be proportionally 
thinner lor its length than is D. ocypus, but not lo 
the degree of D. novaehollandiae. 

Patterson & Rich (1987) pointed out that, al- 
though, the foot of E. gidju is more like thai oi' 
emus than cassowaries, it is more cassowary-like 



than any known species of emu 
(illustrated in Patterson & 
Rich, 1987). In emus, phalanx 
ungualis of digit III is longer 
than thai of digit II, whereas in 
cassowaries it is reversed, wilh 
phalanx ungualis of digit II ex- 
tended into a long spike several 
times the length of phalanx un- 
gualis o( digit III. Emuarius 
does not have phalanx ungualis 
of digit II developed into a 
spike, but il is still longer than 
thai of digit III. 

Cassowaries have digits II 
and IV relatively long com- 
pared wilh digit III (ratios oi 
11:111 and IV:III, respectively, 
without phalanges unguales 
0.82, 0.76). Both are substan- 
tially reduced in relative length 
in emus (ratios as above - 0.55, 
0.63). Both digits II and IV of 
Emuarius are comparatively 
longer than those in emus (digit IV lo a lesser 
degree), but not lo the extent seen in cassowaries 
(0.57, 0.68). Patterson & Rich (1987) suggested 
that the reduction of digit IV and particularly of 
digit II in E. gidju, compared wilh the highly 
cursorial D. novaehollandiae, appears lo parallel 
similar reductions in other groups of lerresirial 
birds and mammals. 

In comparison wilh the pedal phalanges of cas- 
sowaries, those of emus are dorsoplaniarly com- 
pressed. Emuarius is somewhat intermediate, 
with its phalanges substantially more 
dorsoplaniarly compressed than those of casso- 
waries, but less compressed than (but more sim- 
ilar lo) the condition in emus. 

Campbell & Marcus (1991) staled 1he least 
shaft circumference of either [femur or libiotar- 
sus] can be a reliable indicator of the weight of a 
fossil bird\ From their equation, and measure- 
ments off. gidju bones, predicted weights of this 
species were 21kg based on the femur and 19kg 
based on the tibiotarsus. The least circumference 
oi' the tibiotarsus is ahiiosl always at or distal lo 
the midpoint of the bone; in most birds il is in the 
distal third (Campbell & Marcus 1 99 1 ). No libio- 
tarsal specimen oi E. gidju is complete, although 
the length of that measured is about a third of the 
predicted length for this bone. Il is possible that 
the least circumference occurs on the missing 
section of libotarsus, and the value given here 
would have to be adjusted. The predicted 



238 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 2. Measurements (mm) and measures of relative proportions of 
hindlimb bones of Recent emus and cassowaries and of Emuarius gidju. 
For calculation of predicted measurements of E. gidju and of measures of 
relative proportions, see text. 



Species 


FMR 


TBT 


TMT 


Source 


TBT/ 
FMR 


TMT 
/FMR 


TBT/ II 
TMT 


Dromaius 
novaehoUandiae 


203 


401 


383 


Patterson & 
Rich. 1987 


198 


189 


105 


Dromaius 
baudinianus 


164 


293 


234 


Morgan & 
Sutton. 1928 


179 


143 


125 


Dromaius ater 


178 


331 


274 


Morgan & 
Sutton. 1928 


186 


154 


121 


C.asuanus 
unappendiculatus 


236 


388 


306 


Richetal.. 1988 


164 


130 


127 


Casuartus 
casuarius 


232 


384 


300 


Richetal., 1988 


166 


129 


128 


Casuarius bennetd 


221 


365 


229 


Richetal., 1988 


165 


104 


159 


Emuarius gidju 


198 


376 


340 


This paper 


190 


172 


111 



weights from the two bones are close enough to 
give an acceptable first estimate for E. gidju. 

DISCUSSION 



Throughout the early Tertiary, much of Aus- 
tralia was covered in closed forest, and the cli- 
mate was considerably more humid than at 
present (Frakes el al., 1987). The dominant veg- 
etation type over much of the continent was 
rainforest. Nothofagus-dom'iwdtQd rainforest 
covered the Lachlan River valley during the late 
Eocene to late Oligocene-early Miocene (Martin, 
1987). Even where closed forests were not pres- 
ent, gallery rainforest probably occurred along 



Casuarius 
bennetti 



watercourses. It was not until at 
least the middle Miocene (c. 15 
mya) that drying of the climate 
began and open habitats started 
to appear on a large scale. 
There is no evidence of grass- 
lands before the end of the late 
Miocene to Pliocene (Martin, 
this volume). 

The graviportal locomotion 
of Casuarius is associated 
with movement through the 
dense vegetation of the Aus- 
tralo-Papuan rainforests. In- 
creased cursoriality seems 
correlated with the appearance 
of more open habitat, in which 
sustained running could take 
place. (While able to run if 
required, cassowaries have 
limited opportunities in such habitat to work up 
and sustain a reasonable degree of speed because 
sufficiently open areas are restricted.) Morpho- 
logical correlates with cursoriality include pro- 
portional lengthening of the tibiotarsus and 
tarsometatarsus, and a reduction in the relative 
size of digit II. Conditions between the extremes 
of the states found in Casuarius and Dromaius 
are suggestive of levels of cursorial ability inter- 
mediate between theirs, although at what point 
along the scale cannot be determined. This, in 
turn, suggests the possibility that the amount of 
open habitat might be also somewhere between 
that available to cassowaries and emus. 



TAR.'^OMETATARSUS 


TIBIOTARSUS 


FEMUR 


III 










^^^^^M 



Casuarius 
casuarius 



Emuarius 
gidju 



Dromaius 
ater 



Dromaius 
novaehoUandiae 



FIG. 2. Comparative proportions of bones in the hindlimbs oi' Emuarius gidju, two species of Casuarius and two 
species of Dromaius. Note that while the tibiotarsal proportion remains relatively constant, the proportion 
comprising the femur decreases as that of the tarsometatarsus increases. 



HINDLIMB PROPORTIONS OF EMUARIUSGIDJU 



239 




locomotion (Prange el al 
resembled Casuarius. 



1979), more closely 



FIG. 3. 'Combined' length of Leaf Locality proximal 
tarsomctatarsus and Riversleigh distal tarsometatar- 
sus compared with tarsometatarsi of Recent 
casuariids, in cranial view. From left, Dromaius 
navaeliollancUae, D. ater. Emuarius gidju, Casuarius 
cosuarius and C. bennettH. 

Casuarius has been considered more primitive 
than Dromaius on the basis of distribution, habi- 
tat preferences and hindlimb. Schodde & Calaby 
( 1 972) and Schodde ( 1 982) cited the cassowaries 
as elements of the Tumbunan avifauna, which 
represents the earliest lineages of extant Aus- 
tralo-Papuan birds. The emus were placed by 
Schodde (1982) in the autochthonous Eyrean 
fauna, which evolved in response to the opening 
of the Australian habitat. Boles (1991) consid- 
ered Emuarius to be closer to the dichotomy of 
cassowaries and emus than any other known 
taxon. but too cursorially adapted and Dromaius- 
like to have been the common ancestor. It appears 
to mark in the Casuariidae a stage in the transition 
from a graviportal to a more cursorial locoino- 
lion. Although some characters of the hindlimb 
of £. gidju are more similar to either Casuarius 
or Dromaius, many are intermediate between liv- 
ing members of these genera (Boles, 1 99 1 ). Boles 
(1991) drew attention to the fact that the lower 
limb bones were more similar to those of 
Dromaius, whereas the femur, whose proportions 
are more dependent on the bird's mass than its 



There are alternative explanations that accom- 
modate both the cursorial hindlimb proportions 
of £. gidju and the absence of open spaces. One 
possibility is that the ground cover of the rain 
forests was sufficiently open for cursorial ani- 
mals to move rapidly without obstruction. For 
example, modern Nothofagus forests are fre- 
quently open below the canopy, without the dense 
understory of some other rain forest types (pers. 
obs.). 

Neville Pledge (pers. comm.) suggested that a 
situation may have existed similar to that which 
occurred on Kangaroo Island when the dwarf 
emu Dromaius baudinianus was extant. Much of 
the island's vegetation was very thick and would 
have prevented rapid passage of a large animal 
such as the emu. Large mammals, however, 
forced runways through the vegetation, permit- 
ting them to move with comparable, albeit re- 
stricted, ease. The emus apparently took 
advantage of these runways for their own prog- 
ress. Likewise, the large mammals known from 
Riversleigh may have opened similar pathways 
through the thick undergrowth, which could have 
been used by £". gidju. Nonetheless, D. 
baudinianus was also noted for being very fast 
and virtually uncapturable in open areas. 

A small Miocene dromaiid from Alcoota, NT, 
is known from 2 phalanges and 3 unassociated 
trochleae (Patterson & Rich, 1987). These are of 
comparable size with £". gidju. There are slight 
differences in phalangeal morphology, and 
Patterson & Rich (1987) retained these speci- 
mens as Dromaius sp. indct. until more complete 
material is available. The Alcoota palaeohabitat 
has been interpreted as a lake, bordered by sedge 
or grassland, grading to woodland and gully for- 
est (Murray & Megirian, 1 992). This is a different 
environment than that interpreted for the older 
Riversleigh deposits. Even if the Alcoota 
dromaiid proves to be E. gidju, it would have 
limited relevance to reconstructing the 
Riversleigh habitats because of the age differ- 
ences of the deposits. It would, however, suggest 
that E. gidju was preadapted for the more open 
Alcoota environment. 

Models of this species' locomotory mode de- 
pend on extrapolations from living, non-conge- 
neric relatives. These are speculative, and must 
be treated as such, while palaeoenvironmental 
reconstructions based on them require an even 
greater degree of caution. 



24flt 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FEMOROMETATAFSAL tNOE?t 

FIG. 4. Relationship of TMT/FMR and TBT/FMR o\' 

Recent species of Caxttarius (open circles) and 
DvomaUis fcloscd circlcs». Species numbered us lol- 
iows: I. Casnanu:s benne/it; 2. C. cusitanus, 3. C. 
iinuppeniJiciilatus: 4, FJ. haadmu^nis; 5, D. aier. 6, 
D.fioyaehoJIaniiiae, Regression line is TBT/I'"MK = 
0-45 ITMT/FMR|-(- 1 1 3; r=0,84.The fine vertical line 
represenis the inierseciiun ^^ith this line of the 
TMT/FMR ( 1 72) for Emiutrins i^iJjtt as calculated in 
ilie text. Pa^dicicd TBT/FMR is 190. 

ACKNOWLBDGEMENTS 

In addiiion lo acknowledgcmenis in Boles 
( 1991 ) I thank Neville Pledge for discussion on 
the Kangamo Island hahiiai, Lynne Ho and Mau- 
rice Ortega for figures, and Anna Gillespie for 
informaiii)n on E. gidju. The Riversleigh mate- 
rial was collected via an ARC Programme Grant 
lo M. Archer; a grajii from the Department of 
Arts, Sport, the Environineni. Tourism and Ter- 
ritories to M. Archer, S. Hand and H. Godthclp; 
support from the University of New South Wales: 
a grant from the National Estate Pnigrumme 
Grunts Scheme to M. Archer and A. Bartholomai; 
and grants in aid to the Riversleigh Research 
Projecl from Wang Australia Piy Ltd. ICl Aus- 
tralia and the Australian Geographic Society. 

REFERENCES 

ARCHER. M . HAND. S.J. & GODTHELP. H. 1994. 
Riversleigh. 2nded (Reed: Svdnevl- 

ARCHER. M.. HAND. S.. C}ObTHELP. H. & D 
MEGIRIAN. 1989- Fossil mammals of 
Riversleigh, norlhsvestem Queensland: prelimi- 
nary over% icw ol' hioslraligraphv. corretatinn and 
envimnmcnial change. Ausualian Zoologist 25: 
29-65. 

BAUMEL. J.J. & L.M. WfTMER. 1093. Osieologia. 
Puhlicaiions of the Nuttnll Oniitholncical Cluh 
23:45-132. 



BOLES, W-B- 1991. lie\\skm oWromoius gidju Patter- 
son & Rich. 1987. with a reassessment of its 
generic position. Natural History Museum of Los 
Angeles Cotmiy Science Series 36: I95-20S. 

CAMPBELL K.E.'jR. & MARCUS, L. K>7! Tlie 
relationship of hindlinib hone dimensions lo body 
weigh! in birds. Natural History Museum of b;>.s 
Angeles County Science Series 36: 395-41 2- 

FRAKES. LA.. McGOWRAN. B. & BOWLER. J.M. 
1987. Evolution of Australian environments. Pp. 
1-16. In Dyne. G.R. ^ Walton. DAV. (edsl. Fauna 
of Australia vol. lA. General articles. (Australian 
Government Publishing Service: Canberra). 

HOWELL. A.B. 1944. Speed in animals. Their special- 
ization for running and leaping. (Universiiy of 
Chicago Press: Chicago). 

MARCHANT. S. & HIGGINS. PJ. (coonJinalor^K 
1990. Handbook of Australian, New Zealand and 
AnUirciic birds, 1 . (Oxford University Press: Syd- 
ney). 

MARTIN. H.A. 1987.Cainozoic history of Ihc vegeta- 
tion and climate of the Lachlan River Region, New 
South Wales. Proceedings ol the Linnean. Society 
ofNew South Wales 19: 213-257. 

MILLER. A.H. 1963. Fossil raine birds of the late 
Tertiary of South Australia. Record.softhc South 
Australian Museum 14; 4J3 420. 

MORGAN. A.M. & SUTTON. 1 1928. A crilical de- 
scription of some recently discovered K^nes of the 
extinct Kangaroo Island Emu {Dromaius 
dit'tnenianus). Emu 28: 1-19 

MURRAY. P. Si MEGIRIAN, D. 1992. Continuity and 
conlntst in middle and late Miocene vcrtcbrulc 
conununiiies Trom ilie Nonhem Terriiory The 
Beagle 9: 915-218. 

PATTERSON, C, & P.V. RICH. l9S7.The iossii his- 
tory of ihc emus. Drotnaius (A\es: Daimaiinae). 
Records o\' the South Australian Museum 21: 
85-117. 

PRANGE. HD . ANDERSON. J.E & RAI IN. It. 1979. 
Scaling of skeletal mass to body mass in birds and 
mammals, American Naturalist 113: 103-122. 

SCUODDE, R i9S2, Origin, adaplalion and evolution 
of birds in arid Au.slralia. Pp. 191-224- In BrnkcT. 
W.R. & Grcenslade, PJ.M. (eds). Evolution of the 
flora .aid fauna of arid Auslralia. (Peacock Puhli- 
ealions: Frewville). 

SCHODDE, R. & CALABY, J.H. 1972. The biogeog- 
raphy of the Auslralo-Papuan bird and mammal 
faunas in relation to Torres Strait. Pp. 257-3(X). In 
Walker. D. (cl\X Bridge and harrier: the natural 
and cultural heritage of Tones Strait. tAustraiian 
Nalion;il University Press; Canberra). 

VICKERS-RICH. P- 199! Hie Mesozoic and Tertiary 
history of birds on the Australian plalc Pp. 721- 
SOS. In Vickers-Rich. PA^, Monaghan. J.M.. 
Baird. R.F. & Rich, T.H, (eds). Vertebrate pal- 
aeontology of Australasia. (Pioneer Design Stu- 
dio: Melbourne). 



RIVERSLEIGH BIRDS AS PALAEOENVIRONMENTAL INDICATORS 

WALTER E. BOLES 

Boles, W.E. 1997 06 30: Riversleigh birds as palaeoenvironmenlal indicators. Memoirs of 
the Queensland Museum 41(2): 241-246. Brisbane. ISSN 0079-8835. 

Fossilised birds from Riversleigh are used to make a palaeoenvironmenlal reconstruclion. 
Difficulties that hamper this attempt are discussed. From the early to early late Miocene 
deposits, a range of laxa demonstrate aquatic situations; four others are indicative or at least 
suggestive of rainforest; one hints at at least some open spaces; and six arc ambiguous 
because of insufficient morphological variation between taxa with different ecological 
preferences or insufficiently known palaeobiology. The only species thus far identified from 
Pliocene Rackham's Roost Site points to conditions similar to those at Riversleigh today. □ 
Riversleigh, birds, palaeoecology. 

Walter E. Boles. Australian Museum. 6 College Street, Sydney New South Wales 2000, 

Australia: received 4 November 1996. 



Modern birds are excellent habitat indicators. 
Potentially, the Tertiary avifauna of Riversleigh 
could .serve a valuable role in interpreting the 
palacoenvironment. Numerous bird remains have 
been recovered, bui few have been studied (Rich, 
1979; Boles, 1 99 1, 1993a, b,c. 1995, 1997a, b). 

Based primarily on the diverse mammal re- 
mains. Archer ei al. (1989) interpreted the vege- 
tation of Riversleigh during the early to early late 
Miocene as 'dense, species-rich gallery rain- 
Ibresis probably similar to those that persist today 
in mid montane New Guinea'. Archer ct al. 
(1994) concluded that the surrounding Pliocene 
habitat was 'a dry sclerophyll forest or woodland 
with a grassy understorey, probably not too un- 
like the environment that dominates Riversleigh 
today'. This paper reviews available information 
about Riversleigh birds as it might contribute to 
interpretations of Tertiary environments. 

Material (Tabic 1 ) is lodged in the Queensland 
Museum (QM) and Australian Museum (AM). 
The geology and geography of the Riversleigh 
deposits are available elsewhere (Archer et al., 
1989, 1994, 1995;Megirian, 1992). 

INTERPRETATION OF HABITATS: BASIC 
TENETS 

One of the most striking features of the 
Riversleigh avifauna is the large proportion of 
small terrestrial forms compared to other middle 
Tertiary sites in Australia, which are largely dom- 
inated by waterbirds and larger, flightless forms, 
both of which are also present at Riversleigh. 
Among modern Australian terrestrial (non- 
aqualic) birds, the prevailing pattern is consid- 
ered lo be that the more primitive (least 



specialised) members of lineages are found in 
montane and subtropical rainforest and contigu- 
ous wet forests, whereas the more derived laxa 
occur in open habitats. Schodde & Faith (1991) 
considered that *rainforest-inhabiting members, 
particularly in montane New Guinea and subtrop- 
ical Australia, represent ancestral forms from 
which those in scleromorphic vegetation have 
been derived'. These ancestral components have 
been recognised as the Tumbunan fauna 
(Schodde & Calaby, 1972; Schodde, 1982; 
Schodde & Faith, 1 99 1 ). The Tumbunan avifauna 
is now largely centred on 'a Nothofa^us-myr{\c- 
podocarp-dominaied forest of the type once 
widespread across Australia through the mid 
Tertiary' (Schodde, 1982). although some ele- 
ments of this fauna now extend well beyond this 
habitat. Although the Tumbunan-typc habitat is 
now mostly restricted lo higher elevations, a dis- 
iribulion thai is relictual, il was once more wide- 
spread through lower altitudes. Schodde & Faith 
( 1 99 1 ) suggested that 'the subtropical Tumbunan 
avifaunas now present in montane New Guinea 
were widespread in Australia' in the mid-Terti- 
ary. Thus the habitat of Riversleigh during the 
Miocene was probably similar to that retained in 
these present Tumbunan refuges. 

This apparent relationship between level of 
specialisation and habitat has been adopted as a 
basis for palaeoenvironmenlal interpretation. A 
fossil laxon is tentatively considered a likely 
rainforest inhabitant if it is primitive in its lin- 
eage. It may also be considered lo occur in 
rainforest if its affinities are to a group that is 
today confined lo rainforest. If all living species 
occur in a certain habitat then thai habitat is 
considered to most likely for the fossil form. 



mated hindlimb proportions of £. gidju with those 
of Recent Dromaiits and Casuarius. The tibiotar- 
sus and tarsometatarsus of E. gidju were rela- 



lower leg permits these birds lo capture prey from 
hollows and recesses inaccessible to other preda- 
tory birds. Modern species occur in a range of 



242 



MEMOIRS OF THE QUEENSLAND MUSEUM 



There seems little problem with freshwater 
aquatic birds. Within a family, there may be some 
variation in the type of aquatic habitats preferred, 
although the range of differences is substantial in 
only a few instances. Modern habitat preferences 
are used as an indication of the fossils' pal- 
acohabilats, unless otherwise indicated. 

Unless there is some outstanding morphologi- 
cal feature that signals a major shift in its biology, 
a fossil bird is considered to have similar ecolog- 
ical characteristics as its modem counterparts. 
Similarities in morphology between fossil and 
living forms are interpreted to share similar func- 
tions, unless there is evidence to the contrary. 

SYSTEMATIC LIST 

Dromornithidae 

This extinct family comprises 8 species in 5 
genera (Rich, 1979). These were large, Highiless 
birds with major, *ratite'-grade modifications to 
an entirely terrestrial lifestyle. The extent of these 
masks any relationships to other known orders of 
birds, although this family is no longer consid- 
ered closely allied to living ratites (Olson, 19S5). 
Barawertornis tedfordi and Bullockomis 
planeioccnrai Riversleigh as conspicuous faunal 
elements at some sites because of their abundance 
and size. Despite this, they are limited pal- 
aeoenvironment indicators because little is un- 
derstood of their biology. The 2 monotypic 
Riversleigh genera are among the least known. 

At Riversleigh, the 2 species occur together, 
often, but not always, with other large animals 
(e.g., D-Site). At several sites they occur with 
aquatic animals such as lungfish. turtles and croc- 
odiles. Whether this is an indication of a water- 
side association or a taphonomic artefact is not 
known. Nothing in the foot structure is obviously 
modified for entering water, nor has there been 
previous suggestion of an aquatic association. 

Relative proportions of the hindlimb bones can 
be a useful indication of locomolory mode. In 
fewdromornithid species, however, are complete 
specimens known for all major leg elements, and 
these rarely represent the same individual. Re- 
constructions must necessarily be based on the 
better known forms, particularly Genyornis new- 
toni Estimates of leg proportions are based on 
measurements given by Rich (1979). The 
hindlimb proportions of most dromornithids are 
very different from those of Casuarius or 
Dromaiiis, with only llhandornis lawsoni having 
proportions approaching those of living emus; 
Rich ( ] 980) and Vickers-Rich ( 1 99 1 ) considered 



this the most cursorial species. Where known, the 
tarsometatarsus of other species is short relative 
to the other long bones, more like the moas . 

The moas (Worthy, 1991) provide some sug- 
gestions about aspects of dromornithids' life- 
style. Most moas were forest dwellers, almost 
exclusively herbivorous. None seemed adapted 
to cursorial locomotion. Moa remains are often 
recovered in large numbers, indicating that these 
were gregarious birds. Many Australian drom- 
omithid sites yield large numbers of specimens, 
indicating aggregations. Such a concentration of 
animal biomass is analogous to moas and sug- 
gests that dromornithids were herbivorous 
(Vickers-Rich, 1991). 

The dromornithid bill was much heavier and 
deeper than the moas' (Olson, 1985, fig. 3: 
Vickers-Rich, 1991, pi. 4). The skull was larger 
and more robust, with scars indicating broad at- 
tachments for the jaw muscles (P. Vickers-Rich 
pers. comm.). Regardless of what dromornithids 
ale, they were equipped to handle more substan- 
tial food items than were moas. Their bills arc not 
hooked or otherwise suggestive of a predatator. 

Unlike moas, no remains of dromornithid food 
have been found. Large accumulations of 
dromornithid gastroliths(gi/y.ard stones) occur at 
Riversleigh (Archer et al., 1994:79) and other 
sites (Stirling & Zeitz. 1 900; Vickers-Rich. 1 99 1 ). 

By the late Miocene, both graviportal and cur- 
sorial species lived in northern Australia. This 
was taken to indicate both forest and open country 
by Rich & Baird (1986), who did not consider 
dromornithids to have been particularly success- 
ful in invading grasslands. The Riversleigh fossil 
material does not show cursorial modifications. 
It can be tentatively concluded that Barawer- 
rornis and Bullockornis were forest dwellers. 

Casuariidae - emus and cassowaries 

Emuarius Boles, 199! occurs at Riversleigh. 
Living emus and cassowaries Casuarius are quite 
different in their habitat preferences and locomo- 
lory styles. The latter is reflected in the hindlimb, 
suggesting potential for inferring palaeohabital 
from a comparison of £. gidju with living Ibrms. 

Patterson & Rich (1989) found thai the phalan- 
ges o{ E. gidju were between those oiDromaius 
novaehollandiae and Casuarius in morphology, 
although more similar to the former in relative 
lengths and in degree of dorsoplanar compres- 
sion. The fossil form D. ocypus Miller, 1963 had 
a tarsometatarsus that is markedly shorter relative 
to its width than that of A novaehollandiae, but 
longer than Casuarius. Vickers-Rich (1991) in- 



RIVERSLEIGH BIRDS AS PALAEOENVIRONMENTAL INDICATORS 



243 



TABLE 1. Bird families in Riversleigh Tertiary deposits and sites lively longer than in any other 

from which these have been recovered. A=Dromornithidae; casumid, except D. novae hoi landiae. 

B=Casuariidae;C=Phalacrocoracidae;D=Ciconiidae;E=Anatidae; The structure of the phalanges, rela- 

F^Accipitridae; G= Rallidae; H=Cacatuidae; l=Psittacidae^ live width of the tarsometatarsus and 

J=Apodidae^K=Halcyonidae; L= Passentormes (' =Menundae; - proportions of the hindlimb suggest 

=Oriohdae; - = Orthonychidae). f. ! r^ -,. . ,j? 

^ that £. gidju was more cursorial than 

Casuarius and may have approached 
the ability exhibited by D, 
novaehollandiae. E, gidju may have 
had habitat preferences resembling 
those of D. novaehollandiae, i.e., 
largely open country, although some 
rainforest types could possibly offer a 
sufficiently open understorey. 

Phalacrocoracidae - cormorants 

A distal carpometacarpal fragment 
comes from a large cormorant. Be- 
yond signaling a lacustrine situation, 
it is uninformative. 

Ciconiidae - storks 

Stork remains comprise 1 proximal 
and 2 distal tarsometatarsal frag- 
ments, 1 quadrate and a partial skull. 
The tarsometarsi do not belong to 
Ephipphiorhynchus, the only living 
genus in Australia, and are probably 
referable to Ciconia, a genus now 
found in Eurasia, South America and 
Africa. All living storks have associ- 
ations with shallow, slow moving 
water, although they are not restricted 
to aquatic habitats; they do not enter 
heavily forested areas. All eat small 
animals, including vertebrates, and 
some {Leptoptilos) consume carrion. 

Anatidae - waterfowl 

Several specimens have been allo- 
cated to this family but no further 
determination has been made. Sub- 
groups of living waterfowl have cir- 
cumscribed habitat preferences. The 

Riversleigh specimens indicate aquatic, probably 

lacustrine, situations. 

Accipitridae - diurnal birds of prey 

Pengana robertbolesi, a large bird of prey, with 
hyperflexible tarsal joint (Boles, 1993a) is con- 
vergent with the living Polyboroides (Africa) and 
Geranospiza (South America). Mobility of the 
lower leg permits these birds to capture prey from 
hollows and recesses inaccessible to other preda- 
tory birds. Modern species occur in a range of 



SITE 


A 


B 


c 


D 


E 


F 


G 


H 


1 


J 


K 


L 


SYSTEM A 


D-Site 


X 
























D-SITE EQUIVALENT 


Sticky Beak 


X 










X 














SYSTEM A OR B 


White Hunter 




X 




X 


X 


X 


X 










X 


SYSTEM B 


Camel Sputum 


X 


X 


X 








X 






X 




X 


Helicopter 


X 
























Ouiasiie 




















X 




X 


Panorama 
























X 


RSO 


X 














X 




X 




X 


Upper 


X 


X 




















x' 


Wayne's Wok 


X 




X 


















X 


Wayne's Wok II 
























X 


Neville's Garden 


X 






















X- 


SYSTEM '^B 


Bitesantennary 








X 


















Dirks Towers 




X 




















X 


Microsite 
























X 


SYSTEM C 1 


Archie's Absence 
























X 


Henk's Hollow 
























X 


Gag 




X 




















X 


Gotham 
























X 


Jim's Carousel 
























X 


Last Minute 






















X 


X^ 


Ringtail 












X 




X 








X 


Two Trees 
























X 


PLIOCENE 


Rackham's Roost 


















X 






A 



terpreted this to mean that D. ocypiis was less 
cursorial than D. novaehollandiae. Emuarius 
gidju approaches D. novaehollandiae in tarso- 
metatarsal !ength:width ratio more closely than 
does D. ocypus (Boles, this volume a). 

Increased cursoriality in the Casuariidae is 
characterised by an increase in the lengths of the 
tibiotarsus and tarsometatarsus relative to the 
fetnur. Boles (this volume a) compared the esti- 
mated hindlimb proportions of £". gidju with those 
of Recent Dromaius and Casuarius. The tibiotar- 
sus and tarsometatarsus of E. gidju were rela- 



There «:cm& liUic pnjWcnl Willi freshwater this ihcinoblcursorirt) species. Wbca'kjiMWU.lhe 



t. .« i.t^ 



244 



MEMOIRS OF'I HEQIJHHNSLAND MUSEUM 



hatiiUils and do not pcnnit any meaningful cxtiap- 
filarion lo Rivc^sieigh. A remnnil Ira^meiii olihis 
Tamily ij^ol comparable size lo Penqana, but can 
be refeTTCd only (enialively lo (his la\on. 

Rullidac -rails 

There is much rail material, rcp/escniinf mos! 
foielimb and hindlimb (.'lemenis, (rom several 
sites, bur mosi abundantly at White Hunier site, 
possibly from a single individual. All specimens 
represent a medium-si/ed rail about the size of 
living Cailinula tenebtusa. From the shape of the 
c;n'i>t^nteiacarpus and the relative sl^es o! Ihc 
wing and leg cicmenTs. ii appears to have been 
flightless, This rail is probably related lo the 
native-hens Trihtmy\, now usually merged as a 
subgenus o{Cnrllhu4lo. The native-hens comprise 
two living endemic Australian species, one of 
which is fjtghlless, and a flighdess Plcislocene 
species endemic to New Zealand Although 
largel) remaining in the vicinity of water, both 
Australian species freely enter adjacem open 
country. The llighdess Tasmanian wwrntv/i en- 
ters cultivated paddocks, and mainland vi'tirralis 
may move sonic distance from water m semiand 
and arid regions. Species of Galfinnhj arc gregar - 
ious, winch is consistent with the number of 
fossils found at sonic sites. The Rivcrslcigh rail 
indicates the proximtly of wetlands, but little else 
aboul the local environment. 

Cacatuidac - cockatoos 

A I'osirum has been a^ferred lo the modern 
while cockatoos, rafw;/«/( Boles, I993h), Wuhm 
Australia, these species occur from rainforest 
fringes through open foi-est and woodland lo of^ei^ 
arid country. The Riversleigh bifd is considered 
to have been similar to the gfoup of white cock- 
atoos with small bills and rounded, uncoloured 
crests, such as the corellas. These species exhibit 
a c\>nsiderablc range c^f habitat preferences, from 
central Australian arid zone (C*. pastmator) lo 
rainforesis on some islands e.g., Solomon Islands 
(C.dtu'orpsii). This range ofhabiiats occupied by 
modern species renders the Riversleigh sj^cimen 
of hllle value in palaeohahilal reconstruction. 

Psittacidae -parrois 

Two carpomelacarpi and a tarsometatarsus 
iami Rackham's Rtx^st come fnim the living 
Budgerigar Melnp.siftacHS undaUtm, This is a 
good indication (hat the Pliocene habitat al 
Riversleigh was open and lightly umbered. 
Today (his species occursi in \\vz arid, s<2iTii-arid 



and subhumid /ones, ineluding Riversleigh, but 
never far from water. 

Apodidac- swifts 

Humeri, a coracoid and tarsometatarsus of a 
medium-si/ed swifl are close lo a large species of 
swiftlel CoUocolia, the only genus that breeds in 
Australia at present. These species nest in caves, 
but. tlcspiie the number of apparent Riversleigh 
deposits originating from cave lloors, surpris- 
ingly few remains have been found at Riversleigh 
thus far. One of tlic lH)ncs is that of a young bird, 
strong indication that al least some level of local 
ba'cding was taking place. Swifts are no! gotxl 
environmental indicators, because they are ueriul 
feeders, capturing Hying insects above the hahiiai 
canopy, irrespective of \ihal that habitiU may be. 

llalcyonidae - forest kingllshers 

The single specimen available is assigned 10 the 
Halcyonidae, possibly close to TmlirfnupltHs 
(Boles, l^97bl and similar lo more primitive 
\i\'mghli\cyumds{Tan)\sip(eray^ft'!ilionl,Syfnn\ 
which are rainforest inhabiianls (bry. I^Wla.b). 
The Riversleigh fossil resembles what would be 
predicted for an early member of the 
Todimmphus lineage. .Au.stralian Todinwiphus 
occur outside rainforest, but species of the genu.s 
live in rainforests in olher parts of AusU'alasia. 

Passeri formes - songbirds 

These birds are good habitat indicators al spe- 
cific i>r generic level. Songbirds are known from 
aboul 100 specimens from al least 20 sites 
(Boles, 1 095 ); however, only '^ specimens thus far 
provide useful habitai indications. Onhonyx 
kakhwhweri, of which a feniur has been reported 
(Boles. I99.1c), belongs to a genus with the 1 
living spe<ries confined to rainforest of the east 
coast and New Guinea, occasionally entering 
dense bordering vegetation (e.g.. Uinmna). 
Green Waterholc Cave. SH South Australia, the 
site fot 0. hypsilop/ms, never had rainforest, but 
there is evidence for a thick cover of 
Leptospvnnum ( Baird, 1 9S5'), which would prob- 
ably have provided adequate cv»ver. 

The lyiebini Menttra fyawanottles is repiie- 
senled by acarpomelacarpus (Boles. 19*J5). The 
two living species occupy rainforest and* in the 
case of M. novavhollaudiue, coniiguims fatrsi. 
The habitat preferences of the Riversleigh spccieii 
of Orthonyx and Metiura are assumed lo be sim- 
ilar to those of iheir modern congeners. 

A large lt»wer mandible from Neville's Garden 
Site (unpubl. daia) is from the Oriolidac which 



RIVERSLEIGH BIRDS AS PALAEOENVIRONMENTAL INDICATORS 



245 



includes the frugivorous forest birds Oriolus and 
Sphccotheres vindis\ many occur in closed forest. 
The fossil suggests rainforest, but is not diagnos- 
tic. 

TAPHONOMY 



that fossil birds had similar ecological character- 
istics to living counterparts and within a lineage 
more primitive species occur in rainforests, more 
derived ones in more open habitats. 

PALAEOENVIRONMENT 



None of the aquatic or semi-aquatic species, 
except the Gallinulo-iypc rail, show any evidence 
of unusual causes of death or accumulating 
agents. The rails occur in greater numbers at a 
handful ofsites, possibly due to a gregarious habit 
rather than taphonomy. Dromornithid remains 
being very common at some sites may also be a 
result of gregariousness or their corpses could 
have been accumulated during flooding. They are 
often found with large aquatic laxa (e.g., lungfish, 
turtles, crocodiles), and these may represent 
thanatocoenoses. Most other terrestrial species 
are too infrequent to provide any clues, but may 
be best considered chance survival of the remains 
of animals that died for a variety of reasons. 

Exceptions are the small terrestrial forms, par- 
ticularly passerines, which appear to have been 
killed by ghost bats {Macroderma). The living M. 
gigas is a predator of small vertebrates, which it 
captures from the ground or perch. Prey are eaten 
in the roost caves by chewing through the pectoral 
region, manifested in the skeleton as extreme 
damage to the sternum (Boles unpubl. data); dis- 
tal wings and legs are discarded. This is consis- 
tent with the elements that predominate at former 
Macrodenna-iXCCumuViXiQd sites at Riversleigh 
(carpo metacarpus, libiolarsus, larsomeialarsus). 
Unlike northern Australia today, where only a 
single species exists, Riversleigh is known to 
have supported many species of ghost bats during 
the Tertiary (Hand in press). Several sites have 
been identilled as the remnants o^ Macroderma 
roost caves, such as the Miocene Gotham Site and 
the Pliocene Rackham's Roost Site. 

PROBLEMS IN INTERPRETATION 

There are several reasons why the Riversleigh 
birds do not offer the same depth of environmen- 
tal data as living forms. Many specimens are yet 
to be studied. Ordinal level ideniificaiion (Table 
1) is frequently not fine enough for meaningful 
habitat inferences, especially for terrestrial spe- 
cies. There may be insufficient morphological 
differences between related forms occupying dif- 
ferent habitats. Much of the biology of extinct 
groups remain unknown. Basic assumptions 
could be wrong; primary ones employed here are 



The birds show that both aquatic and terrestrial 
habitats were prominent through the early to mid 
Miocene at Riversleigh. Because of the broad 
spectrum of wetland situations in which they 
occur, the cormorant, ducks and rail do not pro- 
vide any clues to the detailed nature of these 
systems. Based on modern habitat preferences, 
the stork indicates shallow, slow moving, lacus- 
trine situations somewhere in the area. 

Several of the better taxonomically resolved 
specimens are consistent with a closed forest. The 
passerines Orthonyx and Menura belong to fam- 
ilies which are today almost exclusively re- 
stricted to rainforest. Living halcyonid king 
fishers occur through most Australian habitats; 
the Riversleigh form, however, is suggestive of 
more primitive, rainforest-inhabiting laxa. 

Possible support for a more open habitat in 
some places comes from Emuarius. Its hindlimb 
proportions are thought to resemble those of 
Dromaius novaehollandiae , a highly cursorial 
animal. If the assumption can be made that the 
similarities in morphological proportions of the 
hindlimbs reflect similarities in function, this im- 
plies an advanced level of cursoriality in 
Emuarius as well. 

The Pliocene environment of Riversleigh, 
based on Rackham's Roost site, was quite differ- 
ent from the Miocene. The Pliocene fossils are all 
small forms, mostly passerine but including a 
small extant parrot, Melopsittacus undidams. 
This species is widespread, in arid and semiarid 
woodlands and scrublands, including the 
Riversleigh area. It requires proximity of water. 
Thus it suggests that Riversleigh in the Pliocene 
was probably very much like it is today. 

The Riversleigh early to mid Miocene environ- 
ment probably included shallow water at a num- 
ber of different sites, with some surrounding 
rainforest and some more open forrests. 

ACKNOWLEDGEMENTS 

Comparative material has been made available 
by the curatorial staff of the Australian National 
Wildlife Collection. Museum of Victoria, 
Queensland Museum, South Australian Museum, 
United Slates National Museum and University 



246 



MEMOIRS OF THE QUEENSLAND MUSEUM 



of Kansas Museum of Natural History. Valuable 
discussions were provided by R.F. Baird, the late 
G.F. van Tets, P. Vickers-Rich, N. Pledge, M. 
Archer and A. Gillespie. Support came from an 
ARC Grant; the University of New South Wales; 
Department of Arts, Sport, the Environment, 
Tourism and Territories; the National Estate 
Grants Scheme; Wang Australia; ICI Australia 
and the Australian Geographic Society. 

LITERATURE CITED 

ARCHER, M.. HAND, SJ. & GODTHELP. H. 1994. 
Rivcrsleigh 2nd edition. (Reed: Sydney). 

1995. Tertiary environmental and bioiic change in 
Australia. Pp. 77-90. In Vrba. E.S., Denton, G.H., 
Partridge, T.C. & Burkle, L.H. (eds), Paleocli- 
male and evolution, with emphasis on human 
origins. (Yale University Press: New Haven). 
ARCHER, M., HAND. S.. GODTHELP, H. & MEGIR- 
lAN, D. 1989. Fossil mammals of Riverslcigh, 
northwestern Queensland: preliminary overview 
ofbiostratigraphy, correlation and environmental 
change. Australian Zoologist 25: 29-65. 
BAIRD, R.F. 1985. Avian Ibssils from Quaternary de- 
posits in 'Green Waterhole Cave', south-eastern 
South Australia. Records of the Australian Mu- 
seum 37: 353-370. 
BOLES. VV.E. 1 99 ! . Revision oWromaius gidju Pauer- 
son and Rich, 1987, with a reassessment of its 
generic position. In Campbell, K.E., Jr (ed.), Pa- 
pers in avian paleontology honoring Pierce 
Brodkorb. Natural Hi.story Museum Los Angeles 
County Science Series 36: 195-208 

1 993a. Pengcma rohertbolesi, a peculiar bird of prey 
from the Tertiary of Riversleigh, northwestem 
Queensland, Australia. Alcheringa 17: 19-25. 

1993b. A cockatoo (Aves: Psitlaciformes: 
Cacatuidae) Irom the Tertiary of Riversleigh, 
northwestem Queensland, Australia, with com- 
ments on the value of the rostrum to the syslem- 
atics of parrots, ibis 135: 8-18. 

1993c. A logrunner Orthonyx from the Miocene of 
Riversleigh, northwestern Queensland. Emu 93: 
44-49. 

1995. A preliminary analysis of the Passeriformes 
Ibrm Riversleigh, northwestern Queensland, 
Australia, with the description of a new species 
of lyrebird. Courier Forschungcn-Inslitut 
Senckenbcrg 181: 163-170. 

i997a. Hindlimb proportions and locomotion of 
Emuarius gidjtt (Patterson and Rich. 1987) 
(Aves: Casuariidae).Memoirsof the Queensland 
Museum 41: 235-240. 

1997b. A kingfisher (Halcyonidae) from the 
Miocene of Riversleigh, northwestem Queens- 
land, with comments on the evolution o\' king- 
fishers in Australo-Papua. Memoirs of the 
Queensland Museum 41 : 229-234. 



FRY, C.H. 1 980a. The origin of Afrotropical kingfish- 
ers. Ibis 122:57-72. 
1 980b. The evolutionary biology of kingfishers ( Al- 
cedinidae). Living Bird 18:1 13-160. 

HAND, S.J. In press. Macrodenna mahigam, a new 
Miocene megadermatid (Microchiroptera) from 
Australia, with broader comments on 
megadermatid evolution. Geobios. 

MEGIRIAN. D. 1992. Interpretation of the Miocene 
Carl Creek Limestone, northwestern Queensland. 
The Beagle 9: 219-248. 

OLSON, S.L. 1985. The fossil record of birds. Pp. 
79-238. In Fiimer, D.S., King, J. R. & Parkes, K.C. 
(eds), Avian biology, vol. 8. (Academic Press: 
New York). 

PATTERSON, C. & RV. RICH. 1987. The fossil his- 
toid of the emus, Dromaiits (Aves: Dromaiinae). 
Records of the South Australian Museum 21: 
85-117. 

RICH, P.V. 1979. The Dromomithidae. an extinct fam- 
ily of large ground birds endemic to Australia. 
Bulletin of the Bureau of Mineral Resources Ge- 
ology & Geophysics- Australia 184: 1-196. 
1980. The Australian Dromomithidae: a group of 
extinct large raiites. Natural History Museum of 
Los Angeles County. Contributions to Science 
330:93-103. 

RICH, P.V. & BAIRD, R.F. 1 986. History of Australian 
avifauna. Current Ornithology 4: 97-139. 

SCHODDE. R. 1982. Origm. adaptation and evolution 
of birds in arid Australia. Pp. 191-224. In Barker, 
W.R. (feGreenslade, P.J.M. (eds), Evolution of the 
Horaand fauna of Eirid Australia. (Peacock Publi- 
cations: Frewville). 

SCHODDE, R. & CALABY, J. 1972. The biogeogra- 
phy o\' the Australo-Papuan bird and mammal 
faunas in relation to Torres Strait. Pp. 257-3(30. In 
Walker, D. (ed) "Bridge and Biirrier: The natural 
and cultural history of Torres Strait.'. (Australian 
National University Press; Canberra). 

SCHODDE. R. & FAITH, D.P. 1 99 1 . The development 
of modern avifaunulas. Pp. 404-412. In Bellet al. 
(eds). Acta XX Congressus Intemaiionalis Or- 
nithologici. (New Zealand Ornithological Con- 
gress Trust Board: Wellington), 

STIRLING, EC. & ZEITZ, A.H.C. !9(K). Fossil re- 
mams of Lake Caiiabonna. I. Genyomis newto/ii. 
A new genus and species of fossil slmlhious bird. 
Memoirs of the Royal Society of South Australia 
1:41-80. 

VICKERS-RICH. P. 1991 . The Meso/oic and Tertiary 
history of birds on the Australian plate. Pp. 721- 
808. In Vickers-Rich, P.V.. Monaghan, J.M., 
Baird, R.F. &. Rich, T.H. (eds). Vertebrate pal- 
aeontology of Australasia. (Pioneer Design Stu- 
dio: Melbourne). 

WORTHY, T.H. 1991. An overview of the taxonomy, 
fossil history, biology and extinction of moas. Pp. 
555-562. In Bell ei al. (eds). Acta XX Congressus 
Intemaiionalis Omithologici. (New Zealand Orni- 
thological Congress Taist Board: Wellington). 



A NEW OLIGOCENE-MIOCENK SPMCIES OF BURRAMYS (MARSUPIALIA, 
BURRAMYIDAE) FROM RIVERSLEIGH. NORTHWESTERN QUEENSLAND 

J. BRAMMALL AND M ARCHER 

Brammall. J. &. Archer, Ni. 1997 06 30: A new Oligoccne-Mioccne species of BurtWW 
iMarsupiah;*, Burrumyidae) from Rivcrslciah, norlhvvcslcm Queensland. Memaitx of thr 
Qneenskmii Museum 41(2): 247-268. Brishane- ISSN 1)07^-8835 

Bunwnvs is abiindanl in Ihe Oligocenc-Miocene ai River^lciph. nonhwestcrn Qticensiiind. 
Bitrrainys bntlyi sp. nuv. ly rcprescnied by over 150 denlarv and maxillary Irai-'iiifius and 
isolated icelh from 22 siles. 5»rni/m'^ appears to be morphological ly conservaiive, with luily 
minor meirical variaiicm between specimens of i?. bnayi from differeni sites and relatively 
few features distinguishing Miocene. Pliocene and Recent species. Phylogenelic aruilyses 
suggest that B. hruhi is the plesiomorphic sisler-group to all other species offiurmmys, with 
B w'akrfifldi sister-groiip \o the clade comprising B. triradiotm and B. /mrxHis. \3 
Burratnyiihw. Burnimys bnayi. h'ivenlei^lt. Oltgocene. Mim'vne. 

J. Brammall <fi M. Ardtrr, School of Biolojiical Sctefice, Utih'erSity^ of^ew Sottfh W<tU\%. 
New South Walet 2052. Austnilu^; n'Cdi*ed4 November 1996. 



Hurramys was represented only by Pleistocene 
fossils of /?. panus from Wombcyan Caves, 
NSW (Broom, IK96) and Pyramids Cave. Viclo- 
ria(Wakenekl, l^)6())uniil 1966 when the Moun- 
tain Pygmy-p<\ssimi. B. parvna, was disco\'ered 
alive at Mount Hotham. Victoria (Anon., 1%6; 
VVarneke, 1967). Two more fossil species oi' 
Htirmmys have been identified; early Pliocene H. 
thradiatus from Hamilton, Victoria (Turnhull el 
al.. 1987) and B. waUfieUU from late Oligocene 
(Woodburne el a).. 1993i Ngama Local Fauna. 
South Australia (Pledge, 1987k Discovers' of 
Miocene Burnimys al Riverslei^'h extends the 
geographic ran^e lar north and provides the first 
si /cable Tertiary sampled 50spccimcns). A met- 
ric analysis of this sample aims to determine taxa 
present and to assess variation Biirntntys hru(yi 
sp. nov. is used as the basis lur an evalualion of 
iniragcncric phyloj:cneiics of Hitrnitfiw 

Dental hoitiology lullows Flower (IS67) for 
premolar numbering and Lucketl (1993) for pw- 
Oiotar/molar boundary and molar number. Tooth 
po.sitions given without .super- or subscript num- 
liets refer to both upper and low er teeth: thus M"^ 
and M4 are individual teeth but M4 refers to both. 
Molar cusp nomenclature follows Archer ( 1984) 
not Pledge ( 1 987 ). Pledge's paraconid is our pro- 
loconid; his protoconid is no! recogni.scd. 

Higher systematic ntMnencIaiure foliowj. Aplin 
Si Archer (1987). Svsieni nomenclature is based 
on Archer el al. (1989) and Greaser ( 1997). Ma- 
teria! retcnred to is housed in the Queensland 
Museum, Brisbane fQMF) or Museum of Viclo- 
tia. Melbourne, (NMVP) Measurements in 
millimetres (mm I are to the nearest 0.01 mm using 
a Wild MMS235 Di^jiial Leni^ih-Measurin^ Set 



attached to a Wild M5A stcrcomicroscope. Molar 
lengths and widths and molar row lengths were 
measured as the maximum dimensions of the 
enamel-covered crown(s) w^ith the teeth in occlu- 
sal view, with lengths taken along ihe an- 
teroposterior axis of the tooth and v\idihs 
measured perpendicular to that axis. For Py in 
dorsal view and P^ in ventral view, maximum 
length was measured parallel to the apical blade 
edge, and anterior, posterior and maximunt 
widths were measured perpendicular to the blade 
edge; buccal and lingual heights were measured 
from the ba.se of the enamel al the saddle between 
the roots, to Ihe median apical edge, parallel to 
the posterior edge of the tooth. Slalislical analy- 
ses were performed using SYSTAT and Kaleida- 
Graph data analysis and graphics applications. 

METRIC ANALYSIS 

DcNpile overall uniformity. Riversleigh 
Burramys material shows some variation in abla- 
tive and absolute premolar and molar si/es. Met- 
ric analysis of dental features allempted to 
identity patterns which might inilicaie se.xuai di- 
morphism, specific or subspecific separation or 
differenliaiionofpopulaiions from different sites. 
Univariate and bivariate distributions and princi- 
pal components analysis were employed. 

Cheektooth dimensions (Table 1) (or Recetn 
H parvus populatu^ns refer to left dcniiiioii ex- 
cept where the right dentition was more complete. 
Standard error iSE) ts u.scd rather than standard 
deviation (SD) because it better indicates reliabil- 
ity of the mean estimate. The cocfricicni of vjiri- 
aiion (CV )= SD divided by mean v 100. 



248 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE L Cheektooth dimensions of Bitrramys species. ResuUs given as: Mean ± Standard Error (No. 
Specimens) Coefficient of Variation (CV%). CV not given where n 2. L = length, AW = anterior width, PW 
= posterior width, MW = maximum width, LH = lingual height, BH = buccal height. 





Riversleigh 


B. wakefieldi 


B. triradiaius 


B. parvus 






Burramys 














Lower teeth 




CV 


CV 




CV 




CV 


P3L 


1.81 ±0.01 (38) 


4.00 


1.64 (1) 


2.58 ±0.05 (4) 


3.68 


2.17 ±0.01 (21) 


1.96 


P3AW 


1.03 ±0.01 (38) 


8.21 


0.86 (1) 


1.04 ±0.01 (4) 


1.85 


0.85 ±0.02 (21) 10.51 1 


P3PW 


1.22 ±0.01 (38) 


5.07 


1.22 (1) 


1.49 ±0.06 (4) 


7.96 


1.32 ±0.01 (21) 


4.11 


P3MW 


1.27 ±0.01 (29) 


5.58 


1.22 (1) 


1.67 ±0.06 (4) 


6.61 


1.39 ±0.02 (21) 


4.79 


P3LH 


1.44 ±0.01 (37) 


5.25 


1.29 (1) 


2.02 ±0.04 (4) 


3.73 


1.92 ±0.01 (19) 


3.23 


P3BH 


1.73 ±0.02 (37) 


5.64 


1.51 (1) 


2.44 ±0.05 (4) 


3.98 


2.22 ±0.02 (20) 


3.05 


Ml L 


1.24 iO.Ol (32) 


3.71 


1-21 (1) 


- 




1.57 ±0.01 (21) 


2.02 


Ml AW 


0.78 ±0.01 (32) 


7.89 


0.86 (1) 


- 




1.00 ±0.01 (21) 


4.23 


MiPW 


0.95 ±0.01 (32) 


5.80 


0.97 (1) 


- 




1.25 ±0.01 (21) 


3.60 


M2L 


1.09 ±0.01 (32) 


3.72 


- 


1.55 (1) 




1.57 ±0.01 (21) 


1.97 


M2 AW 


0.88 ±0.01(34) 


5.55 


- 


1.23 (1) 




1.21 ±0.01 (21) 


2.03 


M2PW 


0.96 ±0.01 (34) 


5.67 


- 


1.32 (1) 




1.32 ±0.01 (21) 


1.90 


M3L 


0.93 ±0.02 (10) 


6.48 


- 


1.32 ±0.04 (2) 


4.29 


1.23 ±0.01 (19) 


1.84 


M3 AW 


0.84 ±0.01 (10) 


4.46 


- 


1.13 ±0.00 (2) 


0.00 


1.06 ±0.01 (19) 


2.25 


M3PW 


0.85 ±0.02 (10) 


6.13 


- 


1.17 ±0.01 (2) 


0.61 


1.07 ±0.01 (19) 


2.76 


M4L 


0.66 (1) 




- 


- 




0.68 ±0.01 (14) 


5.88 


M4 AW 


0.64 (1) 




- 


- 




0.66 ±0.01 (14) 


6.83 


M4PW 


0.50 (1) 




- 


- 




0.52 ±0.01 (14) 


7.82 


Mi_2 


2.30 ±0.02 (27) 


3.37 


- 


- 




3.13 ±0.01 (21) 


1.96 


M1.3 


3.24 ±0.05 (8) 


3.91 


- 


- 




4.34 ±0.01 (19) 


1.34 


M1.4 


3.83 (1) 




- 


- 




4.93 ±0.02 (14) 


1.29 


Uvfer teeth 
















2.01 ±0.02 (17) 


4.89 


- 


2.59 ±0.02 (2) 


1.09 


2.27 ±0.01 (19) 


2.60 


P^AW 


0.93 ±0.01 (17) 


4.32 


- 


0.91 ±0.02 (2) 


2.32 


0.75 ±0.02 (19) 10.39 1 


p3pw 


1.20 ±0.01 (17) 


4.93 


- 


1.63 ±0.05 (2) 


4.79 


1.13 ±0.01 (19) 


3.82 


P^MW 


1.25 ±0.01 (17) 


4.41 


- 


1.63 ±0.05 (2) 


4.79 


1.24 ±0.01 (19) 


2.44 


P^LH 


1.58 ±0.02 (17) 


4.67 


- 


2.32iO.02(2) 


1.22 


1.92 ±0.01 (16) 


2.76 


P^BH 


1.65 ±0.02 (17) 


4.44 


- 


2.16 ±0.01 (2) 


0.98 


2.06 ±0.02 (18) 


3.08 


m"^ L 


1.12 ±0.02 (14) 


4.86 


- 


- 




1.51 ±0.01 (19) 


1.94 


m"^ aw 


1.16 ±0.02 (14) 


5.25 


- 


- 




1.40 ±0.02 (19) 


7.08 


m'^pw 


1.17 ±0.01 (14) 


4.14 


- 


- 




1.45 ±0.01 (19) 


4.01 


M^ MW 


1.39 ±0.01 (14) 


3.90 


- 


- 




1.68 ±0.01 (18) 


2.77 


M^L 


0.98 ±0.01 (8) 


3.45 


- 


1.22 (1) 




1.45 ±0.01 (19) 


1.53 


M^ AW 


1.16 ±0.01 (8) 


2.84 


- 


1.34 (1) 




1.56 ±0.01 (19) 


2.20 


M^PW 


0.93 ±0.01 (8) 


2.40 


- 


1.10 (1) 




1.27 ±0.01 (19) 


3.37 


M-'^L 


0.86 ±0.03 (3) 


6.43 


_ 


- 




1.09 ±0.02 (18) 


8.17 


M-^ AW 


0.93 iO.02 (3) 


2.84 




- 




1.19 ±0.03 (18) 10.83 


M^PW 


0.72 ±0.03 (3) 


7.86 


- 


- 




0.88 ±0.02 (18) 10.71 


M^L 


0.67 ±0.02 (3) 


5.68 


- 


- 




0.77 ±0.01 (13) 


4.86 


M^ AW 


0.71 ±0.02 (3) 


4.97 


- 


- 




0.74 ±0.02 (13) 


8.81 


M^PW 


0.46 ±0.03 (3) 


10.80 


- 


- 




0.51 ±0.01 (13) 


9.49 


M^-2 


2.11 ±0.03 (8) 


3.40 


- 


- 




2.96 ±0.01 (19) 


1.85 


M^-3 


2.98 ±0.08 (3) 


4.55 


- 


- 




4.07 ±0.02 (18) 


1.63 


M^-4 


3.55 ±0.03 (3) 


1.33 


- 


- 




4.77 ±0.03 (13) 


1.86 



CV is less than 1 1 throughout and usually less 
than 6 (Table 1 ). Following Simpson ct al ( 1 960), 
this degree of variation indicates an unmixed 
sample, although Gingerich (1974) cautions 
against uncritical application of this absolute CV 
criterion and recommends greater emphasis on 
relative variabilities of different teeth. In approx- 



imately 80% of measurements B. parvus has a 
lower CV than the Riversleigh sample, but the 
interspecific differences in CV are generally not 
great. CVs for B. triradiatiis fall within approxi- 
mately the sanie ranges as those for the 
Riversleigh and Recent specimens, but arc de- 
rived from very few specimens and are therefore 



NEW OLIGOCnNE-MIOCENE BURRAMYSmOM RIVERSLEIGH 



24y 




5 


B 


u 




c 




1 ' 




i ' 




"fl - 




. 2 




o 


•.li^^^BM 


z 


^li^H 


1 





va2 1.26 
Ml length 



1,34 1 40 





1.00 



1.08 1.12 
Mg length 



1.16 1.20 




1.60 1.90 
P3 length 



1.12 1.19 1-2G 1.33 

P3 posterior width 



1.40 




2,23 2.31 2.39 
M] -2 length 



47 2.55 




P3 buccal height 



69 2.00 



System A 



System B 



= System C 



FIG. I. Frequency hisiogranis lor sonic lower looih mcu.surcmcnts of Rivcrslcigh Burnwiys specimen.^ All 
nicasuremenis inmm. 



nol considered reliable. Total variation (as indi- 
cated hyCVs) suggests 1 species oI7?//mmm' in 
the Rivcrsleigh sample spanning greater vuriulion 
than the sample oF Recent B. paivus. 

Where variation between taxa is small (as ks 
likely with small-bodied ia\a), it may be ob- 
scured by cpigenetic morphological variation, by 
looih wear or by nieasureineni error: metric diC- 
Terences between closely related taKa iu-e most 
likely lo be delected by examining structures with 
the lowest levels o\' such varialitm. M2, in the 
centre of the P^ -M4 tooth row. is in that sense the 
most functionally integrated oi' these teeth; it may 
therefore be expected to be least variable 
(Gingerich, 1974). vSimilarly. total molar row 
lengths may he more tighil> conirolled than the 
lengths of individual molars. M: dimensions and 
molarrow measurements (including pariial molar 
row measurements such as M1.2 length I are gen- 
erally the least variable measurements in B. 
parvus and the Riversleigh .sample: P3 length is 
also relatively constant (Table 1 ). Thus analysts 
ol the Riversleigh sample was focused on P3 and 
Mio, although all other measurements were ex- 
amined. 

Frequency histograms for some measurements 
are bimodal, while others are either unimodal i>r 
perhaps incipientiy bimodal. Mi-> length (Fig. 



IC), with CV=3.37 is bimodal. M^ length (Fig. 
IB; CV=3.72) and P3 buccal height (Fig. IF; 
CV=5.64) are considered bimodal. though nol 
with certainty. Mi lengtli (Fig. lA; CV=3-7h 
may represent a bimodal distribution but could 
equally be a sample from a unimodal. normal 
distribution; P3 length (Fig. 1 D; CV=4.00) and P3 
posterior width (Fig. I E; CV=3.n7) dislriliutions 
could each Ixr described either as havmg 2 or 3 
peaks, or as representing single normal distribu- 
tions. Kolmogorov-Smirnov Lilliefors lesis indi- 
cale that some of the univariate distributions 
differ signillcanlly trom normal (Table 2) and 
comparison with Table 1 shows that these include 
several with low variation. Thus univariate fre- 
quency distributions hint that the sample repre- 
sents more than one population, but do not 
provide a basis for subdivision. 

Bivariate plots (Fig. 2) suggest no clear divi- 
sions other than those evident in the univariate 
di.stribuiions. such as the apparent bimodalily of 
M2 length (Fig. IB, 2B)- They show that speci- 
mens from Systems B and C have ovcriapping 
distributions, but that for some measurements, 
specimens from System C sites are, on average, 
smaller than specimens from System B sites. This 
is .so for Ml length and P3 length iFig. 2A. 2C, 
2D) and lo a lesser extent forM: length (Fig. 2B). 



250 



MEMOIRS OF THE QUEENSLAND MUSEUM 



MO 

1.05 

I 1.00 

I 0.95 

<i> 

(/) 

g. 0.90 

0.85 

0.80 
1 

2.00 
1.90 



J3 



• •• 



8^ 

m- 



■B < 



n 




10 1.15 1.20 1.25 1.30 1.35 1.40 
Ml length 



1.02 1.04 1.06 1.08 1.10 1.12 1.14 1.16 1.18 
Mj length 



1.60- 



1.50 



T:- H-* 

- :...#...: 

....A. ..•[_, ;_ 


.'m^. I... 


: C 




i^ 1 


- w^ 

Q 

1 i [ ] ' 1 ! ' 



2.05 

2.00 

1.95 

JO 1-90 

1 1■85^ 

dT 1.80 
1.75 
1.70 



1.65 



D 



-^■* -H 

....■•I •..; 



,65 1.70 1.75 1.80 1.85 1.90 1.95 2.00 2.05 
P3 length 



1.10 1.15 1.20 1.25 1.30 1.35 1.40 
Ml length 



ffl = System A • = System B D = System C 



FIG.2. Bivariate plots for some lower tooth measurements of Riversleighfiwrramj^ Specimens. All measurements 
inmm. 



TABLE 2. Column 1, Kolmogorov-Smimov Lilliefors tests for normality. Probability (P) values below 0.05 
indicate a difference from normality significant at the 95% level. Columns 2 and 3, mean values for Systems 
B and C respectively. Column 4, Students t-tests; P indicates a significant difference between Systems B and 
C. Abbreviations as for Table 1 . 





Lilliefors Test (P) 


Mean - System B 


Mean - System C 


T-test (P) 


P3L 


0.088 


1.82 


1.79 


0.266 


P3AW 


0.175 


1.03 


1.05 


0.474 


P3PW 


0.455 


1.22 


1.22 


0.928 


P3MW 


0.678 


1.26 


1.29 


0.211 


P3LH 


0.215 


1.43 


1.44 


0.923 


P3BH 


0.904 


1.72 


1.76 


0.254 


Ml L 


0.002 


1.25 


1.21 


0.071 


Ml AW 


0.193 


0.79 


0.76 


0.268 


MiPW 


0.072 


0.96 


0.95 


0.625 


M2L 


0.009 


1.09 


1.09 


0.903 


M2 AW 


0.009 


0.87 


0.91 


0.041 


M2PW 


0.585 


0.95 


0.98 


0.111 


M3L 


0.593 


0.93 


0.98 


0.458 


M3 AW 


0.177 


0.85 


0.81 


0.394 


M3PW 


0.038 


0.85 


0.81 


0.458 


Ml-2 


0.054 


2.32 


2.27 


0.226 


Ml-3 


0.001 


3.25 


3.16 


0.529 



NEW OLIGOCENE-MIOCENE BURRAMYSFROM RIVERSLEIGH 



51 




• = System B □ = System C A = C+ (Encore Site) 



FIG. 3. Specimens of Burramys from various sites ai 
Riversleigh plotted on principal component axes ob- 
tained using 11 measurements from P3 and M].:. 
Eigenvectors recorded in Table 3. X = first principal 
axis. Y = second principal axis, Z (perpendicular lo 
page) = third principal axis. Solid line encloses spec- 
imens from System B sites. Dashed line encloses 
specimens from System C sites, including Encore 
Site. Dotted line excludes from System C 'aberrant' 
specimen QMF30104. indicated by arrow. 

but appears not to be the case for P3 buccal height 
(Figs 2C). M2 posterior width shows the opposite 
trend (Fig. 2B), whereby System C specimens arc 
on average larger than System B specimens. 
Student's t-tests show these differences to be 
non-significant at the 95% level (Table 2), but a 
principal components analysis employing dimen- 
sions of P3 and M|.2 (Fig. 3, Table 3) confirms 



that total variation is explained partly by System 
C specimens being smaller than system B speci- 
mens. Eigenvectors for component 1 are all pos- 
itive (Table 3). indicating that this is a general 
'size component'; specimens scoring high on the 
first component (i.e. falling further towards the 
positive, or right-hand side of the X-axis in Fig, 
3) are larger than those to the left. Although there 
is considerable overlap between Systems B and 
C, the centre of mass of the System B distribution 
is further to the right than that for System C. 

Specimens from Encore site (younger than Sys- 
tem C, ?early late Miocene) cluster at one ex- 
treme of the System C distribution, with the 
exception of a single large aberrant specimen 
QMF30104 from Gag Site (Fig.3). In the System 
B-Sysiem C continuum (Fig. 3) the cluster of 
Encore Site specimens falls on the 'older' (Sys- 
tem B) end of the System C spectrum. 

Despite the apparent trend of mean difference 
between specimens from Systems B and C, spec- 
imens from both Systems are present in each of 
the apparent peaks of the univariate distributions 
(Fig. lA-F). This suggests that the underlying 
structure of the sample is not simply anagenetic 
change tracked from the older System B sites to 
younger System C sites, though such may have 
occurred. The bimodality of several of the fre- 
quency histograms may rellect sexual dimor- 
phism and/or 2 roughly contemporaneous taxa. 
This suggestion is also supported by data plotted 
against sites arranged in estimated siraiigraphic 
order (Fig. 4A-F.) Although samples from indi- 
vidual sites are inadequate to compare wilhin- 
and belween-site variation statistically, variation 
between sites is only a little greater than that 
within Upper site, provenance of the largest sam- 
ple. Caution is therefore necessary when interpre- 
ting apparent between- or across-silc trends (such 



TABLE 3. Results of principal components analysis using 1 
specimens from Riversleigh. Abbreviations as for Table 1. 



measurements of P3 and M1-2 of Burramys 



Component; 


1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


11 


Eigen Value: 


4.971 


2.177 


1.179 


0.803 


0.638 


0.497 


0.336 


0.230 


0.097 


0.049 


0.022 


Percent 


45.194 


19.790 


10.716 


7.299 


5.801 


4.519 


3.056 


2.092 


0.883 


0.447 


0.203 


Cumulative 
























percent 


45.194 


64.984 


75.699 


82.998 


88.799 


93.318 


96.374 


98.466 


99.350 


99.797 


100.000 


Eigenvectors: 
MiL 


0.304 


-0.297 


0.076 


0.510 


-0.328 


0.126 


0.172 


-0.398 


0.088 


0.304 


0.376 


MjAW 


0.323 


-0.178 


-0.285 


0.040 


0.397 


-0.500 


0.507 


-0.020 


-0.033 


0.188 


-0.282 


MiPW 


0.387 


0.005 


-0.265 


-0.199 


0.131 


-0.356 


-0.414 


0.033 


0.006 


-0.188 


0.623 


M2L 


0.209 


-0.489 


0.152 


-0.278 


0.293 


0.353 


-0.240 


0.294 


0.277 


0.426 


-0.060 


M2AW 


0.203 


0.368 


-0.472 


-0.204 


-0.053 


0.535 


0.381 


0.193 


-0.138 


0.152 


0.213 


M2PW 


0.362 


0.150 


-0.390 


0.127 


-0.107 


0.168 


-0.380 


-0.299 


0.338 


-0.150 


-0.518 


M1.2 


0.324 


-0.403 


0.142 


-0.106 


0.051 


0.290 


0.099 


-0.117 


-0.507 


-0.567 


-0.111 


P3L 


0.363 


0.015 


0.179 


0.117 


-0.512 


-0.184 


0.118 


0.666 


0.169 


-0.164 


-0.117 


P3PW 


0.188 


0.344 


0.215 


0.617 


0.510 


0.146 


-0.184 


0.248 


-0.204 


0.018 


0.016 


P3LH 


0.322 


0.303 


0.313 


-0.303 


-0.219 


-0.159 


-0.198 


-0.191 


-0.468 


0.461 


-0.187 


P3BH 


0.250 


0.332 


0.498 


-0.258 


0.211 


0.056 


0.313 


-0.267 


0.486 


-0.218 


0.112 



232 



MEMOIRS OF THE QUEENSLAND MUSEUM 



1.40 




J 1.10 

CM 

^ 1.06 

1.02 
2.55 

5 2.45 

^ 2.35 

X 2.25 

2-15 
2.10 

2-00 

I) 1.90 

c 

-«1.80 
1.70 



-B 

- ffl 

' 1 


HI 


4i^' 


• • 

• 

m" 


a 


1 
i * 

™r- 






i— 

• r 

-T — r 


• f • 








1 

a 
— 1 — I 


• 

I 1 


— r 



c_ 

- ■•I— 


ITT- 

Tzrfrz 


.lA.„i.„....J_..L i - 


t 
1 

......g. 

1 r 


1 — 1 — ! — r i i i 



D 



D i 







System A 



System B D - System C 



FIG 4. Rivcrslcigh Bumoms: size measures againsi 
siles in slraligrnphic sequence (Archer el ul., 1989). 
Disumce-sonhorizoniahiXLs arbitrary. Sites: l=Whiie 
Hunter: 2=Crcaijcf s Rampans: 3=0utasilc: 4=RS0: 



as size decline over time) as being significant. 

Although site S (Fig.4) includes Ten Bags Site, 
Mike's Potato Patch and Upper Site, most speci- 
mens are from Upper Site and these span ihe 
range of variation at site 8. 

If two tnorphotypes are present they are both 
ivpresenled in Upper Site {Fig.4). Muirhcad 
( 1 994 ) demonstraled size-guilding comparable to 
ihal in Recent mammal communities among 7 
Upper Site bandicoot species separated on si/e; 
this is possibly due to competitive displacement 
of laxa that eat size-variable foods such as seeds 
or insects wiihin a single community. Thus, 
Upper Site probably represents a single diverse 
community and the 2 morphotypes o\' Hurra my s 
could be sexual dimorphs or sympalric laxa. Wc 
reject sympalry because of morphological consis- 
tency of specimcfis tailing near the peaks in the 
Ml-: length distribution; the si/e difference be- 
tween those peaks is too small (Roth, 1981) lo 
represent 2 species in different niches at the same 
level of a food wch. The ratio of the second peak 
lothe first (Fig. 1A-C)is 1.06, short of the often- 
cited cutoff value o^ 1.3. (Roth (1981) showed 
that the 'constant ratio rule' is empirically unsub- 
stantiated, bul suggested thai character displace- 
ment is unlikely lo be indicated by ratio values 
lower than 1.3.) 

Challenging the likelihood of either sexual di- 
morphism or sympatry is the fad that some spec- 
imens are m the higher peak u\' apparently 
bimodal distributions for some measurements, 
bul the lower peak for others; whereas other 
specimens remain in one peak or the other for all 
or most measurements. There appears to be no 
combination o[ features that can be used lo sub- 
divide the sample; this is supported by the multi- 
variate analysis (Fig. 3) which fails to divide the 
sample. Ageneral trend to declining si/.e through 
Systems B and C (Figs 2.3.4A,C) is evident, bul 
some Fncorc Site specimens suggest reversal of 
the trend. 

SUMMARY 

Riversleigh liurramys .specimens may repre- 
sent two populations. Patterns of variation also 
suggest a dine of dereasing si/e through lime: 
however, small sample si/cs and uncertainly of 
relative ages limit the reliability o\ ihis observa- 

5=W;iync'TWor^>=?anic^puiuin. Neville's Garden 
and Dirk's Towers; 7=lnabeyancc; S=Ten Bags, 
Mike's Potato Patch and UpperSitc; 9=KangarCKi Jaw; 
10=Gag; ll=Lasi Minute; 12=Main Site; 13= Jim's 
Jaw; 14-Wang; l5=Encorc. 



NEW OLIGOCENE-MIOCENH BURRAXfYSVROKA RtVERSLElGH 



2Si 



lion. Eltwo pnpulaiicMVs have heen sampled, mag- 
nitude ami disiribulion of viiriiiliun MiggtrsI thai 
ihcsc arc males a.nd Icmalcs of I species. Exiani 
populations of B. parvus are nut dcnially difMi»r- 
phic (BrammalK unpuhl.), hut rhcir iilpine hal>iial 
is far removed froiii ihe Miocene niinforesi envi- 
ronmenialRiversleigh(ArchercIal., \9M, \9^\) 
m it is not possihlc lo infei thai Rece»i and 
Miocene Hurraniys share pupoluiion siruciureb. 
Wc recognise a single new species. 

SYSTLMATIC PALAEONTOLOGY 

Class MammaJia Linnaeus. I75S 
Supercfthorl Marsupialia Ilh^er. IH|f 

Oaler Dipa>lodonna Owen. I H66 

Sufviffdmilv Buiraniyoidea Bmoiri IHyS 

Kimily BurramyidacUnL)om. 189H 

Burramys Broom. 1896 

Burramys brulvi sp. nov. 
Oigs 5-9: Tables L4) 

KTYMOIOGY For the laic Aninir Brutv who. to- 
gether with his duughlci tJaiiic Clurkc. hcli»cLlcollc(.-i 
m;iT!y specimens and discovered Braty <& the BcasiSiie 
ail ihcGag Ftaa*uu. 

MAIHKIAL. lioloiypc QMK30Hn (Pig. Sk a \c\\ 
denary <DEN| with I). P1-3, M| 2 and alveoli for l> 
and M:^-4. The tip ol li is missing, us aic \Uc condyhir. 
nn^ii liir n?ici coronoid processes. Paraiy pes QMF30 1 7(> 
(Fig. 6). R DHN wjlh F: ,v.M 14. Hrokc-n ii!ilLTit)rUt 1*2 
aiut missing (he ascentling ivimns *mij condylar, anguhir 
;mdconnioi(Jpnvesscs.QMF30riOl (pig. 7k L miiAilla 
with P"' ^. M ' ^ uihI palalc medial lochfckiet'lh. Types 
from early Iti mid MIoccpl' Ijppv^r Sue on GtHllhL'lp 
Hill. DSiic Plateau. 

Other material: SYSTEM A - White Hunter Siic. 
QMI-21144 RM-.; QMF:3500. DEN ui(h RP^ SY.S- 
TEM B - Camel Sputum Sue. QMF2i)732. DEN wiih 
RVJi, Pv. QMr2()735. R DFN. QMr20736. \M-: 
QMF3060(3. maxilla >Mth LP* ■^. M*; OMF3OI07. 
m-:NwuhLlt,P2-^,M|.r.QMF3(}ll(),DFN\vilhRli. 
P?.j, Ml-:. Injbcyancc Siic: QMF3007i). pjf^fv ^iih 
U»1. M\A. Mike's Pol;iU) Palcli Siic; yMi-2()759. 
DfN vvtih LM:: QMr2(nhi). I M '; QMF2()7o I , Pi or 
P\ Neville sGardcn Sue; OMI-2()7 1 S. DliN with KP^. 
M I . (,>Mr2074N. I .M2; QMF2(W()2. DliN with Rli . Pi. 
Mi; QMF2:^34w, DEN w.ih I P3. Ml---; QMF2337fi. 
DEN with RPi. Ml. QMF235tl. DEN wiih RPr 
QMI 24261. maxiil;. wiih RP-^ QMF300Ht), maxitkt 
wiih KP--\ M'-, QMr30092. maxilla wuh RP- \ 
M'--*; gMF3()l 13. DEN wiih RPv M|; QV1F31)1 14. 
( f^^: OMF3iU32. DEN wiih LP\, Mi v, OMF3U27I. 
KNr. OulasUe; QWrZUim, I DI:N; yMF30(KS(». 
DEN wiihl.li. Pr Ml 2. RSOSilc: OMF3nOK|, DEN 



wiih I.P^, Mi-v. OMF3O084. DEN wiih Rlj. Pi. M|. 
QMF3(H)44, maMJla wiih RP-: nMF30l4<K I.Pv 
OMF30(4l. LP"*; OMF30I41 RP\ Ten Bags Site: 
OMF23502. DFN with LPv Mi l!pper SiiC 
QMF20774, DFN wiih Rli: QMF20775, DEN with 
LI I, P3; OMF20776. DHN willi RP^. yMF20777, 
DEN with RMv QMF2l)7SS, maxilla wiih RM*'^. 
OMF207S(i, DliN wiih tA1|-^, Vy. QMF207S7. max- 
illa wiih LM'. P- \ QMf"207HS\ maxilhi wiih I.M'; 
OMF3tK)K2, DEN with l.P;. Mi-^: L>MF3(){)S3. DEN 
wiih RP-'.^,M)--':OMI-^()()S^, DEN with IJi.P^,M| 
QMF3()aS(:. DEN wuh Rli. P"- x Mi-^ 0MF3tM:iSL 
nuiMtla wilh LP-'^ QMF300SS, masilla wiih EP^'-^; 
OMF30C^M. maxilla wuh LP- ^ M*-*: QMF30095. 
niuxilla wiUf RP^ M'""^; QMF3(KW6. maxilla wilU 
LP-"^; QMF3()007. maxilla v^ith RP-"^: OMF30(m. max- 
illa with KP*. M'. C)MF30(W. maxilla with RP\ 
OMF30I0I. maxilla with LP- ^ M' -: QMF301i>2. 
DEN wiUi Lli. P1.3. Mu2: QMF30)O3, maxilla with 
LP-\ M': QMF30I06. DEN wiih Rli. P^-v. 
QMF30ill, DEN wim Ll|. P: %. Mi: OMF301I2. 
DHN wiih RIk P^>; QMF3()1 17, DI-.N w.ih RP^, M|-r. 
OMF30ltK,DENwuhRli.P^.M|:OMF30tP^,DEN 
with RM-* \: gMr30I20. DEN with LIM. M| 2- 
QMF30I2l,i:»ENwilhLI|,P^;QMI'3()l22J)r'.NNk'ith 
Lli. P.v. OMF30I23. DEN woh I Pi. Mi-j: 
OMF30124. DEN wuh KP^. M | v. QMF30I 25. DEN 
with RP<, M|; OMF3(M27, DEN wiih I.li. P?. 
OMF3012S. DEN with RP3: QMF3(1I2»). R DEN: 
gMF'^OHO. 30131. I. DEN; QMF30I33. DEN with 
RPi Mi-r QMF30I38, maxilla wi(h LP-: 
QMF31)I39, RP-; QMF30!46, 3UI4S, 31)149. 30152 
LP^: QMF30I47. 30150. 30154. 301 55. 301 79, 30182 
LP-^; QMF3015I, 30153, 30174. 30IS0. 301X4 RP-^ 
OMF301f)0. LM2: QMF30I 64-301 07. LM'; 
QMF3()lhS, 30173. 30177 RM'; yMF3()f76, DI:N 
wiih RP:.5. M1.4: OMF30[SI. 301X3 RPi; 
OMF30IK5, 30iyO RM't, QMF301S6. KM-. 
QMF30IK7.l.l|;QMF10US.RIl.Q.MF3()l«y,RM> 
Waync^s Wok Sue: OMF20725. maxilla with RP': 
OMF2072(.. maxilla wuli KM'*-: OMF20737. imXtl- 
larv (ragmcni with RP\ OMF207"^S, Dt'.N with RM|; 
OMF20744. DEN wuh KMj. P^. OMF20745. DtX 
with LM2 V. QMF20746. DEN with RM1-2: QME 
22KI6. maxiihi wuh RP--\ M' •^. QMF:H)108. DEN 
with RP2-V Mi-r. OMF3nn6, DFN wilh LP-^. Mi 
Wavuc's Wok 2 Sue: QMF30I00. DEN wuh Rl 1. P^. 
Mi-v OMF30175, LP-^ SYSTEM n OR C - (Men ol 
-Xgcs 1 Sue: QMF20'H)5. R DEN. Clcli of Ages 2A 
Siie- QMr-22772. maxilla wilh KPl M'. Ciell ot A^l-s 
4 Site. QMF20767_ RP'' QMF20S33. RP^: 
QMF20S36. KP;: QMF23200. RPV SYSTEM C - 
Encore Site: OMF20752. 1 Mv. QMI'2()7S^ ].l»^; 
QMF20754. LM| , yMF209O4, DEN wuh RM | -». P^ 
y, OMI*23462. DEN wilh KMu2. Pt. OMF2a334. 
DFN with I.Mj.v. QMF24424. DFN wiih LMi; 
OMF24420. DEN wuh LI 1 . P^. M:: QMF24552. RP\ 
QMF24727. DEN with Lli. P^, M1.2 Clag Silt- 
OMF30t)7K. DEN Willi Kin. QMF300')?, maxilla >\ilh 
IP^- QMr^O|04 DFN wilh LIk M|-2, V\; 
QMF30I34.LDEN' OMF30t3?. DFN wilh I .P^, Mr 



234 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 5. A-C. Biirramys brutyi n. sp. holoiype QMF30102. Lel'i deniary with li P2-3 Mio in (A) buccal, (B-B') 
occlusal siercopair and (CJ lingual views. D-F, Burramys bruryi paralype QMF3009I. Left maxilla with P*^*^ 
M '"* in (D) buccal, (E-E') occlusal siercopair and (F) lingual views. Scale = 2mm. 



OMF30137, LP^ QMF30156, LMr QMF30I57. 
RM'; QMF3()I58, RM|; QMF3016I. LM^ 
QMF30170, RP\ QMF30171. RP^ Henk's Hollow 
Site: QMF30172, LP\ Jim's Jaw Site: QMF30178, 
DEN with RPv Kangaroo Jaw Site: QMF301 15. DEN 
with RP^,, M1.2. Last Minute Site: QMF30I05. DEN 
with R!|, Mi-^ Pxi; QMF301I6. DEN with RP^, 
M1-2: QMF30I43. LP^ QMF30144. RP^; 



OMF30145, LP-^ apical fragment; QMF30I62. RM^ 
QMF30163. RM^ QMF30i69, DEN with RP^, Mi 
Main Site; QMF30109. DEN with RPv Ringtail Site 
OMF20756. RP^ QMF20757. maxilla with RM^- 
P^. Wang Site: QMF20763, maxilla with LP-"^ 
QMF20766. DEN with RMi, ?\, QMF30272. RP^ 
AGE UNCERTAIN - Greaser's Ramparts Site 
QMF2077I.LP^ 



NEW OLlGOCENfe-MIOCENfc BURRAMYSYROM RIVERSLRIGH 



155 




FIG. b Butramyshrutyt paraiype QMF30 1 76: occlusal 
sxcrcopaif of right demary fragmeai with P:-3 ami 
Mi.4. Scale = 2nim - 

DIAGNOSIS. Differs from B. thradiatus and B. 
parvus in being smaller, in having upper and 
lower pltigjaulactiid P3 .smaller and with fewer 
(5-6) cuspules and associated ridges and in hav- 
ing 2-nKUcd upper and lower M4. Dcnlary and 
maxilla more robusi dian m B. par\-us, wiih 
smaller palatal vacuities, shorter I2-P: interval 
and less reduced posterior molars. P.^ with larger 
crown and larger posterior root than that of B. 
wakefielili and diverging less from anteroposter- 
ior molar row axis. Pi-2 double-rooted: single- 
rooted in B, wakejleldi. Distinguishable from B. 
wcikefieldi and B. parvus by Mi cusp morphol- 
ogy: proioconid more lingual in B. Mtikefiehli 
than h. parvus or B. hntni; meiaconid more an- 
terior in B. parvus than B. brntyi or 6. wukefividi, 

COMPARATIVE DESCRIPTION. Thedenlary 
oiB. bruryi is subequal to that of B. wakefieldi in 
si/.e and shape. Both are more robust ihan thai of 
B. parvus but slightly less so than ihui of B. 
iriradiuius. The leading edge of the ascending 
ramus of H. bratyi is considerably more robust 
and rises at a sleeper angle from the horizontal 
axis of the denlary than does ihatof 5./?^/n'(/^. but 
not quite as steeply as that of B. triraduituK. The 
1:-P: interval is shorter in B. bruni than in B. 
paryus but is not as short, relative to the length o\' 
the ramus, as that ofB, frfradiatus, 

Lowerdeniiiion. Ii is long, slender and procum- 
bent, with the tip curved upwards and slightly 
Iwisted. It is slightly less procumbent in B. brut^n 
ihanini?.pt/n'«.y. The crown of li isbusally about 
the same dorsoveniral thickness in H. bntrvi and 



B. par\nis hut a little thicker in B. tnradUUus. l\ 
of B. brun'i ihins abruptly about half way along 
lis exposed length, with the anterior half of the 
tooth being narrower than the posterior half. In 
lateral view Ii of iJ. hrurs'i is more curved than in 
the other species. 

h has not been idenlitlcd in B- hrutyu B- 
wakefieldi or B. niraduvits. In B. pantis Ij is 
small and stngle-rooied. insertmg into a shallow 
alveolus directly behind the posterior alveolar 
margin of li Its crown inclines forwartJ looverlie. 
Ii posierobasally. In some specimens o\ B. bruryi 
there appears to be the remnant of a small alveo- 
lus in the fragile region between li and Pi. sug- 
gesting a small, single-rooted h. 

Pi is sinall, 2-rooied and cap-like, the crown 
swelling beyond the roots in all directions. There 
is a minor ridge along the iintcroposierioraxis of 
the tooth, with the crown sloping away from tlie 
crest on each side towards the lingual and buccal 
margins respectively. In dorsal view it is almost 
circularinoutline, being slightly wider than long. 
The crt)w n does not extend as far beyond the roots 
posteriw'iy as it does in other directions. In B. 
paribus the crown is shorter and flatter than in B. 
bruryi and is also procumbent, rising slightly at 
its anterior end to overlie the posterior end of h; 
it is ovoid in dorsal view (slightly longer an- 
teroposleriorly ) and its posterior end is reduced. 
The anlenot root of P| mserts anterobuccal lo 
the posterior root- The posterior alveolus is closer 
to the anterior alveolus of P: than it is to the 
antcrioralveolusofPi.insertingslighllyiingually 
and anterior lo the anterior alveolus of P:. The 
septum separating the posterior alveolusofPi atid 
the anterior alveolus of P: frequently breaks 
down so that they form a single cavity. In some 
Specimens, theret'orc, there may appear to be only 
three alveoli in the region which had been occu- 
pied by the 4 roots of Pi and P^. Even with the 
septum intact, the arrangement of alveoli tnight 
suggest that the posterior alveolus of Pi and the 
anteritw alveolus of P2 belonged to the same 
tooth. Whereas in B. bruryi the alveoli of Pi and 
P: are closely but unevenly spaced, in B. parvus 
the 5 alveoli uf 1:, Pi and P:- are evenly spaced 
and in the adult animal there is a small gap 
between Pi and Pz (in subaduU or young^jr ani- 
mals Ihe teeth are closer together). 

P2 is similar in shape but a little larger than P|. 
The slight anieropo.sterior crest lies at an angle 
(lingual posteriorly) across the alveolar margins. 
directly above an imaginary Ime joining the cen- 
tres of the P: alveoli Posteriorly the crown ex- 
tends beyond and Tises above the rooi.. 



256 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 7. Lower cheekteelh 1)17?. /jn/n*/ in occlusal view. 
A-C. QMF 30102. A, LPv B-B\ LMi. CC\ LM\ 
D-D'RM3ot"QMF30100.E-E\RM4ofQMF30176 
B-E stereopairs. Scale = 1 mm. 

Icrminaling in a small cuspulc and abulling P-^. 
Anieriorly, the crown extends slightly beyond the 
root. Lingually and buccally the crown swells out 
and falls away to a rounded point on each side. 
The buccal, ventral apex is slightly higher and 
more anteriorly located than the lingual apex, so 



FIG. 8. Left upper cheekteeth of B. hrtiivi paraiype 
QMF3009 1 in occlusal view. A. P-\ B-B\ M ' . C-C\ 
M-. D- D', M^"^, B-D stereopairs. Scale = 1 mm. 

thai the crown is somewhat twisted. In B. parvus 
Pz is larger and relatively longer, with a crown 
that extends further beyond the roots, particularly 
anteriorly, giving the anterior end oi the tooth a 
shelf-like appearance in lateral view. The crest is 
less clearly defined than in B. hnityi and approx- 
imately parallel to the axis of the h-P: interval. 



NEW OLIGOCENE-MIOCENE BURRAMYSFROM RIVERSLEIGH 



257 



The crown oCPt shows less lingual-buccal asym- 
metry than in B. brutyi. The posterior end of the 
crown rises higher and more steeply than in B. 
hnityi with a distinct hump above the posterior 
root of the tooth, posterior to which the crown 
increases only shghtly in height. The Pz of B. 
triradiattis is similar to, but larger than, that of fi. 
bruryi. It is wider but shorter than ?: in B. parvus 
and almost circular in dorsal view. Although it 
protrudes beyond the roots in all directions, it is 
flatter than in B. brutyi and B. parvus. As wilh B. 
brutyi, the buccal side is displaced ahead of the 
lingual side and as with B. parvus, in lateral view 
the crown has an anterior 'lip'. The anteroposter- 
ior crest is poorly developed, A P2 (NMV 
PI 800 16) assigned to B. triradiatus by Turnbull 
et al. (1987) is considerably larger than and dif- 
ferent to P2 in the Holotype. It is 1 -rooted, in 
contrast to P2 in the Holotype, which has 2 or 3 
roots. NMV PI 800 1 6 could possibly be a B. 
triradiatus ?~. P2 is not known from B. wakefieldi 
but appears to have been I -rooted. 

The plagiaulacoid crown of P^ is longer and 
taller in B. brutyi than B. wakefieldi, larger in B, 
parvus and larger again in B. triradiatus. P3 o'i B. 
brutyi has 5 or 6 dorsal cuspules and associated 
ridges. The anterior edge of P3 rises vertically in 
B. brutyi, curving back dorsally to an almost 
horizontal serrated crest. The anterior profile is 
straight in B. wakefieldi, but leans backwards 
slightly as it rises to an also horizontal crest. The 
anterior root descends from the crown more an- 
teriorly and buccally in B. wakefieldi than in B. 
brutyi. In B. triradiatus and B. parvus, the ante- 
rior profile of P3 curves forward then backward 
as it rises, giving the corrugated tooth a 'fanned' 
appearance and increasing the length of the dorsal 
edge. In B. triradiatus the anterior root curves 
forward slightly as it rises, with its convex profile 
continued by the crown. In B. parvus the root rises 
vertically to the base of the crown, then the crown 
expands gently forward. The P3 blade is slightly 
concave lingually and convex buccally. The ex- 
posed portion of the anterior root of P3 protrudes 
further beyond the jaw^ margin buccally in B. 
brutyi than in B. parvus. It is also in high relief in 
B. wakefieldi and B. triradiatus. In B. parxnis, ihe 
posterior end of the crest has shifted lingually and 
backwards (relative to its position in B. brutyi). 
Thus the anterior angle between the long axis of 
the P3 crest and the molar row is greater in B. 
parvus than B. brutyi, as is the angle between this 
crest and its underlying roots. The posterior root 
of P3 is also smaller buccally in B. parvus than in 
B, brutyi and is smaller again in B, wakefieldi 



because the posterior end of the crest and hence 
the direction of the bite force in that region has 
shifted lingually. The anterior end of P3 is more 
attenuated in B. parvus than in the other species. 
Some specimens of fi. brutyi have cracks running 
from the dorsal cuuing edge basally and hack- 
wards, stopping near the base of the crown. P3S 
of each of the other species have similar cracks. 
They are particularly frequent and extensive in B. 
triradiatus. The P3S of/?, triradiatus also gener- 
ally show more wear on the anterior end of the 
dorsal cutting edge than is evident in the other 
species. 

Lower molars are bunodont in Burraniys. They 
differ mainly in size. Mi cusp morphology and 
degree of reduction of M4. Some unworn molars 
of B. brutyi are slightly crenulate, unlike other 
species of Burramys, but since crenulation is rare 
in B. brutyi and since molars of the other fossil 
species are poorly known, this feature is noi re- 
garded as diagnostic. The molar gradient is 
greater in B. par\us than in other species. 

Ml is approximately the same size in B. brutyi 
and B. wakefieldi and is larger in B. parvus. It has 
two roots in each oi' these species. M| is not 
known from B. triradiatus but judging from its 
alveoli was 3-rooled and relatively small, with 
M4< Mi< M3< M2. The irigonid rises more stee- 
ply against P3 in B. brutyi and B. wakefieldi than 
in B. parvus, with Ihe proloconid taller in com- 
parison to the metaconid, P3 and Mi are therefore 
more disparate in height in B. patrus than in B. 
brutyi or B. wakefieldi. Posteriorly, the crown 
extends further beyond the roots in B. parvus Ihan 
in the other species. h\B. wakefieldi ihe Qwiocon'id 
is particularly tall. In all species, the Mt 
postmetacristid is continuous with the longitudi- 
nal axis of the dorsal crest of P3. In B. brutyi and 
B. pan'us the premctacristid swings buccally to 
meet Ihe postmetacristid, creating a disjunction 
between the P3 crest and the lingual crests of M 1. 
The poslprolocristid/premetacristid angle is more 
obtuse at the metaconid in B. brutyi than B. 
parvus because the metaconid is more posteriorly 
positioned in the former than the latter. The break 
in the P3-M1 blade system is therefore, longer in 
B. brutyi than in B. pannis. In B. wakefieldi the 
protoconid is more lingually positioned so thai 
the crests associated with the P3 and Mi pro- 
toconid, metaconid and cntoconid form an almost 
straight line. 

M2 is smaller in B. brutyi than B. triradiatus or 
B. parvus. M2 of the latter is slightly longer and 
narrower than that of B. triradiatus. It is propor- 
tionately shorter in B. brutyi than B. parvus and 



25S 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 9. B. bntni sp. nuv. A-C. Icfl Pi-: with 1: alveolus and anlerobasaj porlion of P3. holotype QMF30102 in 
(A-A') lingual. (B-B') occlusal and iC-C) buccal views. D-D\ left P- -^ and anierior portion of M', paratype 
QMF30091 in lingual view. A-D stcreopairs. Scale = I mm. 



NEW OLIGOCENE-MIOCENB BURRAMYSFROM RIVERSLEIGH 



259 



TABLE 4. Measuremcnls of B. brutyi types. From 
holotype where possible. M3-4 lengths and widths, 
M1.3 and Mi-4 from paralypc QMF30176. All upper 
tooth measurements from paralype QMF30176. Ab- 
breviations as for Table 1. 



Lower teeth \ 


P, L 


0.48 


Pi MW 


0.55 


P2L 


0.62 


P2MW 


0.64 


P3L 


1.76 


P3AW 


1.07 


P3PW 


1.23 


P3MW 


1.31 


P3LH 


1.45 


P3BH 


1.76 


M, L 


1.21 


Ml AW 


0.83 


MiPW 


0.97 


M2L 


1.10 


M2AW 


0.88 


M2PW 


0.94 


M3L 


0.98 


M3AW 


0.85 


M3PW 


0.80 


M4L 


0.66 


M4AW 


0.64 


M4PW 


0.50 


M1.2 


2.30 


Mi_3 


3.18 


Mi_4 


3.83 



Upper teeth | 


P^L 


1.01 


P^MW 


0.60 


P3L 


1.91 


P^AW 


0.92 


p3pw 


1.21 


P^MW 


1.21 


P^LH 


1.56 


P^BH 


1.64 


M^L 


1.11 


M^ AW 


1.17 


M^PW 


1.21 


M^ MW 


1.39 


M2l 


0.96 


M^AW 


1.13 


M^PW 


0.89 


M^L 


0.82 


M^AW 


0.90 


M^PW 


0.66 


M^L 


0.64 


M^AW 


0.67 


M^PW 


0.40 


M^-2 


2.07 


Mi-3 


2.90 


M^-'* 


3.51 



very slightly shorter than B. triradiatus. M2 is not 
known for B. wakefieldi. It has two roots in each 
species except forfi. triradiatus, in which it has 
three. In a few {<5%) oi' B. hrntyi specimens the 
anterior alveolus has, ventrally, a septum (or re- 
mainder thereoO subdividing it basally into 2 
compartments, suggesting a root bifurcated al its 
lip. This condition may be intermediate between 
the 2- and 3-rooted conditions. In B. brutyi and B. 
/?«n'M.s there is frequently a small cuspid halfway 
along the lingual margin of the crown, at the 
junction of the poslmetacristid and the prc- 
entocristid. Sometimes the cuspid is not clearly 
differentiated from the postprotocristid. It is the 
same size in both species even though the tooth 
is larger in B. parvus. The cuspid is not evident in 
B. triradiatus, although there is a small dorsal 
protuberance on the anterior end of the pre- 
entocristid of NMV P 1 58628. In each species the 
postprotocristid curves lingually from the pro- 
toconid before straightening and running approx- 
imately parallel to the tooth axis until interrupted 
by the transverse hypoconid-entoconid lophid. 
Postprotocristid curvature is less extreme in B. 



brutyi than the other species. The cristid obliqua 
lies parallel to the tooth axis, forming a 
posterobuccal cingukir pocket between itself and 
the postprotocristid. In all species the buccal 
cusps are bulbous. The hypoconid causes the 
posterobuccal comer of the tooth to extend be- 
yond its basically rectangular outline. The lingual 
cusps are slightly ahead of the buccal cusps, 
skewing the sides of the tooth slightly. They are 
more crescentic than the buccal cusps and, to- 
gether with their associated crests, form a blade- 
like structure. 

M3 is similar to, but smaller than, M:. Cusps are 
lower and basins shallower, with the crown sur- 
face showing more wear than M2. M3 is smaller 
in B. brutyi than the other species, is slightly 
larger in B. triradiatus than B. pafrus and is not 
known from B. wakefieldi. Interspecific compar- 
isons of M3 are as for M2 except that in B. brutyi 
and B. parvus, but not B. triradiatus, the Ungual- 
buccal skew is slightly more pronounced than in 
M2. In all species M3 is slightly shorter an- 
teroposteriorly than M2. In B, triradiatus the pro- 
toconid and hypoconid of M3 are subcqual 
whereas in M2 the hypoconid is larger. In M3 
there is a distinct cleft dividing the rounded pro- 
loconid and hypoconid. 

M4 in B. brutyi and B. wakefieldi has 2 roots, 
whereas in B. triradiatus it has 3 and in B. parvus 
1 . While most specimens of fi. brutyi have 2 roots 
or alveoli for M4, some have 3 and a few had I 
root. Such variation is not evident in the B. 
triradiatus or B. panms, M4 is not known for B, 
wakefieldi or B. triradiatus hut the alveoli of ^. 
triradiatus suggest that it was far less reduced 
than in B. parvus and possibly less reduced than 
in B. brutyi. M4 is low-crowned, with low cusps 
which quickly wear down. It is smallest and most 
degenerate in B. parx'us. 

Upper teeth oiBurratnys anterior to P- have not 
been recognised from Riversleigh or Hamilton, 
so discussion o\' the upper dentition will be lim- 
ited to P-"'' and M' ■*. Skull fragments and upper 
teeth of B. wakefieldi are unknown. The upper 
dentition of B. triradiatus is known only from 
isolated teeth. 

Maxilla. Palatal vacuities are smaller in B. brutyi 
than B. parvus. The anteroventral opening of the 
infraorbital foramen is also smaller (and less 
round) in B. brutyi. as are foramina in the a!- 
isphcnoid and squamosal. Known bones of the 
skull are more robust in B. brutyi than in B. 
parvus. In both species the maxilla is swollen 
around the P-^ alveolus, between the lachrymal 



360 



MEMOIRS OF THE QUEENSLAND MUSEL M 



and the infraorbital Tor^men. This swelling Is 
mon' exlcnsive in B. parvtt'i than B. hntni. wi(h 
P-^ and the iUUcrior limit of the niolor row hcgin- 
ninc furihcr forwanl in the living species. In 
ventral view, ihcanlcromcdiai limit oi* the /ygo- 
inalic iireh in B. hrutyi is level with a point mid- 
way between the proiocone and prottvonule oi" 
M'- In H. parvus it is midway hctwccn ihe 
uieiiKonuk Jitid prouuone o!" M' llio upper 
mular gradient is steeper in B. panuts than in the 
otlici* species. In H. hrutyt Ihe molar row rotates 
buccally around the muMlIa Iruni Iront to back, 
to a greater degree than occurs in B. parvus. 

Upper (icntition. P^ of B. btntyi is 2'rooicd and 
sunilar lo, alllutugh slightly larger than. Vi. A 
waak cresi runs from a small cuspule iii the hi^ih- 
csi point on the crown, which is midway along 
the raised posterior edge, to Ihe anlenor base of 
the crown, l.ingually and huccally the crown 
slopes towards the roots. The base ol the crown 
expands linguallyovct ihcposietior root, extend- 
ing Ihc crown outline posrcrolingually. This 
swelling is less pronounced in B. parvus, in 
crown view llie tooth is tcardn>p-shapcd, hemg 
just wider than the transverse diarrieter of ihc 
anterior kkh. Iii B. parvus, by Ciintriisi, the 2- 
rooicd P"^ emwn expands hcyond the rotns for Us 
whole length (more so posteriorly than anteri- 
orly). In boih species the crown is parallel in the 
edge ol the medially inclined pulule. lormmg an 
angle with P-^ and ihc molar row, Ahhou^^.h P- for 
B. triradiauis has not been idcniilied. the small, 
.sjngle-rootedtooih(NMV PIK0I)I6) (Jeiermined 
bv Turnbuil el al. (1987) tc» be a P2, is similar to 
P^N of B. brutyi and B. parws and is inlerprcled 
here lo be P-. 

In B. hrutyn. as jn B. trinidiativi and B. parvus, 
P^ is similar tc» Pv Regarding P^ anieroposierinr 
length. B. bruryi <B. pairus <B. triradiaius. P^ is 
dorsovenlrally sJiortesl in B. brutyi and slightly 
lallct in B irtrfuIi(ifus\\YiiU B parvus. I1 is simi- 
larly shaped in all three species but in B. brutyi 
Ihc crown decreases in anleropc»sierior length 
from base tn occlusal edge, whereas in B. par\'us 
and B. triradiarus the ventral edge ol the blade is 
at least as long as base of the crown. In anterior 
view, P ' of B. ht uiyi is as wide as that n\'B. parvus 
at its base, but tapers more rapidly and is hence 
ihickcf at the <x:clusal edge and more robust in 
appearance. In B. triradiarus the tooth is thicker 
basally than m B, parvus because of a broatler 
cinp.ulum (see below). Ii is thicker for most of its 
height hul tapers lo almost as thin an edge as docs 
B, panm P\ Whereas the deniary turns medially 



immediately anterior lo Pj, the maxi]Ia of 
Burmmys turns medially only anterior lo P^ P^ 
Iherefore docs noi appear to turn out from Ihe 
molar ri)w as much as P3; its crest is approxi- 
mately parallel to the molar row. Consecjuendy U 
does not have 10 retract anierobasally (as with ^^) 
10 insert into the hone and unlike P? its anterior 
edge, seen in lateral view, may appear lo extend 
slighdy forwartl basally. Probably as a conse- 
quence ol this, the bucc«l-convt?xity/lingual-con- 
cavity is, in all species, less pronounced ilvin in 
P(. hi occlasal view, P^ of B. brutyi is basically 
rectangular, bul with the anlenor end curving lo 
a rounded puini ;ind the posterior corners 
rounded. In B. parvus and B. triradiatus it is more 
ovoid, the anterior end again being a little nar- 
rower than the posterior end. and pointed. There 
is a narrow cingulum, poorly developed at the 
anlenor end. along ihe lingual and buccal sides o\' 
the crown. The cingulum is very weak in B. 
brutyi, slightly belter developed in B. parvus aiul 
significantly better developed in /?. iriradiatus. In 
this Jipecics the cingulum is sometimes 
emphasised lingually by a vertical wear facet that 
lerniinales abruptly al the cingulum. In B 
tnradiatus and lo some e\tein in B. parvus The 
second and sometimes third lingual ridges merge 
into the first which forms a curb that arcs back 
low ard ihe cingulum. This curb is loss prominent 
in H. brutyi, in which ridges approach the cingu- 
lum wilhoul merging. 

In all species M' has 3 roots — a larger lingual 
and 1 smaller buccal roots. In B. brutyi M* is 
wider ilian it is long. In B. parvus and B, 
ttiradiatus il is about as wide as long, in all 3 
species there is a swelling anierobuccal to the 
paraconc such that the antcrobuccal corner of the 
toolh IS a litlle larger than the poslerobuccal cor- 
ner. In B. I)tuiyi\\\crc is a distinct buccal Lingular 
basin or shelf at the intersection of the posi- 
paracrista and the premeiacrisia extending back 
to the level of the metacone and forward nearly 
as far as the paracone. It is sometimes delimited 
anteriorly by a small crest running buccaliy from 
the paracone. In B. par\*us this pocket is tittle 
more than a sloping cingular shelf. In B. 
triradiutus il is a narrow cingulum following the 
rounded paracone and metacone buccaliy 
(Turnbuil cl al., ]VK7. fig, 5A), The cciolopli of 
B. brutyi is roughly parallel to the anteroposterior 
a\i.s of the itioth. As wilii /^ parx'us, ihe paracime 
issignificanily larger than the other lusps, retain- 
ing its height as the tooth wciU's. The pnoioconule 
and melaconule are less developed in B. brutyi 
and B, triradiatus than in B. pamn. Hence in 



NEW OLIGOCENE-MIOCENE BURRAMYS FROM RIVERSLEIGH 



261 



occlusal view, M' ofB. brutyi is basically rectan- 
gular, with the anterior and posterior ends of the 
tooth parallel. In B. pannis it is longer and more 
curved lingually than buccal ly because of the 
inllated protoconulc and melaconule. In occlusal 
view there is an indentation between the paracone 
and metaconc in B. brutyi and B. triradiatus, 
whereas in B. parvus the crown outline between 
those cusps is almost straight. 

M- is rectangular in fi. bnayi (shorter an- 
tcroposteriorly) and considerably smaller than in 
either fi. triracliatus or B. parvus, in both of which 
it is about as wide as it is long. In all species it has 
3 roots and a small cingular pocket anierobuccal 
to the paracone, bounded lingually by a short 
preparacrista that runs perpendicularly from the 
anterior edge of the tooth to the paracone. The 
buccal cusps and their associated crests are blade- 
like in comparison to the more rounded lingual 
cusps. In B. triradiatus the buccal cusps are more 
pointed and the lingual cusps more rounded than 
in B. brutyi or B. parvus. The transverse lophs are 
also taller and consequently the cingular and cen- 
tral basins deeper. The protocone and metaconule 
are more approximated than in other species, as 
are the paracone and meiacone. In unworn spec- 
imens of B. parvus the relative cusp heights are 
as reported for B. triradiatus (Turnbull et al. 
1987): protocone exceeds metaconule while 
paracone is subequal to the meiacone. In worn 
M-s of B. parvus the lingual cusps are lower so 
that the paracone exceeds the metacone which 
exceeds the protocone which is subequal to the 
metaconule. This pattern of cusp wear appears to 
be the same in B, brut}'i. 

M^ o\' B. brutyi is similar to M- but is smaller, 
proportionately a little narrower (because the lin- 
gual cusps are less bulbous) and with cusps a lillie 
lower. The posterior cusps are more reduced than 
the anterior cusps and the metaconc, in particular, 
is relatively lower. The metaconule is slightly 
further forward than m M- so thai the postero- 
lingual corner of the tooth is more rounded in 
occlusal view. This feature is similar to the con- 
dition in B. triradiatus and, even more so, to the 
condition in B. parvus. The transverse lophs, pre- 
and post-cingula and their associated basins soon 
wear down to the level of the central basin. M^ is 
most reduced posteriorly in B. parvus and least 
reduced in B. triradiatus. 

M"^ is larger, both relative to other molars and 
absolutely, in B. brutyi than in B. parvus and is 
also less posteriorly reduced. The posterior cusps, 
especially the metaconule, are markedly reduced. 
The anterior cusps, although low and rapidly 



worn, are distinct in unworn teeth and remain 
distinguishable until late wear stages. Although 
the cusps, their associated crests and basins are 
low and quickly levelled, worn M'^s of B. brutyi 
shows more surface morphology than those of fi. 
parvus, in which even newly-erupted M'^s are 
almost featureless. M*^ has 3 roots in B. brutyi. 
Ride (1956) reports a double-rooted M^* in B. 
pan-US but it appears from the specimens exam- 
ined thai the basic condition in B. parvus is a 
3-rooted M"^, perhaps with a reduced number of 
roots in some specimens. 

INTRAGENERIC PHYLOGENETIC 

ANALYSIS 

Thirty-five characters were investigated for 
their potential to contribute to an analysis of the 
relationships between species of Burramys. 
Cercartetus nanus, C. lepidus, C. caudatus and 
C. concinnus were used as the primary outgroup 
since Burramys and Cercartetus are sister groups 
(Archer, 1984; Aplin & Archer, 1987). Tricho- 
surus caninus. T. arnhemensis, T. vulpecula^ 
Spilocuscus maculatus i\\\6 Phalangercarmelitae 
were used as a secondary outgroup because DNA 
hybridisation indicates that burramyids and 
phalangcrids are sister groups (Springer & 
Kirsch. 1 989). Character numbers refer to Table 
5; unnumbered characters are not included in the 
analysis. 

1 . Body si/e. Jaw lengths suggest that B. bruiyi and B. 
wakefieldi were of similar body size. B. iriradiatus and 
B. parx'us are larger and approximately the same size 
as each other. Cercartetus lepidus, the smallest of its 
genus, is also regarded as the most primitive (Archer, 
1984). Phalangerids are larger than burramyids but this 
is likely to be a derived condition; diverse taxa exhibit 
a general tendency for increasing body size over lime 
(Maurer el al., 1992). The small size oi' C. lepidus 
suggests that larger size is apomorphic within 
Burramys. In our discussion of character states, a 
morphological feaiure is regarded as large only il its 
greater size is independent of increased body size. 

2. Robustness. The dentary and maxilla of R pcinnis 
are more slender than those of other Burramys, despite 
being larger. All species of Cercanelus have similarly 
slender jaws. Trichosurus, Spilocuscus and Fhalanger 
are more robust than Cercartetus or B. pan-us, but 
being several limes larger ihan burramyids. thcv do not 
form a useful comparison in this regard. The slender- 
ness of Cercartetus suggests that increased robustness 
is apomorphic in burramyids. 

3. Length of I2-P2 interval. The interval occupied by 
I2, Pi and P2 is longerrelalive to jaw length inB. parvus 



262 



MEMOIRS OF THEQUEHNSLANDMUSBUM 



TABLE 5. Characters and character polarities for imragcneiic phylogenciic analysis of Butramys species. 
Plesiomorpiiic ^iwVc denoted by 0; ? indicates that intbrmalion on character is unavailable. A and B indicate 
alternative derived states. 



Character 


B. brutyi 


B. wakefieldi 


B. iriradiaius 


B. parvus 


1 Body size 








1 


1 


2 RobuBticity 


I 


1 


\ 





3 Length of I2-P2 interved 


1 


1 


2 





4 Length of Ij 





? 


2 


1 


5 Basal thickening of Ii 


1 


? 


1 





6 Number of roots P|_2 





lA 


IB 


p 


7 Arrangement of alveoli Pj.o 


1 


?• 


? 





8 Size of P3 


1 


1 


2 


1 


9 Size disparity between P3 roots 


1 


3 


2 


2 


1 Numberof ridgesp3 


I 


1 


3 


2 


1 1 Curvature of P3 anterior profile 


B 





1 


1 


1 2 Concave/ convex P3 








1 


1 


13 A rched dorsa 1 edge P3 


JO 





1 


2 


1 4 Divergence of P3 from molar row 


I 


i 


2 


2 


1 5 Transverse compression P3 


1 


1 


3 


2 


1 6 Distinct M^ talomd and trigonid 


1 





7 


1 


17 Ml protocorxid position 





1 


? 





18 Ml metaconid position 


t) 





7 


1 


1 9 Relative length lower molars 


2 


2 


1 


t) 


20 Neomorphic cuspid 


1 


? 





1 


21 Loph(id)developmentM2.3/ M*^"^ 


1 


? 





I 


22 No. roots M1.3 








1 


Q 


23 No. roots M4 





a 


lA 


IB 


24 Reduction of M4 


1 


1 





t 


25 Size of maxillary vacuities 

26 Anterior limit P^ 





? 


7 


1 


1 


? 


7 





27 Rotation of upper molar row 


1 


? 


7 





28 Inflation <ii lingual cusps M^ 





1 


6 


1 


29 Lingual displacement of M^ 





? 





1 


paracone 











than in Li, bvutyi or B, niradUitus, with B iriradiaius 
the shonesi. This region is inconiplcic in the holotype 
o\'8. witki'Jleldi but appears to be ubout the same length 
as in B. hnuxi. This interval is relatively long in 
Ci^rcaruins and phalangerida,- indicating that ihis is the 
plesiomorphic slate. 

4. 1] length. 1| is longer in B. parvu.s than B. hrulyi and 
longer again in B, trinuiiatus (unknown in B 
wakefieldi). It is shorter m Cercurlcttts than Burramv.s 
iind is shorter in phalangerids. A long I) is regarded as 
apomorphic. 

5. Thickened base o\ i 1 , In B. hniiy/ and B. iriradiutus 
1 1 is thick basally (thicker in B triradiums) and im- 
mediately begins to taper; approximately half way 
along the exposed portion of the looih it tlnns markedly 
then atlcniiaies to the tip. In B, parvus li tapers grad- 
ually without marked reduction at a particular point. In 
Ccrciineiiis. Tncho\uru\, Spiloatscus and Phalani;er 
1) does not change suddenly rn diameter, suggesting 
that a basally thickened 1 1 is apomorphic. 

Shape of Pi Pi is not known lor B wakefichli or B 



mradidtu.s. In B. brutyi W is small, rounded and similar 
to Pz, In B. panu.s P\ is intermediate between the 
cap-like P: and the slightly elongate, procumbent l2- 
In C. caudatus and C lepidus P| and P: arc both 
bution-like and upright: in C nanus and C. concinnus 
Pi resembles I2 almost as much as P2. Thihosttms and 
/^/laAm.tjtv species have extensive diastenuiia. lacking 
P| and P2 analogous to those oi' burramyids. ll is 
therelorc unclear which state of Pi is more 
plesiomorphic and although this character may be phy- 
iogenctically significant, a satisfaciory polarity assign- 
inenl cannot be made. 

6. Number ofroois P| and P2. Burramys brunt and B. 
pontt.s have double-rooted P| and P2- Burratnys 
inradiatus has a triple- rvuMcd P2 and double- or triptc- 
roolcd P|; the numt^cr o\ rovls is not clear due ti> 
damage m the available material P 1 and P2 each appear 
to have been smglc-n>oied in B. Mukefifldi- P1-P2 ot 
rf/'C(yrf(^fi(-vpossess .sometimes one and sonicitmes two 
roots. P|-2 of Truhosutvs and Phaiun^cr are either 
cMrenicty reduced or absent. OiUgroup analysis does 
not resolve the polarity of this character. The nt)rmal 
marsupial premolar condition is two-rooted .so this is 



NEW OLlGOCENE-MiOCENE /?fy/?/?>^A/y5 FROM RIVERSLEIGH 



163 



taken to be the plesiomorphic condition. Btirramys 
wakejleldi and B. triradiams are interpreted as having 
alternative derived stales. 

7. Arrangement of Pi-2 alveoli. In B. paiTus the alveoli 
of P|-2 arc in a straight line between I2 and P3. In B. 
hrutyi the anterior alveolus o^ ?2 is lingual to its pos- 
terior alveolus and the posterior alveolus o^ P] is lin- 
gual to its anterior alveolus. B. wakefieldi and B. 
triradiams have different numbers of roots lor P1.2 
from B. hrutyi and B pan'us. so their alveoli are not all 
homologous. In all species ofCercareiusihe alveoli of 
P|-2 lie in a straight line: this is also the case for 
Trichosuriis, Spilocuscus and Phakmger (where the 
teeth occur). Linearly-arranged alveoli are therefore 
thought to be plesiomorphic for burramyids. 

8. Size of plagiaulacoid premolar. The sectorial premo- 
lar of Cercanetus and phalangerids (and Mi of C. 
concinnus) is smaller than ihat o( Burramys. It is there- 
fore assumed that an enlarged plagiaulacoid premolar 
is synapomorphic for Burramys and apomorphic 
wilhin the genus. Although P3 of B. pan'us is larger 
than that of B. wakefieldi or B. brutyi, log-scaled plots 
of P.^ buccal crown surface area against jaw length 
(unpubl. data) suggest that P3 of B. pannts is neM 
disproportionately large for its body size. P3 of B. 
triradiatus, on the other hand, departs significantly 
from the line of best fit for P3 size agamsi body size. 
bcingdisproportionately large. P3 of R wakefieldi id\h 
below the line, suggesting that it is disproportionately 
small, but Siudentized residuals do not show its depar- 
ture from the line to be significant. 

9. Relative sizes of anterior and posterior roots of 
plagiaulacoid premolar. Buccally, the posterior root of 
P3 is smaller, relative lo ils anterior root and crown, in 
B. wakefieldi than in other Burramys. The posterior 
root of P3 is smaller (relative to the anterior root and 
the crown) in B. pan'us than in B. hrutyi. The anterior 
root o\' the large P3 oi' B. triradiaius is massive; al- 
though the posterior root is comparatively small, the 
disparity is not as great as that in B. wakefieldi. In 
Cercarteius and phalangerids. the anterior and poste- 
rior roots of the sectorial premolar are subequal; this is 
thought to be the plesiomorphic condition. 

10. Number of ridges on plagiaulacoid premolar. In B. 
brutyi and B. wakefieldi there are 5 or 6 ridges on each 
of the buccal and lingual faces of P3 and 5 or 6 
associated dorsal cuspules. The lack of posterior and 
weakness of anterior cuspules in the holoiypc o\' B. 
wakefieldi appears to he the result of extreme wear on 
the formerly serrated tooth. In B. pan'us there are com- 
monly 7 ridges and cuspules and in B. ihradiaius, 9. 
Phalangerids with smaller, unridged P3s are thought to 
he more plesiomorphic than lho.se with larger, ridged 
P3s (Flanncry et al.,1987); all have fewer ridges and 
cuspules than Burramys. Cercarteius nanus and C. 
caudafus have a single sharp dtirsal cusp on the secto- 
rial prenK)lar and C. concinnus one main cusp on its 
premohiriform Mj. A larger number of ridges and 



cuspules is regarded as more derived wilhin Burramys 
and a synapomorphy of the genus. 

1 1. Curvature of anterior profile of P3. In lateral view, 
P3 of fi. u'aAry7f/r// and B. /?n/rv7 has a relatively straight 
(approximately vertical) anterior profile. In B. 
triradiatus and B. imrx'us the crown expands anteriorly 
to produce a curved profile. The sectorial P3 of 
Cercanetus does not curve forward anteriorly (al- 
though the aulapomorphic premolariform Mi of C. 
concinnus docs). Anterior curvature may be associated 
with increased P3 size, with enlargement having been 
achieved by anterior extension of the crown. However 
P3S of T. caninus and T. vulpecula. which are curved, 
are smaller than those of Spilocuscus and Phalanger, 
which are less curved. Size and curvature are therefore 
not necessarily linked. It is possible that Miocene P3S 
represent primary enlargement of the tooth without the 
functional elaboration of other species, in which the 
inllated anterior edge may disperse stress, increase 
occlusal area, or perform some other function. A 
curved anterior profile is regarded as apomorphic 
within Burramyidae. 

12. Lingual concavity/buccal convexity of P3. The P3 
blade of Burramys is concave lingually and convex 
buccally ( particularly anteriorly ). The contrast between 
lingual and buccal curvature is least pronounced in B, 
wakefieldi and B. brutyi and more pronounced in B. 
tmr\'us and slightly more in B. triradiatus. As with 
anterior profile curvature, this feature occurs in 
Tricfiosurus hut not in Cercanetus, Spilocuscus or 
Phalanger. It is regarded as apomorphic. 

13. Arching of dorsal edge of P3. The dorsal edge of 
P3 is arched in B. pan'us: in the other species it is 
straight, but in B. triradiaius there is a slight curvature 
at the anterior end of the blade. The sectorial teeth of 
Cercanetus do not have a dorsal blade edge homolo- 
gous w^ith that o( Burramys and so do not provide a 
useful comparison. The dorsal edge of P3 is straight in 
phalangerids and this is assumed lo be the plesio- 
morphic condition. 

14. Divergence of P3 from anteroposterior axis of 
molar row. In Burramys, the longitudinal axis of P3 
departs from the ramus such that it forms an angle with 
the anteroposterior molar row axis. This angle is largest 
in B. wakefieldi and is larger in B. pan'us and B. 
triradiaius than in B, brutyi. In Cercarteius the longi- 
tudinal axis of the lower sectorial tooth is parallel to the 
anteroposterior axis of the molar row and wilhin 
phalangerines. a more oblique placement of P3 is re- 
garded as apomorphic (Flannery et al., 1987). Diver- 
gence of P3 from the anteroposterior axis of the molar 
row is a synapomorphy of Burramys: w\{h\n Burramys, 
the plesiomorphic condition is taken to be a less diver- 
gent P3. 

15. Transverse apical compression of P3. In anterior 
view, the crown of P3 of Burramys tapers from the 
base, attenuating dorsally then terminating apically 



264 



MEMOIRS OF THE QUEENSLAND MUSEUM 



with a serrated longitudinal median ridge. This trans* 
verse apical compression is least pronounced in the 
Miocene species and most pronounced in B. 
triradiatus. Crowns of the sectorial premolars of 
Cercartetits, Trichosurus, Spilocuscus and Phakmger 
are less attenuated than those of Burramys. Increased 
dorsal transverse compression is synapomorphic for 
Burramys. Laterally compressed P3s are regarded as 
more derived than those with thicker apices. 

16. Distinction of talonid and trigonid of M|. In B. 
wakefieldi the talonid and trigonid of Mi are clearly 
demarcated in occlusal view by lingual and buccal 
indentations. In B. brutyi and B. parxnts the talonid and 
trigonid are less distinct. Mi is not known for B. 
triradiatus. Taionids and trigonids are more distinct in 
Cercartetus than in Burramys, indicating the 
plesiomorphic stale. The fused talonid and trigonid 
departs further from primitive tribosphenic morphol- 
ogy. Alternatively, the structure of Mi in B. wakefieldi 
could be auiapomorphic, with the crests defining the 
talonid and trigonid functioning primarily as buttresses 
for the anterolingual crests which may, in this animal, 
have extended the function of P3. However, the former 
hypothesis is preferred. 

17. Lingual displacement of protoconid of Mi. The 
protoconid of Mi is displaced further lingually in B. 
wakefieldi than in B. brutyi or B. pairus so thai in B. 
wakefieldi the crests associated with P3 and the Mi 
protoconid, mctaconid and entoconid form an almost 
straight line. The position of the protoconid is variable 
in Cercartetus and phalangerids. In the primitive 
tribosphenic molar, the protoconid is a buccal cusp, so 
lingual displacement is regarded as apomorphic. 

18. Anterior displacement of metaconid of Mi. The 
paraconid is absent in Burramys and the most anterior 
lingual cusp is the metaconid. In B. par\'us the 
metaconid is more anterior than in B. wakefieldi or B. 
brutyi, narrowing the gap in the P3-M1 crest. In the 
Phalangeridac and Cercartetus position of the 
metaconid relative to the protoconid is variable. Oul- 
group analysis docs not resolve the polarity of this 
character. The metaconid of B. pan'us occupies the 
position that in a plesiomorphic (tribosphenic) molar 
would have supported the paraconid, so the anteriorly 
displaced metaconid is regarded as apomorphic. 

Inclination of Mi trigonid against P3. The trigonid of 
Ml rises more steeply against the posterior face of P3 
in B. brutyi and B. wakefieldi than in B. parvus. Neither 
Cercartetus nor phalangerids give a clear indication of 
the polarity of this character. It has developed a number 
of times in phalangerids and pilkipildrids and is prob- 
ably homoplasious. 

1 9. Relative length of lower molars. M2-4 o\' Burramys 
differ in their lengths (relative to widths) such that B. 
brutyi <B. triradiatus <B. par\us. M2-4 are not known 
for B. wakefieldi but judging by their alveoli, they were 
of similar proportions to those of B. brutyi. In 



Cercartetus, Trichosurus, Spilocuscus and Phalanger, 
the molars are relatively long, implying that this is the 
plesiomorphic condition. 

20. Ncomorphic cuspid at intersection of 
postmetacristidand preentocristidof M2-3. In B. brutyi 
and B. parx'us there is usually a small neomorphic 
cuspid approximately halfway along the lingual mar- 
gin of the crown, at the junction of the posimetacristid 
and the preentocristid. This cuspid is not present in the 
few available lower molars of B. triradiatus, nor in 
Cercartetus. Trichosurus, Spilocuscus or Phalanger, 
suggesting that it is apomorphic m the Burramyidac. 

Lingua! cusps skewed ahead of buccal cusps of M2-3- 
In B. triradiatus, the lingual cusps of M2-3 arc ahead 
of the buccal cusps, skewing the sides of the teeth 
slightly. This skew is less evident in B. brutyi and 
slightly less again in B. parrus. Cercartetus caudatus 
is about as skewed as B. parvus and is the least skewed 
of species of Cercartetus, with C. namis and C. con- 
cinmis showing about the same, increased degree of 
skew. The amount of skew on the molars is variable 
within phalangerids, ranging from very minor to quite 
pronounced. Outgroup analysis gives no clear indica- 
tion of whether skewed molars are plesiomorphic or 
derived in Burramys. 

2 1 . Transverse loph(id)s of M:>-3 and M^"^. The trans- 
verse lophs and lophids of M-'-^ and M2-3 are more 
complete in B. triradiatus than in B. brutyi or B. parvus, 
such that the central basins and the pre- and post-cingu- 
lar basins of the teeth are deeper and more clearly 
deHncd in the Hamilton species. Cercartetus lacks 
transverse loph(id)s but this is probably apomorphic 
for the genus; lophs and lophids are w ell formed on the 
molars of the more plesiomorphic phalangerids. 
Burramys triradiatus is thought to be relatively 
plesiomorphic in possessing more complete molar 
lophs and lophids. 

22. Number of roots Mi -3. M 1.3 are double-rooted in 
B. wakefieldi, B. brutyi and B. panus, but in B. 
triradiatus dTQ 3-rooted, TurnbuU ct al. ( 1 987) regarded 
the 3-rooted condition as a plesiomorphic retention. 
However, Cercartetus, phalangerids and virtually all 
marsupials have 2-rooled molars, making the 
plesiomorphic retention of 3-rooted lower molars by /^. 
triradiatus seem unlikely. The 3-rooted lower molars 
of R triradiatus are interpreted as autapomorphies. 

23. Number of roots M4. M4 is single-rooted in B. 
pairus. double-rooted in B, wakefieldi and B. brutyi, 
and has 3 roots in B. triradiatus. M4 of Cercartetus 
(where it occurs) and phalangerids has 2 roots. The 
single-rooted and three-rooted M4 of S. parvus and B. 
triradiatus irespccuwdy) are interpreted as alternative 
apomorphic stales derived from a 2-rooted 
plesiomorphic condition. 

24. Reduction of M4. M4 is most reduced in B. parx'us 
and least reduced in B. triradiatus, with the Miocene 



NEW OLIGOCENE-MIOCENE BURRAMYS FROM RIVERSLEIGH 



265 



species intermediate. Cercartetus lepidits and C. 
caudatiis (apparently the most plcsiomorphic 
Cercanetus) have M4. though reduced: in C. nanus and 
C. conc'mnus M4 is absent. In Tnchosurus M4 is sub- 
equal to Ml -3; in Spilocuscus and Phakmger (which 
are generally more derived than Trichosurus) it is 
slightly smaller than the anterior molars. Reduction of 
the posterior molars occurs frequently and indepen- 
dently. Since primitive members of both outgroups 
have less reduced M4, and since reduction of the molar 
row posteriorly is commonly a derived stale, more 
reduced IVI4S are interpreted as apomorphic. Although 
M4 reduction correlates with M4 root number in 
Burramys, it is treated as a separate character since, as 
demonstrated by the relative sizes and number of roots 
of Ml -3 in the different species, there is not necessarily 
a connection between molar size and number of roots. 

25. Enlarged maxillary vacuities. Maxillary vacuities 
are larger in B. pan'us than in B. brutyi. The vacuities 
o{ Cercartetus do not resolve this character. Vacuities 
are less extensive in phalangehds than in B. parvus, so 
a less evacuated palate is regarded as plcsiomorphic. 

26. Anterior limit ofP-^ relative to zygomatic arch. P-^, 
and therefore the anterior of the upper molar row, is 
further forward on the maxilla relative to the jugal 
portion of the zygomatic arch in B. parx'us than in B. 
bruiyi. In C. concinnus and C. caudaius the teeth are 
further forward than in Burramys; in C. nanus (and 
possibly also C. lepidus) the anterior extent of the teeth 
is similar to that in B. pan'us. In S. maculatus, P. 
cannelilae, T. arhemensis and T. vulpecula. the 
cheekteeth commence further forwards. The polarity of 
this character is not immediately evident, particularly 
as there are a variety of slates within Cercartetus: 
however the anterior disposition of the teeth in 
phalangerids would argue for that being the 
plcsiomorphic condition. 

Enlarged P-^ cingulum. The P3 cingulum is slightly 
more developed in B. parx'us than B. brutyi and signif- 
icantly more pronounced in B. triradiatus. It possibly 
developed in conjunction with the enlargement oi P-^ 
and the generation of greater bite forces at the P3s. 
functioning as a stopper for P3 during premolar func- 
tion (as indicated by posterolingual wear facets that 
stop abruptly at the cingulum in B. parvus and B. 
triradiatus) and also protecting the gums from hard 
food particles sectioned by the premolars. P-^s of 
Cercartetus and phalangerids are not sulTiciently sim- 
ilar to those oi Burramys io have homologous cingulae. 
so outgroup comparison cannot polarize this character. 
If the enlarged cingulum is linked to P^ size it is not an 
independent character . 

Anterior attenuation of P-\ In dorsal view, P^ is more 

ovoid and in particular, more attenuated anteriorly, in 
B. triradiatus and in B. parvus than in B. brutyi. P-^ is 
insufficiently similar in Cercartetus and phalangerids 
to Burramys to be useful in determining polarity of this 
character. Anterior attenuation may be associated with 



P^ size and is probably linked to anterior inilation of 
P3; it is not treated as an independent character. 

27. Posterobuccal rotation of molars rotate around 
maxilla. Uppermolarrow rotation is greater in fi./jn/rvi 
than in B. pan'us. The upper molars do not rotate 
buccally in a posterior direction in Cercartetus, but 
they do in phalangerids examined, tJsing Cercartetus 
as the primary outgroup and applying the principle of 
commonality, the rotating molar row of fi. brutyi would 
be interpreted as more derived than the dental arcade 
of B. parvus. 

Pronounced anterobuccal cingular basin M^ In B. 
brutyi there is a cingular basin on the anterobuccal 
comer of Mr, in B. triradiatus there appears to be a 
narrow cingular pocket and in B. pan'us the pocket is 
little more than a sloping cingular shelf. Inflation of the 
anterobuccal comer of M^ is a synapomorphy for 
Burramys. It seems that the degree of definition of the 
anterobuccal pocket is inversely related to lingual shift 
of the paracone. Therefore, it is not treated as a separate 
character. 

28. Inflation of lingual cusps of M'. No maxillary 
material is available for B. wakefieldi. In each of the 
other species of Burramys, the lingual side of M' is 
enlarged by a protoconule on the lingual margin, ante- 
rior to the protocone. Both protoconule and mctaconule 
are more inflated in B. pan-us than in other species and 
in association with this, lingual cusps of B. panus lie 
closer to the lingual edge of the tooth than in the other 
species. There is no protoconule in C. lepidus or C. 
caudaius: it is present (relatively undeveloped) in C. 
concinnus and perhaps in a rudimentary state in C 
imnus. There is no protoconule in Trichosurus, 
Strigocuscus or Fhalanger. An enlarged protoconule 
is considered apomorphic. 

29. Lingual displacement of paracone of M' . The M' 
paracone of 5, parxus is displaced lingually so that the 
ectoloph is oblique with respect to the anteroposterior 
axis of the tooth. In B. brutyi and B. triradiatus the 
ectoloph is approximately parallel to the tooth axis, 
with the paracone more buccal. The M ' paracone is not 
displaced lingually in Cercartetus or phalangerids, in- 
dicating that this is the plcsiomorphic condition. 

A Wagner analysis was performed using both 
ACCTRAN and DELTRAN algorithms of PAUP 
(Swafford, 1989). Wagner parsimony allows re- 
versal or convergence to construct trees with the 
fewest steps. Where reversal or convergence 
would produce an equally parsimonious solution, 
ACCTRAN accelerates character transforma- 
tions, favouring reversal, whereas DELTRAN 
delays transformations, favouring convergence 
(Wiley et al., 1 99 1 ). Characters were ordered and 
a hypothetical ancestral Burramys, having all 
character states 0, was used to root the analysis. 

ACCTRAN (Fig. 10) or DELTRAN opiimisa- 



266 



MEMOIRS OF THE QUEENSLAND MUSEUM 



1^ 



CO 



a 1-2 

[6] O^IB 
[8] 1-2 
[ipl 2 — 3 
^ 2-3 
1—0 
1—0 
[22| 0-1 
[231 0— 1A 
[241 1-0 



^ 

m 






--[2] 1 — 

[T[ 1—0 

1] 1^0 
m 1-2 

lH 1-0 

23] 0— IB 

24] 1-2 

23 0—3 



■[6] 0— 1A 
[T| 2— 3 
E3 2-3 

[H 1-0 

\V7\ 0-1 



H] 0-1 

[2§ 0—1 
[27| 0-1 



-a 0-1 
[Tol 1 — 2 
[n] 0-1 
[lU 0—1 
M 0-1 
[m 1-2 

[18] 0-1 
[191 2—1 



--H 0-1 
[9] 1-2 
[g 1-2 
i25l 0—1 



0—1 
0—1 
0-1 
0—1 
0—1 
0—1 
0—1 
0—1 
0—1 
0-2 
0-^1 
0—1 
0—1 



FIG. 10. A phylogenelic hypothesis of intrageneric 
relationships of Burramys. Apomorphies listed at 
nodes; character numbers in boxes refer to Table 5. 
Character stale transformations indicated by arrows. 

tion generated a single most parsimonious tree. 
The topology of this tree is identical for both 
algorithms. Burramys parx'us and B. triadiatus 
form a clade to which B. wakefieldi is the 
plesiomorphic sister group; B. brutyi is the 
plesiomorphic sister group to a clade containing 



all other species of Burramys. For some charac- 
ters, the path of transformation differs depending 
upon whether transformation is accelerated or 
delayed. There are several convergent character 
states in the DELTRAN tree, no convergences 
and more reversals in the ACCTRAN tree. When 
transformation is delayed the following character 
states arise convergently in B. brutyi and B. 
parvus: loph(id)s of M2-3 reduce; neomorphic 
cuspid appears on M2-3; and Mi talonid and 
trigonid become less distinct. Basal thickening of 
Ii occurs independently in B. brutyi and B. 
triradiatus. The relative length of the lower mo- 
lars decreases independently in B. brutyi and B. 
wakefieldi. With delayed transformation B. 
par\'us reverses to a more plesiomorphic state of 
reduced robusticity and relatively long molars, 
and the relative size of M4 in B. triradiatus in- 
creases secondarily. These reversals also occur 
when transformation is accelerated, as do the 
following: in B. parvus and B. triradiatus the 
relative length of lower molars increases (to a 
greater degree in B. pan'us)\ in B. pan'us the I2-P2 
interval increases; in B. triradiatus loph(id)s de- 
velop on M2-3 and the neomorphic cuspid disap- 
pears from M2-3; and in B. wakefieldi the talonid 
and trigonid of Mi are relatively distinct from one 
another. 

Although a single most parsimonious tree was 
generated by this analysis, another tree only one 
step longer placed B. wakefieldi as the 
plesiomorphic sister-group of the other three spe- 
cies, and B. brutyi as the plesiomorphic sister- 
group of the B. triradiatus + B. pannis clade. A 
bootstrap analysis using a branch and bound 
search with 100 repetitions, to place confidence 
estimates on clades (from ACCTRAN) found the 
node defining the B, triradiatus + B. parvus clade 
to be supported 84% of the time, but the node 
separating B. brutyi and B. wakefieldi occurred in 
less than 50% of repetitions. Using DELTRAN 
the B. triradiatus + B. parvus clade was supported 
78% of the time, and the node separating the other 
3 species from B, brutyi was supported by 55% 
of repetitions. In both cases, the node separating 
B, brutyi and B. wakefieldi is poorly resolved. 

DISCUSSION 

Burramys brutyi is the only species of 
Burramys at Riversleigh and is not known else- 
where, it is represented by > 1 50 specimens from 
23 Sites in Systems A, B and C; it is one of the 
most widely distributed (spatially and tempo- 
rally) marsupials at Riversleigh. Its earliest oc- 



NEW OLIGOCENE-MIOCENE BURRAMYSFROM RIVERSLEIGH 



267 



currence at late Oligocene (Myers & Archer, 
1997) White Hunter Site is of similar age to the 
type locahty o\B. wakefieldioxx Mammelon Hill, 
Lake Palankarinna, South Australia (Woodbunic 
eta!., 1993). 

Metric analyses did not reveal any significant 
size variation between sites; variation within sites 
being as great as between sites. This persistence 
in unchanged form from the late Oligocene 
through much of the Miocene suggests an un- 
usual degree of ecological stasis for the species. 
Fossil Biirramys in Victoria, South Australia 
and NW Queensland shows that small existing 
populations of B. parvus arc remnants of a pre- 
viously more diverse and far more widespread 
lineage, now apparently in decline. This fact 
urges particular conservation concern for the ex- 
tant species. Although populations of fi. pan'us 
are apparently stable, they are threatened both by 
habitat disturbance and greenhouse warming, 
which could jeopardise their ability to survive 
(Geiser & Broome 1993). 

ACKNOWLEDGEMENTS 

Comparative material was made available by J. 
Dixon, L. Frigo, T. Rich and E. Thompson, Mu- 
seum of Victoria; N. Pledge, South Australia 
Museum; and J. Wombey, C.S.LR.O., 
Gunghalin. The vital support of the following 
organisations is also gratefully acknowledged: 
the Australian Research Council (grants to M. 
Archer); the National Estate Grants Scheme 
Queensland (to A. Barlholomai and M. Archer); 
the Department of Environment, Sports and Ter- 
ritories; the Queensland National Parks and 
Wildlife Service; the University of New South 
Wales; IBM Australia Ply Ltd; ICI Australia Pty 
Ltd; the Australian Geographic Society; Wang 
Australia Pty Ltd; the Queensland Museum; the 
Australian Museum; Ml Isa Mines Ply Ltd; Sur- 
rey Beatty & Sons Pty Ltd; the Riversleigh Soci- 
ety Inc.; the Royal Zoological Society of New 
South Wales; the Linnean Society of New South 
Wales; and many private supporters. 

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TWO NEW BALBARINE KANGAROOS AND LOWER MOLAR EVOLUTION 

WITHIN THE SUBFAMILY 

B.N. COOKE 

Cooke, B.N. 1997 06 30; Two new balbarine kangaroos and lower molar evolution within 
the subfamily. Memoirs of the Queensland Museum 41(2):269-280. Brisbane. ISSN 0079- 
8835. 



Lower Jaws and teeth of Nambaroo couperi sp. nov. and Wumroo dayamayl gen. et sp. nov.. 
fossil balbarine kangaroos from the late Oligocene White Hunter Site of Riversleigh. are 
described. Mi trigonid cuspid homology in Hypsiprymnodon is re-interpreted such that a 
reduced protoconid is recognised, the anterobuccal cuspid is regarded as the protostylid and 
the anterolingual cuspid as the metaconid. The evolution of lophodont lower molars within 
Balbarinae is examined on the bases of this interpretation and information supplied by the 
new species. O Riversleigh. kangaroo, Balbarinae. Nambaroo, Wururoo, cuspid homology. 
lophodonty. 

B.N. Cooke, School of Life Science, Queensland University of Technology. GPO Box 2434, 

Brisbane, Queensland 4001. Australia: 18 December 1996. 



The Balbarinae Flannery, el al., 1982 was 
erected for a group of fossil macropodids in 
which the M| protolophid is markedly com- 
pressed. It assumed phylogenetic significance 
when Flannery (1989) proposed that baibarines 
were ancestral to both slhenurines and 
macropodines. 

Baibarines form a major component of the fos- 
sil kangaroo fauna of Riversleigh and appear to 
have had a wide distribution in the Oligocene- 
Miocene of Au.stralia. Three genera and 8 species 
have so far been named, but it is more diverse than 
this. Bcdbaroocamfieldensis Flannery et al.. 1 982 
is known from Bullock Creek and Balharoo sp. 
Flannery et al. (1982) from the Kangaroo Well 
Local Fauna, both in the Northern Territory. 
Three species of Nambaroo Flannery & Rich, 
1 986 were described from the Tarkarooloo Local 
Fauna of South Australia. Unnamed baibarines 
have been reported by Flannery (1989) from the 
Kuljumarpu Local Fauna of South Australia and 
Woodburne et al. ( 1993) from the Eladunna For- 
mation of South Australia. Riversleigh baibarines 
include Bcdbciroo gregoriensis Flannery et al.. 
1982 and 3 species of Ganawamaya Cooke, 
1992. The present paper includes descriptions 
based on lower jaws and teeth of a new species 
of Nambaroo and a new genus and species of 
balbarine. 



outlined by Szalay (1969), following Ride 
(1993). Terminology follows Van Valen (1966), 
Szalay (1969) and Butler (1990). However, the 
'anterior cingulid' is restricted in use to that com- 
ponent of the macropodoid lower molar anterior 
cingular shelf lying lingual to the anteriorly di- 
rected paracristid. Trecingulid' is refers to that 
component lying buccal to the paracristid. Van 
Valen (1966, 1994) used 'precingulid' for the 
anterior cingular shelf anterobuccal to the 
paracristid of plesiomorphic mammalian lower 
molars. The distinction is made here because 
lingual and buccal components of the 
macropodoid anterior cingular shelf are demon- 
strably of different origins. The buccal compo- 
nent is the more plesiomorphic since it occurs in 
plesiomorphic baibarines such as Nambaroo 
couperi sp. nov. More derived baibarines such as 
Wiiriiroog^yx. nov. and fifl//7^roo demonstrate the 
development of the neomorphic lingual compo- 
nent via lingual displacement of the paraconid 
and lingual extension of the paracristid. Supra- 
generic classification follows Aplin & Archer 
(1987). QMF denotes Queensland Museum fossil 
collection catalogue numbers. Measurements arc 
in millimetres. 



METHODS 



SYSTEMATICS 



Molar homology follows Luckett (1993). Pre- 
molar homology follows Flower (1867). Homol- 
ogy of molar structures has been determined by 
reference to a generalised tribosphenic pattern. 



Family MACROPODIDAE Gray, 1821 
Subfamily BALBARINAE Flannerv. 

Archer & Plane. 1982 
Nambaroo Flannery & Rich, 1986 



270 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Nanibaroo couperi sp. nov. 
(Figs 1,2, 5 A; Table I) 

DIAGNOSIS. Nambaroo with a hypoconulid at the 
posterior, buccal base of the entoconid on Mi and 
marked convexities along the lateral margins adjacent 
to the ends of the interiophid valley on all lower molars 
except M4. 

MATERIAL. Hololype QMF3040I. a partial right 
dentary consisting of the entire horizontal ramus, most 
of the angular process and portion o'i the ascending 
ramus to the level of the damaged condyle. P3 and M 1-4 
are preserved; from White Hunter Site, Hal's Hill, D 
Site Plateau, which has been correlated (Myers & 
Archer, this volume) with the Ngama Local Fauna from 
the Tirari Desert which Woodbume et al. (1993) have 
shown to be late Oligocene, about 24 to 26My. 

ETYMOLOGY. For Patrick Couper, Queensland Mu- 
seum, for his assistance during the course of this re- 
search. 

DESCRIPTION. The holotype is a fragment of a 
right dentary consisting of the entire horizontal 
ramus, most of the angular process and portion of 
the ascending ramus to the level of the damaged 
condyle(Fig. 1 A,B). Dorsal edgeof the diastema 
delineated by a ridge with matrix-filled alveolus 
for a very small I2 or analogous tooth at anterior 
end. Horizontal ramus twisted, with mesial sur- 
face inclined slightly dorsally below P3 and 
slightly ventrally below M4. Mandibular sym- 
physis extending as far posteriorly as the level of 
the anterior margin of P3. On the buccal surface 
the anterior mental foramen located below and 
slightly anterior to P3, with a much smaller pos- 
terior mental foramen below the hypolophid of 
M2. Horizontal ramus deepest below Mi, with its 
zone of most ventral protrusion below M3. Ven- 
tral margin straightest below P3-M3, curving up- 
wards below the diastema and more steeply so 
posterior to M3. Buccal margin of the masseteric 
fossa straight so that the entrance to the masse- 
teric canal is 'D' shaped in cross section. Ventral 
margin of the masseteric fossa low on the ramus, 
well below the level of the molar row. Inferior 
dental canal recessed into the lingual wall of the 
masseteric canal but not partitioned from it. Be- 
cause of the confluence of the two canals, forward 
extent of the penetration of the masseier difficult 
to determine, but the gradient of anterior canal 
constriction suggesting insertion no further for- 
ward than M3- Lingual border of the angular 
process in the same vertical plane as the lingual 
margin of the horizontal ramus but the angular 
process extending more posteriorly than the 



ramus. Large portion of the floor of the pterygoid 
fossa lost. 

Ascending ramus rising at 100° relative to the 
plane of the molar row. Condyle situated 9.4mm 
above the molar row, a transversely elongate 
structure, broader lingually, tapering to the buc- 
cal side, obliquely inclined to the plane of the 
ascending ramus. 

Dentition. Molar row straight in both occlusal and 
lateral view. P3 Hexed slighdy buccally out of 
alignment with molar row. Occlusal surfaces of 
anterior molar lophids inclined slightly buccally. 
Those of more central molars more or less hori- 
zontal. Hypolophid of M4 inclined lingually. 
Slight increase in molar size posteriorly. 

P3 gracile, short and blade-like with horizontal 
occlusal margin. In occlusal view with an ellipti- 
cal outline with the occlusal crest occupying ap- 
proximate midline, although curving lingually 
posteriorly. Five cuspids, of which most posterior 
largest, occupying the occlusal crest. Lingual and 
buccal transcristids associated with each cuspid, 
although those on lingual surface partly obscured 
by wear. Anterior and posterior margins of crown 
delineated by vertical cristids. 

In occlusal view Mi with a rounded anterior 
margin and lateral convexities low on the crown 
adjacent to the ends of interiophid valley. Pro- 
tolophid shorter than hypolophid: protoconid po- 
sitioned on approximate midline of tooth, 
Protoconid taller than metaconid, buttressed 
buccally by a prolostylid about same height as 
metaconid. Paracristid running almost directly 
forward to anterior margin where it meets ante- 
rior edge of aprecingulid which descends steeply 
to buccal margin of tooth. No anterior cingulid, 
trigonid basin open anterolingually. Cristid ob- 
liqua inclined slightly lingually, descending an- 
terior face of the hypoconid, turning anteriorly to 
cross interiophid valley and terminating al base 
of protostylid. 

Entoconid taller than hypoconid. Occlusal cresl 
of hypolophid forming shallow 'V with lingual 
arm steeper than buccal. Low point of hypolophid 
slightly lingual of midline. Preentocrisiid de- 
scending to interiophid valley Hoor from apex of 
entoconid. Wedge shaped prominence at poste- 
rior base of entoconid, interpreted here as a 
hypoconulid because its position corresponding 
to that occupied by hypoconulid in other marsu- 
pials and, as in these animals, contacted by the 
posthypocristid. Short lingually displaced, diag- 
onal poslhypocristid and hypoconulid forming 
posterobuccal border of small fossette in poste- 



TWO NEW BALBARINE KANGAROOS 



271 




FIG. 1. QMF3040I, Holotype oi Nambaroo couperi sp. nov. A, 
occlusal view. Scales = 10mm. 



buccal view. B, lingual view. C, stereopairof 



rior face of entoconid. Rounded, lingual border 
probably representing postenlocristid. From base 
of hypoconulid a hypocingulid extending across 
about half width of posterior base of hypolophid. 

M2 larger than Mi, approximately rectangular 
in outline, but with lateral convexities of crown 
base adjacent to ends of interlophid valley. Pro- 
tolophid and hypolophid about equal in length, 
lingual cuspids taller than buccal cuspids. Occlu- 
sal margins of both lophids shallowly concave. 
Paracristid running anlerolingually to anterior 
margin from which enamel has been lost, al- 
though sufiicient remains to indicate that an an- 
terior cingulid ran between paracristid and 
lingually positioned premetacristid. Short, stee- 
ply sloping precingulid extending from anterior 
end of paracristid to buccal margin of the tooth. 
Cristid obliqua runing anterolingually across 
interlophid valley to join posterior base of pro- 



tolophid at about midline. Preenlocristid short, 
not reaching floor of interlophid valley. 
Postentocristid runing vertically down posterior, 
lingual edge of entoconid to meet low and some- 
what irregular hypocingulid which runs trans- 
versely across half width of the posterior face of 
hypolophid. No posthypocristid. 

M3 similar to M2 but metaconid and hypoconid 
more nearly equal in height, hypocingulid less 
developed and cristid obliqua interrupted adja- 
cent to base of protolophid. 

M4 differing from more anterior molars in the 
following: protolophid longer than hypolophid; 
no obvious convexities of crown base adjacent to 
ends of the interlophid valley; some wrinkling of 
the enamel within trigonid basin; precingulid less 
obvious; rounded postmetacristid descends pos- 
terior face of metaconid to floor of interlophid 
valley; cristid obliqua interrupted before contact- 



272 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 1 . Dental parameters for QMF303401 , Holotype of Nambaroo couperi sp. nov. 



Cat. No. 


P3 


Ml 


M-) 


M^ 


M4 


1 


mw 


h 


1 


pw 


hw 


I 


pw 


hw 


I 


pw 


hw 


1 


pw 


hw 


1 QMF30401 


5.2 


2.9 


3.9 


5.2 


2.6 


3.3 


5.4 


3.5 


3.9 


5.6 


3.9 


4.1 


5.4 


3.8 


3.6 



ing base of protolophid and ridge of enamel runs 
transversely across floor of the interlophid valley 
from anterior end of cristid obliqua to buccal 
margin; hypoconid taller than entoconid (which 
is slightly damaged); no distinct posteniocristid; 
hypocingulid in form of rounded, transverse 
prominence crossing posterior face of hypolopid. 

DISCUSSION. N. couperi is similar in size to N. 
saltavus, N. tarrinyeri and N. noviis which were 
described by Flannery & Rich (1986) from the 
Tarkarooloo Local Fauna of South Australia. 
Apart from the plesiomorphic hypoconulid, N. 
couperi is in other aspects of talonid morphology, 
more derived than N. saltavus. It lacks a 
postentocrislid which is present in N. saltavus, 
and has a well-developed hypocingulid crossing 
buccally from the hypoconulid on the posterior 
base of the hypolophid, present also in N. tar- 
rinyeri and N. novus but undeveloped in N. 
saltavus. The M 1 cristid obliqua contacts the pro- 
toslylid of M couperi as it does in N. saltavus, a 
condition which Flannery & Rich ( 1 986) consid- 
ered plesiomorphic for macropodoids. In A'. 
novus the cristid obliqua contacts the base of the 
protoconid and in N. tarrinyeri the cristid obliqua 




64.3 




FIG. 2. Dimensions of QMF3040I. holotype of 

Nambaroo couperi sp. nov. A, buccal view. B, lingual 
view. 



bifurcates and contacts both protostylid and pro- 
toconid. Ml Irigonid morphology in N. couperi 
is most similar to N. novus in that both have an 
anteriorly directed paracristid, lack a paraconid 
and have a protostylid which is closely associated 
with the protoconid. 

Wururoogen. nov. 

TYPE SPECIES. Wururoo dayamayi sp. nov. 

DIAGNOSIS. Balbarines with a large, trenchant 
P3 and a posterobuccally inclined enamel ridge 
(the 'protostylid crest' ) descending from the apex 
of the protoconid of Mi. 

ETYMOLOGY. Gulf coast aboriginal vn/n<. a long 
lime ago (Breen, 1 981), and rcc a common Australian 
diminutive for 'kangaroo'. Masculine. 

Wururoo dayaniayi sp. nov. 
(Figs 3, 4, 5B; Table 2) 

DIAGNOSIS. As for genus. 

MATERIAL. Holotype QMF19820, A fragment of the 
horizontal ramus of a right dentary, extending from the 
anteriorendoflhcdiastemato the posterior of the molar 
row and preserving P3 and M1-4. from White Hunter 
Site, Hal's Hill. D Site Plateau, which has been corre- 
lated (Myers & Archer, 1997) with the Ngama Local 
Fauna from the Tirari Desert which Woodbume ct al. 
(1993) have shown to be late Oligocene, about 24 to 
26My. 

ETYMOLOGY. Waanyi daya. chop; mayi tooth, for 
the tall, robust plagiaulacoid premolar of the holotype. 

DESCRIPTION. Only small portion of anterior 
margin of ascending ramus present, remainder of 
dentary posterior to M4 lost. Horizontal ramus 
deepest below M3/M4, tapering gently anteriorly, 
ramus twisted so that mesial face inclines dorsally 
below P3 and slighly ventrally below M4. Low 
ridge running length of dorsal margin of very 
shon diastema. Mental foramen well below and 
slightly anterior to P3, small posterior mental 
foramen below hypolophid of M2. Mas.sctcric 
canal elliptical in cross section, extending for- 
ward at least as far as anterior of M2, from which 
position anterior of canal blocked by undissolved 



TWO NEW BALBARINE KANGAROOS 



273 



matrix. A narrow canal, Ihe inferior dental canal 
or branch thereof, partitioned from lingual side of 
masscleric canal by ihin lamina of bone anterior 
to M4. Mandibular symphysis extending posteri- 
orly to below mid point of P3. Molar row concave 
in lateral view and straight in occlusal view. 
Molar size increasing from M| to M3 but M4 a 
little smaller than M3. 

Dentition. P3 large, robust, plagiaulacoid, occlu- 
sal edge well above occlusal plane of molar row, 
long axis flexed slightly buccally out of align- 
ment with line of molar row. Viewed occlusally 
with a convex buccal margin and slightly concave 
lingual margin. Lingual face of crown inclined 
at steeper angle than is buccal. Occlusal margin 
on approximate midline for most of length, but 
slightly lingually displaced at posterior end. Six 
cuspids on occlusal margin, all bul most posterior 
having associated lingual and buccal trans- 
cristids. Cristids descending anterior and poste- 
rior margins of crown from corresponding 
cuspids. 

Ml roughly rectangular in occlusal outline bul 
with distinct lingual convexity in crown ba.se 
adjacent 10 inierlophid valley. Anterior margin 
abutting posterior buccal margin of P3. Pro- 
lolophid markedly shorter than hypolophid with 
protoconid set on approximate midline. Lingual 
cuspids positioned closer to margin and with 
sleeper lateral walls than buccal cuspids. Both 
prololophid and hypolophid crests concave ante- 
riorly, posterior faces of both lophids verlical. 
Protoconid taller than meiaconid. Paracristid 
straight but anterolingually inclined as descends 
from protoconid apex lo a prominence positioned 
just posterior to anterior margin. A short but 
clearly defined ridge is directed lingually from 
apex of prominence, ending abruptly anterior to 
midpomt of prololophid. While the prominence 
may represent the paraconid, an alternative view 
preferred here is that this ridge represents a lin- 
gual extension of the paracristid, and its termina- 
tion the paraconid (sec Fig. 7c). Line of 
paracristid continued by a ridge running anteri- 
orly from anterior prominence for very short dis- 
tance and at steeper angle to anterior margin. 
From here a continuous ridge descends 
ventrobuccally on antenor margin, forming bor- 
der of narrow precingulid which ends before 
reaching buccal margin. No premetacrislid or 
postmetacristid. An enamel ridge, the *pro- 
tostylid crest' descends posterobuccally from 
apex of protoconid for about half height of thai 
cusp where it is contacted by ascending, anterior 



portion of cri.stid obliqua. Cristid obliqua de- 
scends anterolingually from apex of hypoconid 
before turning anteriorly to cross wide inier- 
lophid valley and ascend diagonally on posterior 
of protoconid. 

Entoconid taller than hypoconid which shows 
evidence of wear: enamel breached on lingual 
side of apex. Preentocristid descends directly an- 
teriorly from apex of entoconid to floor of inier- 
lophid valley. Posthypocristid short, extremely 
lingually displaced, originating at lowest point of 
hypolophid crest, closer lo entoconid than 
hypoconid, descending ventrolingually to meet 
postentocristid just above base of entoconid. In- 
verted triangular fossette enclosed laterally by 
posthypocristid and postentocristid. Below junc- 
tion of postentocristid and posthypocristid is 
short, prominent enamel ridge, descending at dif- 
ferent angle to postentocristid and which may 
represent a reduced hypoconulid. This ridge 
forms lingual margin of broad, horizontal 
hypocingulid extending across two-thirds of base 
of hypolophid. 

Broad wear facets on posterior face of pro- 
lolophid extend across link between protostylid 
crest and crisiid obliqua; similar facets on poste- 
rior face of hypolophid extend across 
posthypocristid. Facets bear fine vertical striae. 

M2 rectangular in occlusal outline but with 
slight concavity of lingual margin adjacent to end 
of inierlophid valley. Prololophid and 
hypolophid subequal in length, occlusal crest of 
hypolophid slightly more anteriorly concave than 
that of prololophid. Lingual cuspids set closer to 
lateral margin and with steeper lateral walls than 
buccal cuspids. Metaconid taller than protoconid 
which shows evidence of wear: enamel breached 
lingually adjacent lo apex. Paracristid directed 
anterolingually as descends from protoconid 
apex to anterior margin. Prominenl prc- 
metacristid runs slightly buccally as descends 
from metaconid apex lo anterior margin. Short, 
broad anterior cingulid enclosed between these 
cristids. Short precingulid buccal lo paracristid. 
In posterior view occlusal crest of prololophid 
forms shallow *V' with low point located closer 
to protoconid than metaconid. Cristid obliqua 
forms thick enamel ridge as crosses inierlophid 
valley, tapering somewhat anteriorly as ascends 
short distance on lingual side of posterior face of 
protoconid. No postmetacristid or preentocristid: 
broad inierlophid valley widely open lingually. 
Floor of inierlophid valley considerably more 
elevated on lingual side of cristid obliqua than on 
buccal side which slopes steeply towards crown 



274 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 3. QMFl 9820, Holotype of Wururoo dayamayi gen. et sp. nov. A, buccal view. B, lingual view. C, stereopair 
of occlusal view. Scales = 10mm. AR number is an informal system used in the Vertebrate Palaeontology 
Laboratory, University of New South Wales. 



base. Entoconid and hypoconid subequal in 
height. No posthypocristid. Thick ridge of 
postentocristid continuous with similarly promi- 
nent hypocingulid which descends slightly ven- 
trally as crosses posterior base of the hypolophid. 

M3 with longer anterior cinguiid, lingual cus- 
pids taller than buccal cuspids, protolophid 
slightly longer than hypolophid and thickened 
ridge, representing remnant of posthypocristid, 
descending vertically from low point of occlusal 
crest on posterior face of hypolophid. Less worn 
than is M2. Cristid obliqua can be seen to arise 
from more anterolingual position relative to apex 
of the hypoconid. 

M4 smaller than M3 with much narrower ante- 
rior cinguiid. Lacking precingulid buccal to 
paracristid. Hypolophid markedly shorter than 
protolophid. Metaconid taller than protoconid, 
crest of protolophid formed chiefly by buccal 



crest from apex of metaconid, descending to meet 
lingual flank of protoconid at a point directly in 
line with anterior end of cristid obliqua. Cristid 
obliqua originates lingual to hypoconid apex and 
runs directly anteriorly across interlophid valley. 
Hypoconid and entoconid subequal in height but 
entoconid set somewhat anterior to hypoconid. 
Apex of entoconid damaged during life and bro- 
ken edges of enamel subsequently smoothed as a 
result of wear. Hypolophid crest with a narrow 
V-shape in posterior view with lingual arm run- 
ning slightly anteriorly towards damaged apex of 
entoconid. No preentocristid but very thick 
postentocristid running ventrobuccally on poste- 
rior face of entoconid, merging with equally 
prominenlhypocingulid which crosses half width 
of hypolophid base. 



TWO NEW BALBARINE KANGAROOS 



275 



TABLE 2. Dental parameters for QMF 19820, Holotype of Wururoo dayamayi gen. et sp. nov. 



Cat. No. 


P3 


Ml 


Mt 


M:i 


Ma II 


1 


mw 


h 


! 


pw 


hw 


I 


pw 


hw 


1 


pw 


hw 


1 


pw 


hw 1 


QMF19820 


8.6 


4.9 


67 1 


5.9 


4.1 


4.2 


6.5 


4.8 


4.5 


6.9 


5.1 


4.7 


6.7 


4.3 


4.1 1 



DISCUSSION. P3 is much larger and more mas- 
sive than that in Nambaroo or Ganawamaya 
Cooke, 1 992, and is similar to that in undescribed 
Riversleigh species of Balbaroo. The large P3 in 
the latter balbarines is more similar in profile and 
size relative to molars to propleopines and 
hypsiprimnodonlincs than to macropodids. The 
diastema in W. dayamayi is also much shorter 
than in Nambaroo or Ganawamaya. The shorter 
diastema and markedly more robust P3 may be 
indicative of a greater reliance on the use of 
premolar shearing action in food collection 
and/or processing. The similarity of 
plagiaulacoid premolars in more derived 
balbarines, propleopines and hypsiprimno- 
dontines may be the result of convergence in 
species placing a similar emphasis on premolar 
shearing. However, premolars in these groups are 
similar in form to those of phalangerids and it is 
likely that the palgiulacoid premolar form is 
plesiomorphic for macropodoids. If robust, 
plagiaulacoid premolars are plesiomorphic, 
within Balbarinae the markedly more gracile pre- 
molars of Nambaroo and Gamiwamaya would 
represent an apomorphy for a clade containing 
these two genera. 

The lingually displaced Mi poslhypocrislid 
seen in W, dayamayi is also present in species of 
Nambaroo and in Ganawamaya aedicidis Cooke, 
1992. lis widespread occurrence among 
plesiomorphic balbarines supports the view of 
Flannery & Rich (1986), that lingual displace- 
ment of the posthypocristid has played an import- 
ant role in hypolophid formation in Balbarinae. 
The connection of the cristid obliqua to the pro- 




FIG. 4. Dimensions of QIV1FI9820, holotype of 
Wuntroo dayamayi gen. et sp. nov., buccal view. 



loslylid crest resembles the connection between 
the cristid obliqua and the discrete proiostylid of 
Nambaroo saltavHS as noted by Flannery & Rich 
(1986), and suggested by them to represent the 
plesiomorphic slate of this character in 
macropodids. 

The Ml precingulid in other undescribed 
plesiomorphic balbarine species (pers. obs.) re- 
ceives the poslerolingual cuspule of P-^. The pre- 
cingulid is low in W. dayamayi, suggesting that a 
prominent poslerolingual cuspid is to be expected 
on its P\ The lingual ridge associated with the Mi 
paraconid forms the margin of a small anterior 
cingulid which receives the posterior end of the 
P^ occlusal margin, as indicated by signs of wear 
on the posterior face of the ridge. The broad wear 
facets on the posterior faces of the Mi lophids 
indicate broad contact with the lophs of M' while 
the orientation of the striae indicate greater verti- 
cal rather than lateral relative movement between 
lophs and lophids. 

CUSPID HOMOLOGY AND EVOLUTION 

OF LOWER MOLAR MORPHOLOGY IN 

BALBARINAE 

Differing views of cuspid homology and evo- 
lution of lower molar morphology among 
macropodoids are examined and alternative 
hypotheses proposed to determine homology of 
structures on lower molars of plesiomorphic 
balbarines from Riversleigh and to elucidate the 
course of molar evolution within Balbarinae. 
Since interpretations of cuspid homology in 
Hypsiprymnodon have been central to wider ar- 
guments pertaining to macropodoid molar evolu- 
tion, such interpretations of previous authors are 
reviewed first and a new interpretation presented 
which incorporates evidence provided by the new 
species described herein. 

The posierobuccal proiostylid crest in VV. 
dayamayi is very similar to thai which descends 
from the dominant trigonid cuspid to a tiny buccal 
cuspid in dP? of Hypsiprimnodon moschatus. 
Ride ( 1 96 1 ) identified the dominant cuspid as the 
metaconid and the liny cuspid as the protoconid, 
but has since modified this interpretation (Ride, 
1993), considering the central trigonid cuspid 
posterior to the paraconid on dP3 10 be a 



276 



MEMOIRS OF THE QUEENSLAND MUSEUM 



more posterior molars. Molars 
posterior to Mi haveabuccaily 
positioned protoconid which 
has a paracrislid running to the 
anterior margin. The 
metaconid of these molars is 
linguaily positioned and has an 
associated premetacristid. 
Both cristids arc present on Mi 
and bear the same relationship 
to each other as they do on 
more posterior molars. The 
metaconid is distinct but the 
protoconid, from which the 
paracristid originates, may be 
considerably reduced and is 
more centrally positioned, the 
FIG. 5. Posterior occlusal views of Mi A, QMF303401, holotype of trigonid thereby becoming lal- 
Nambaroocoupensp.nQ\.B,QMF\9^26M\oiypcofWitruroodayamayi erally compressed in a manner 




VX- 




gen. et sp. nov.. 

parametaconid and the small cuspid posterobuc- 
cal to this to be the protoconid, with the normal 
relationship of protoconid, metaconid and 
paraconid preserved on Mi, the parametaconid 
having been lost in this tooth. 

Archer (1978) raised the possibility that the 
anterobuccal cusp in Mi (his M2) of//, moschatus 
(and other macropodoids) may be Ihe homo- 
logue of the phascolarctid protostylid and not the 
protoconid' and further that 'the tiny cusp ob- 
served by Ride (1961) on Mi ( Ride's dP4 and 
here dPs) of Hypsiphmnodon may be the serial 
homologue of this protostylid'. Archer & Flann- 
ery (1985) and Flannery & Rich (1986) also 
identified the postcrobuccal cuspid of the Mi 
trigonid of//, moschatus as the protostylid having 
here an anterior cristid descending to the anterior 
margin, and the cuspid lingual to this, also with 
an anterior cristid, as the protoconid, the 
metaconid being displaced posterolingually. 

The interpretation of trigonid cuspid homology 
on dP3 offered by Archer (1978) is accepted here 
but none of the above views regarding Mi 
trigonid morphology in //. moschatus are upheld. 
For reasons explained below, the most buccal 
cuspid-like structure associated with the pro- 
tolophid is accepted as the protostylid but the 
lingual cuspid of the protolophid is regarded as 
the metaconid. 

As Ride (1993) noted, constraints imposed by 
functional interactions with premolars can alter 
the topography of teeth at the premolar/molar 
boundary, so cuspid homology of Mi in //. 
moschatus has been examined with reference to 



closely resemblmg that seen m 
balbarines. In unworn //. 
moschatus specimens (QMJ10233, QMJ9327, 
QMJ145), the paracristid originates from a slight 
elevation, interpreted here as the reduced pro- 
toconid (Fig. 6F). A posterobucccal crest links 
this to a larger cuspid which is thus the protostylid 
in the sense in which the term is used by Van 
Valen (1966), and Butler (1978): it occurs in the 
same position as the protostylid of other diprotod- 
ont marsupials, e.g., pseudocheirids and 
phascolarclids and Nambaroo, a position in 
which it fulfils the function ascribed to it by 
Butler (1978), i. e., 'to shear against the an- 
terolingual surface of the paracone of the corre- 
sponding upper tooth' (a function which could 
not be fulfilled by the 'one or more protostylids' 
indicated by Ride (1993), as occurring on the 
precingulid anterobuccal to the Mi trigonid in the 
propleopines, Jackmahoneya and Propleopus). 

Ride (1993) indicated 3 cuspids immediately 
posterior to the paraconid of dP3 of//, moschatus: 
the large, central cuspid identified as the 
parametaconid, a posterolingual metaconid and a 
very small postcrobuccal protoconid. The 
posterolingual cuspid is accepted here as the 
metaconid but since the large, central cuspid has 
a cristid running anteriorly from its apex toward 
the paraconid, the cuspid is here regarded (as it 
was by Archer, 1 978) as the protoconid, its cristid 
as the paracristid and the small postcrobuccal 
cuspid as the protostylid, homologous with that 
which also occurs on dP? in undcscribed 
Riversleigh specimens of Nambaroo (pers. obs.), 
the protostylid on Mi of //. moschatus and the 



TWO MEW BALBaRIN^ ICANGAROOS 



277 



dP/3 






M/1 






O Paraconid 

O Protoconid 

^ Protostylid 

a Parametaconid 

D Metaconid 



FIG. 6. Interpretations of irigonid cuspid homology in Hypsipn'mnodan maschatus. A & B. Ride (1993). C, 
Archer (1978). D. Archer & Flannery (1985). E. F. Cooke (herein). 



proiostylid crest on M i of Wururoo and on dF3 o\' 
bulungamayine species (Cooke, 1997). 

The homology of the buccal cuspid on the Mi 
proiolophid of propleopines is less clear. Ride 
(1993) labelled this cuspid as the protoconid and 
ihc cuspid immediately lingual to it as a 
ncomorph. the pafametaconid. Flannery & 
Archer ( 1985) labelled the buccal cuspid as the 
proiostylid and the cuspid immediately lingual to 
il as the protoconid. While it is clear that the 
prolo.slylid, as a discrete cuspid or as a protostylid 
crest, is common among plesiomorphic 
macn>podoids, it is unusual in this group for it to 
have acquired a cristid mimicking iheparacrislid 
in linking to the anterior cingulid. If Flannery &. 
Archer are con'ect, this would constitute a syn- 
apornorphy for propleopines as would the 
neomoiphic parametaconid in Ihe interpretation 
of Ride. 

The suggestion of Archer & Flannery (1983) 
that Ihe buccal cuspid of the protolophid in Mi of 
propleopines and potoroines was the protostylid 
and that the metaconid wa.s losi in potoroines but 
retained in propleopines was enlarged upon by 
Flannery & Rich (1986), They suggested that in 
poloroids the protostylid formed the buccal pro- 
tolophid cuspid and the protoconid the lingual 
cuspid, in contradistinction to macropi>dids in 
which the protostylid is lost and the buccal cuspid 
is the protoconid and the lingual cuspid the 
metaconid (lost in potoroids other than pro- 
pleopines). The plesiomorphic balbarincs, 
Namharoo, Wttruroa and ikmawanuiya certainly 



indicate that the protostylid has been reduced and 
ultimately lost in balbarines, however there is less 
evidence to support a belief that it has been re- 
tained in potoroids. The Mi (Cooke, 1997) pro- 
tostylid of Palaeopotorous priscus Flannery <& 
Rich. 1986. bears a similar relalionship to the 
protoconid and cristid obliqua as docs that of W. 
tnoschatus, exhibiting a condition intermediate 
between the distinct cuspid of the protostylid of 
Nambaroa and the protostylid crest of W. 
dayamayi. It thus may represent an early stage in 
Ihe reduction of the protostylid among potoroids. 
In the proposals of cusp homology in 
macropt:)dids and potoroids referred to above, the 
condition occurring in H, nwschatus is crucial 
becati-se il is used in both cases to represent the 
iniermediale condition between a phalangcrid an- 
cestor and more derived potoroids. If, as has been 
proposed here, the reduced cuspid from which the 
paracrislid originates on the short Mi pnmjiophid 
of /y, maschatus is the protoconid, then there is 
no reason to suppose that this is not also the case 
in other poloroids and the sequence, Pal- 
aeopoioroHS - Hypsiprimnodon - Potorous, used 
by Flannery & Rich (1986) to illustrate their 
argument can be taken as indicating a transition 
involving reduction of the protostylid. 

The protostylid is also exhibited in varying 
degrees of development within phalangcrids. A 
very siinilar structure to ihe Mi protostylid crest 
of W. dayamayi occurs in Phalange r inter- 
castellanus and joins the crjstid obliqua. In 
Tru'hosurm vulpecida there ts a straight ridge 



m 



MEMOIRS OF Tl IE QUEENSLAND MUSEUM 



pec 







0hc 



.-Ac picsiomorphic For (he group. In 

pseudocheirids among Petauroidea and 
Nambarao among Macropodoidea the 
prolostylid is fully developed but in most 
phal anger ids and in Palaeopotnnnts. 
Hypsipritmwdon and Wururoo the poien- 
tial is not as fully expressed. This expres- 
sion has independently been suppressed 
among more derived macropodoids (w iih 
ihc possible exception of propleopines) 
and t>ihcr phalangeridans. 

Wuno'oo is more derived than 
Nambarao in terms of Mi morphology: 
the protostyiid has been reduced, a 
neomorphicanicnorcingulid has begun to 
form and the Irigonid basm has become 
partly enclosed. Namhawo and \V. 
dayatnayi jndicaic initial stages of a trend 
among balbarines towards the dcvelop- 
meni of an anterior cingulid and the enclo- 
sure of the Irigonid basin. ConcomUani 
with thus trend is a decrease in the relative 
importance oflheprecingulid which *KCU- 
pies much o( the Mi anterior margin in 
plesiomurphic species such as N. coupert. 
Hypolophid formation in balbarines 
proceeded as ouilmcd by Klanncry & Rich 
^. ^ (1986). involving elevation and lingual 

displacementoftheposihypocrisiid which 
FIG. 7. Evolution of lower molar morphology in balbarines contributes a buecal component to the 
illustrated by RM). A, hypothetical macropodoid ancestor, B, hypolophid, the lingua! component con- 
Namhamo cottperi. C. WUiruroa daxumai. D, Batbawo grade, iribuled by a buccally directed crisi id from 
Abbreviations: Ac=amenor cingulid; co=cnsiid obliqua, ihe enloeonid. The latter crislid is not 
Ec=cnioconid; Hc=hypocingulid; Hd=hypoconulid; necessarily a neomorphic eniohvpo- 
Hy=hypocomd; Mc=meiacon,d; Pa=paracon.dj Pc=pre- ■ ^^ ^^-^ ,993 However; the 
cmgulld; pec=poslentocnsiid, phc=posihypocristad: Pr=pR> . v 1 .u xx i- k\ i 

toconid;Pr.Votostylid;prc=protosty|idcr^t hypoconuhd on the M, ot .V. voupen and 

'^ Its apparent presence in a reduced lorm on 

Mi of W- ilayamayi indicates the addi- 
tional involvement of this cu.spid in hypolophid 
formation in picsiomorphic balbarines (Fig. 7). 
The hypoconulid m such tonus is contacted by a 
lingually di.splaced. diagonal posthypocristid. 
An inverted triangular fossciie occurs on the lin- 
gual posterior face of the hypolophid. bounded by 
the posthypocristid and theposteniocristid iwhen 
present). A shallow fossettc develop.s buccal to 
the posthypocristid and a ncomorptiic cingulid, 
the hypocingulid according to the detlnition of 
Butler (1990), develops running transversely 
across the posterior base of the hypolophid from 
the hypoconulid or. when that structure has been 
lost, from the ventral end of the posthypocristid, 
as m N. saluivus. The posthypocristid is lost in 
more derived species such as tht>se o\' Balharop 
in which the postentocrisiid links to a transverse, 



descending posierobuccally to the base iy{ the 
protoconid where it links to the cristid obliqua. In 
T. cnnlnus there is a ridge on the protoconid 
resembling that in T. vulpecula, but it does not 
contact the cristid obliqua which independently 
ascends the posterior face of the protoconid. 
Flannery & Rich (1986) indicate a discrete pro- 
tostyiid in Phalangerveslitus and a similar situa- 
tion occurs in Spilocuscus nnicidatus, but in some 
specimens of this species the protostyiid may be 
reduced to a ridge on the poslerobuccal Hank of 
the protoconid. Flannery & Archer (1987) sug- 
gest a protostyiid on Ml in Sfrij^oruscus reidl and 
Thclwsurus dkksoni, both from System C sites 
atRiversIeigh. 

It would appear that the genetic potential for the 
formation o\' a protostyiid is widespread among 
phalangeridans and may be considered as 



TWO NEW BALBARINE KANGAROOS 



279 



posterior cingulid homologous with the 
hypocingulid. 

The modificalion offered above to the hypoth- 
esis of Flanncry & Rich has been extended by 
Ride (1993) to macropodoids in general but I 
apply it only to balbarines. The bilophodont 
lower molar morphology of bulungamayines is 
derived from a bunolophodont ancestral form by 
different means to those outlined above (Cooke, 
1997) and clearly indicates independent evolu- 
tion of lophodonty within this group. However, a 
hypoconulid positioned low on the posterior, lin- 
gual face of the hypolophid and associated with 
a posthypocristid, is the probable plesiomorphic 
condition for all macropodoids The basal 
macropodoid may well have had a bunolophod- 
ont lower molar morphology similar to that of 
Palaeopotorous or of phalangcrids, but with a 
more distinct protoslylid buccal to the protoconid 
and a hypoconulid contacted by the 
posthypocristid at the posterior base of the en- 
toconid. The lingual component of the balbarine 
hypolophid would represent a reduction of the 
buccal cristid of the enioconid which forms the 
transverse posterior lophid of bunolophodont 
macropodoids. 

The hypoconulid in plesiomorphic balbarines 
and its absence in any known potoroids of com- 
parable age suggests that balbarines diverged 
early from the main stem of macropodoid evolu- 
tion and independently and probably rapidly 
evolved lophodonty, the better to exploit a herbi- 
vore niche. The combination of plesiomorphic 
dental characters present in balbarines, including 
lateral compression of the Mi trigonid and the 
presence of both a hypoconulid and prolostylid 
on this tooth, contrasts markedly with the absence 
of such characters in bulungamayines. There has 
been parallel (and probably later) evolution of 
lophodonty within Bulungamainae and the ab- 
sence of plesiomorphic molar characters such as 
those indicated above, suggests that 
bulungamayines may be more likely to be ances- 
tral to the highly derived macropodids. 

ACKNOWLEDGEMENTS 

Research grants from the Australian Research 
Council and the University of New South Wales 
have been the primary mechanism for providing 
the research material examined in this study. Ad- 
ditional support for the Riversleigh project has 
come from the National Estates Program grants, 
the Australian Geographical Society, The Austra- 
lian Museum, The Riversleigh Society, ICl FHy 



Ltd, Century Zinc Limited, the Mt Isa Shire and 
private donors. 

I thank the Director and staff of the Queensland 
Museum for provision of workspace and facili- 
ties; Jeff Wright and Alex Cook for preparation 
of photographic prints; Michael Archer and 
David Ride for their helpful discussions; the staff 
and students at the University o'i New South 
Wales involved with the Riversleigh Project who 
have been so willing in their assistance. 

LITERATURE CITED 

APLIN. K.P. & ARCHER, M. 1987. Recent advances 
in marsupial systcmatics with a new syncretic 
classification. Pp xv-lxxii. In Archer. M. (ed). 
Possums and opossums: studies m evolution'. 
(Surrey Beaity & Sons: Sydney). 

ARCHER, M. 1978. The nature of the molar- premolar 
boundary in marsupials and a reintcrpretalion of 
the homology of marsupial cheekteeth. Memoirs 
of the Queensland Museum 18: 157-164. 

ARCHER, M. & FLANNERY, T.F. 1985. Revision of 
the extinct gigantic rat kangaroos (Potoroidac: 
Marsupialia). with a description of a new Miocene 
genus and species and a new Pleistocene species 
of Propleopus. Journal of Paleontology 59: 1131- 
1149. 

ARCHER, M.. GODTHELP. H., HAND. SJ. & 
MEGIRIAN, D. 1989. Fossil mammals of 
Riversleigh, northwestern Queensland: prelimi- 
nary overview of biostraligraphy, correlation and 
environmental change. Australian Zoologist 
25(2): 29-65. 

BREEN, G. 1981. The Mayi languages oi'the Queens- 
land gulf country. Australian Institute of Aborig- 
inal Studies AlAS new series 29. 

BUTLER, P.M. 1978. Molar cusp nomenclature and 
homology. Pp 439-453. In Butler. P.M. & Joyscy, 
K. (cds), Development, function and evolution of 
teeth'. (Academic Press: London). 
1990. Early trends in the evolution of tribosphenic 
molars. Biological Reviews 65: 529-552. 

COOKE, B.N. 1992. Primitive macropodids from 
Riversleigh, northwestern Queensland. Al- 
chcringa 16:201-217. 
1997. New Miocene bulungamayine kangaroos 
(Marsupialia, Potoroidae) from Riversleigh, 
northwestern Queensland. Memoirs of the 
Queensland Museum 4 1 : 28 1 -294. 

FLANNERY, T.F. 1989. Phylogeny of the 
Macropodoidea: a study in convergence. Pp 1-46. 
In Grigg, G. Jarman. P & Hume, 1. (eds), Kanga- 
roos, wallabies and rat-kangaroos. (Surrey Beatty 
& Sons: Sydney). 

FLANNERY. T. & ARCHER, M. 1987. Stri^oatsats 
reidi and Trichosunis dicksoni, two new Ibssil 
phalangerids (Marsupialia: Phalangcridac) from 
the Miocene of north western Queensland. Pp 527- 
536. In Archer, M. (ed.), Possums and opossums: 



280 



MEMOIRS OF THE QUEENSLAND MUSEUM 



studies in evolution. (Surrey Beatty & Sons: Syd- 
ney). 

PLANNER Y, T.F.. ARCHER, M. & PLANE. M. 1982. 
Middle Miocene kangaroos (Macropudoidea: 
Marsupialia) from three localities in northern Aus- 
tralia, with a description of two new subfamilies. 
Bureau of Mineral Resources Journal of Austra- 
lian Geology and Geophysics 7; 287-302. 

FLANNERY, T.F. & R'ICH, T.H.V. 1986. 
Macropodoids from the Middle Miocene Namba 
Formation, South Australia, and the homology of 
some dental structures in kangaroos. Journal of 
Paleontology 6()(2): 418-447. 

FLOWER, W.H. 1867. On the development and suc- 
cession of teeth in the Marsupialia. Philosophical 
Transactions of the Royal Society of London 
8157:631-641, 

LUCKETT. W.P. 1993. An ontogenetic assessment of 
dental homologies in therian mammals. Pp 182- 
204. In Szalay, F.S, Novacck, M.J. & McKcnna, 
M.C. (eds). Mammal phylogeny. (Springer- Ver- 
laa: New York). 

MYERS, T.J. & ARCHER, M. 1997. Kuwrmtjangmm 
(Marsupialia. Ilariidae): a revised systematic 
analysis based on material from ihe late Oligocene 
of Riversleigh. northwestern Queensland, Aus- 
tralia. Memoirs of the Queensland Museum 41: 
379-392. 



RIDE, W.D.L. 1961. The cheek teeth of 
Hypsiphmnodon moschatus Ramsay 1 876 
(Macropodidae: Marsupialia). Journal and Pro- 
ceedings of the Roval Society of Western Aus- 
tralia 44: 53-60. 

1993. Jackmahoneya gen. nov. and the genesis of 
the macropodiform mohir. Memoirs of the Asso- 
ciation of Australasian Palaeontologists 15:441- 
459. 

SZALAY, F.S. 1969. Mixodeclidae, Microsyopidae, 
and the Insecliyorc-Primaie transition. Bulletin of 
the American Museum of Natural History 140: 
193-330. 

VAN VALEN, L. 1966. Deltatheridia, a new order of 
mammals. Bulelin of the American Museum of 

Natural History 132: 1-126. 

1 994. Serial homology: the crests and cusps of mam- 
malian teeth. Acta Palaeonlologica Polonica 38, 

3/4: 145-158. 

WOODBURNE, M.O.. MACFADDEN, B.J.. CASE, 
J.A.. SPRINGER. M.S., PLEDGE, N., POWER. 
J.D.. WOODBURNE, J.M. & SPRINGER. K.B. 
1993. Land mammal biostraligraphy and mag- 
neiostratigraphy of the Etadunna Formation (late 
Oligocene) of South Australia. Journal of Verte- 
brate Paleontology 14: 483-515. 



NEW MIOCENE BULUNGAMAYINE KANGAROOS (MARSUPIALIA: 
POTOROIDAEjFROM RIVERSLEIGH, NORTHWESTERN QUEENSLAND 

B.N.COOKE 

Citokc. BN. 19970630: New Miocene bulungamayine kangaroos (Marsupial in: PotoroicJac) 
from Rivcrsleigh. northwestern QuccnsIandT Memoirs ofihe Queensland Museum 41{2): 
281-294. Brisbane. ISSN 0079-8835. 

Nowiiigee matnx gen. et sp. nov. and Ganguroo bikimina gen. ct »p, nov. arc described from 
frcshwalcr Miocene System B limestone al Riversleigh^ NW Oucensiand. Subfainilial 
diagnosis ofBulungamayinae is emended. The new species indicate ihai lophodonly was 
achic\cd in buiiingamayineis by ii diffcrcnl prwess from thai in balbannes. Similarities in 
denial morphology between biiUmgamuyines and laie Miocene maeropodids siiggcsi Ihtii 
Biilun^amayinac is anecslral to Macfopodidac. □ Rivvrshigh. kan^nwo, halharinvs. 
Bulungamayinae, hphndonty 

B.N. Cook^, School of Lift Science, Queenskind University of Technology, GPOBi>x24M, 
Brisbane, Queensland 4001, Australia; 17 Decemher I9Q6. 



Ral-kangaroos or potoroids, in the sense of 
Archer & Banholomai (1978) and Barlholomai 
{ l*>78). were unknown in the pre-Pliocene fossil 
rc-jord of Ausinilia until Archer (1 979) desL-ribcd 
Wabularoo tum^htotii as an enigmatic, lophodont 
kangaroo from ihc Riverslcigh Local Fauna of the 
Carl Creek Limestone. Flanncry el al, (I9H2) 
described Bulun^^amaya delicaro from the Carl 
Creek Limestone and placed it and W. nausihtoni 
in the potoroid Bulungamayinae. Cumardec 
pitscuali, also from ihc Carl Creek Limestone Wiis 
described in the same paper hut placed in the 
Fotoroinac. More recent additions to ihe record 
of poloroines include Wakiewakie lawsttni 
Woodbume. 1984 and Purtia mosaicus Case, 
1 9K4. from the Ngapakaldi Local Fauna of South 
Australia and Hettongia tnoyesi Flanncry & 
Archer. 1987, from Two Trees Site at 
Riverslcigh. Flanncry & Rich (I9H6) described 
Gmnardee and indcicrminalc potoruines from 
IhcTarkarooloo Local Fauna ol South Australia. 

Archer & Flanncry (1985) erected Pto- 
plcopinae for Ekaltadeta imu. a giant rat kanga- 
rt)o from Gag Site at Riverslcigh and Pleistocene 
and Pliocene .species of Frop/eopus. Flanncry &. 
Archer (1987) described Hvpsiprimnadon 
hanholoniaii from the Gag Site al Riversieigh 
and Flanncry & Rich ( 1986) reported 
hypsiprimnodontine material from the Tar- 
karooioo Local Fauna. Palacopoioroinae Flann- 
ery & Rich, 1 986 accommodates Palaeopotorous 
pnsciis from the Tarkarooloo Local Fauna. 

The diversity of pre-Pliocene potoroids is such 
that only 2 o\' the more recently discovered .spe- 
cies have been assigned to existing genera and 3 
new subfantilies have been proposed. Of these 



Bulungamayinae Flant>ery ci al, 1Q82. has ai- 
tracied most discussion. Woodburne (1984) and 
Case (1984) argued thai ihc lophodont 
bulungamayincs W. nau^'hroni and H. dclicam 
share characteristics with plcsiomorphic 
macroptidids (Iheir macropodines) such as 
Dorcopsoides fossHis Woodburne. 1 967. 
Dorcupsis and Dorcopsidus and should be in- 
cluded in Macropodidae (their Macropodinac). 
They also argued a similar placement of 
Gutnardee Flanncry et aL ( 1984) identified syn- 
apomorphics which ihey considered united 
potoroids in a monophyielic group and defended 
their placement of bulungamayincs and 6'. 
pascualiwihxn Potoroidae on the basis of several 
of those derived slates. Flanncry & Archer 
( 1 9S7a, b) demonstrated that one suggested syn- 
apomorphic chaiacicr,squamosaM*rontal contact 
on Ihc lateral wall of the cranium, is not mnvcrsal 
within the group and could no longer be consid- 
ered as a potoroid synapomorphy. The slate of 
this character is unknown in new bulungumayinc 
malei iai desci ilxjd l)clow. 

The new species are similar in si/.c and have 
similar premolar morphology to thai of B. 
delicata and together with thai species represent 
a sequence which reveals much nhoui ihc evolu- 
tion of lophodonty within Bulungamayinae 

METHODS 



Molar homology follows Luckeit ( 1993), prcs 
molar homology tbllows Flower (1867). Dental 
descriptive lerminolugy is principally that used 
by Archer (1984) but with some terms adopted 
from S/alay (1969) and Ride (1993), In upper 



283. 



MEMOIRS OP Tl in QUIICNSLAND MUSEUM 



iDoliirs tht' term, 'paracmgulum' is used to indi- 
cate an anterior cingulum hounded laterally by 
the prcpanicrislii and prepruUxTisla as indiciilcd 
in Szalay (1969). Ride used 'prccingulum' for 
this structure. 1 use 'procingulunV foi an nnieriof 
cingulum extending lingually irom the prc- 
protocrista, following S/alay. This structure was 
referred to hy F^idc as the \uiicrulingu:d 
cinguium'. 'Metacinguium' is used lor a pusle- 
rior cnigulum buundtid posk*rulii\gually by the 
poslmciaconulc crivSia; 'paracrisLa' and meia- 
trisla' arc used fur the lingually directed, loph- 
ibrming crislac from the paraconc and mctacone. 
respectively. Use of the lalU'-ricTO) in ihisiiiannet 
is a departure from S/.alay who uses *metacrista' 
synonymously with pUMimelacrisia' . 
Tostmetacrista' is used here in the sense of 
Archer (1984). 

Cusp homoldgy (if upper molars is thai of 
Tedford & WcKidburne ( 19S7). with the posterior 
buccal and lingual cusps designated as meiaconc 
and mclaconulc respectively, and the cuspulc be- 
tween these as the neoincia'conule. Siipragencrie 
classilicatfon fullows Aplin Si Archer (1987). 
Material is housed in the Queensland Museum 
(QMF). Measurements arc in millimcircs. 

SYSTEMATICS 

Supcrfamily MACROPODOIDEA Grav, lS2l 

family POTOROIDAH Gray. IS21 

Subfamily BULUNGAMAYINAE 

Flannery. Archer & Plane 1982. cmend- 

Cuoke. 1997 

Bulungamaymcs have a buccally expanded mas- 
seteric canal eonfluenl over its length with the 
infenor denial canal, the common canal penetrat- 
ing deeply within the denlaty below the molar 
row. The digastric process of the deniary is ex- 
panded so that the ventral margin of the dcntary 
is convex below the molar row. Ii has enamel 
confined lo the buccal surface but extensive on 
Ihat surface and not confined to the ventral por- 
tion as it is in putofDmcs. Vciiiral imd dorsal 
enamel llangcs are present on li. F^ is elongate 
with many fine iranscristids and a bulbous base. 
A small ifiodi, I2 but which may he a small canine, 
is just posterior to the dorsal margm of the li 
alveolus. Molar teeth may be bimolophodttnl or 
lophodonl as defined by Flannery ci a). ( 1984). 

Bulungamayincs differ from hypsiprimno- 
dontines and propleopines by having an elongate 
Pi who.sc occlusal margin in lator;d view is 
straight or concave, rather than a plagiaulacoid P.^ 



with a convex occlusal margin. They djflcr from 
poioroines hy having much more bulbous premo- 
lars, by having iin Ij and a much more extensive 
area of enamel on the buccal surface of Ij. They 
differ from paUieopoToroines by lacking a distinct 
proiosiylid on Mi. 

RLMARKS. Type speciiijcns ol bulungamayines 

erected herein arc far more complete than those 
of previously described species and reveal details 
of anatomical and dental features absent in the 
holotypes of Bulnngamaya delicata and 
Wahtilaroo mmghtoni . This additional informa- 
tion forces the above subfamily revision. 

Nowidgetf gen, nov. 

TYPE SPECIES. Nowidgce matrix sp. nov. 

DIAGNOSIS. Bulungiimayjne with bunolophodoni 
molars. Upper molars with Jilnrge sly I;ircuspC extend- 
ing posici lOfly to close the buccal end cif ihe uucrloph 
region. 

ETYMOLOGY. Wminyi (.is spoKen by Ivy Slinken, 
formerly of Rivcisleigh Station) Nowid^ift, grand- 
mother. 

Nowidgee matrix sp. nov. 
(Figs I. 2; Table I) 

DIAGNOSIS. As for the genus. 

MATERIAL EXAMINED. Hototypc QMr303y0, 
from Camel Sputum, Godthelp's Hill. D-Siie Plateau. 
Paraivpcs QMF19961, 20255, 2276 L 30393, 30394. 
3039.5 from Camel Sputum Site. QMri9937, 20069, 
2(K)80. 30391 fram Wayne's Wok SUe, flat's Hilt, 
D-Sitc Plateau. Both System B sites (Archer el aL, 
1989) of Miocene age. 

ETYMOLOGY. Laiui mairix. mother of an animal, 
a'fers iu iis arice.stral pcsition. 

DESCRIPTION OF HOLOTYPE. Right deniary 
fragment of most of the horizontal ramus to the 
level of M4 and part of the ascending ramus. It, 
P ^ and M 1 .4 preserved. Aseending ramus at ai^out 
I \i)''' to occlusal plane of molar row. Masseteric 
canal eonlluenl with interior dental canal, mak- 
ing it difficult to assess extent of forward pene- 
tration of the masseler, but an tenor wall of 
masseteric fossa e.xlending anteriorly ro about 
level of the M3 protolophid . At this level diame- 
ter of common canal not greatly exceeding that 
o( sulcus representing inferior dental canal in 
posterior, lingual wall of masseteric fossa. This 
suggests anterior pt>rtit)n of common canal occu- 



NEW BlJLlfNGAMAYlNE KANGAROOS 



2^i 



pied chiefly by vessels assuciaicd with denial 
caiuiJ and masscicr noi passing much more ante- 
riorly than this level. Miisseleric lussa buccally 
expanded with Hat surface tor attachment ofsu- 
pcrlicial layci of mas.setei al anlerovcniral bor- 
der, extending dorsally on antcrohuccal niarein 
ofascendii^g ranms. Ventral margin ol horizontal 
nimus gently convex with lowest point hclow 
Mz/'M^, Merual foramen just anterior to P3. be- 
tween root of 1 1 and dorsal margin of the dia- 
stcina; mueh smaller po.slerit»r n^iental foramen 
ventral to protolophid of M2. Posicrovcmrally 
inclined buccinator sulcus between P3 alveolar 
margin and posterior mental foraivien. Diaiitema 
short, as long as P}. Damage to 1 1 alvcolax ma/gin 
iihscuring I: or its alvt^nliis (I2 alve^>lus in CJMF 
19*^37} Mandibular symphysis extending p*.>sie- 
riwly almost to level of Pi posterior margin. 

Dentition. Molar row .straight in occlusal view. F? 
llexed slighlly buccally out of alignmcni with it. 
In lateral view rnolat row concave: occlusal sur- 
f»L'cs ol M: and Mi lyii^g below line joining 
occlusal surfaces of Mi and M4. liffeclii of weiu- 
on molar teeth progressively less obvious to- 
wards posterior of tnolar row Molar si^e in- 
creases iViJin M1-M3 bul M3 larger than M4. 

li broken at anterior end, depth unift>rm over 
preserved length, rising ai approximately 2(7' rcl- 
^live to dorsal margin of horizontal ramus. 
Enamel confined to buccal side, has both dorsal 
<md ventral enamel flanges, veniral being more 
sUungly developed. Dorsal Jlange forms occlusal 
niargin. Circular cmsS'Seclion close to alveolus 
becoming more elliptical anteriorly. 

P^ blade-like. 5Wt longer than M|. Occlus^d 
outline sciiiilunar vviih .straight lingual margin 
and convex buccal margin. Occlusal crest slightly 
lingual to midline. Ilexes lingually at posterior 
end. Six small cuspids on occlusal margin ante- 
rior to longer, posterior cuspid. Transcnslids as- 
sociated with each o\'(^ minor cuspids and anterior 
and posterior margins of blade delineated by ver- 
tical cristids. Lingual surface of occlusal blade 
more steeply inclined than buccal. 

Mi almost square in occlusal outline but tmr- 
fowcr anteriorly than posteriorly. Protolophid 
shorter than hypolophid: protoconid closer to 
midline than is hypoconid Lingual cuspids taller, 
more sharply angular and closer to adjacent lat- 
eral margin than rounded, buccal counterparts 
which arc also more worn. Lingual surfaces ver- 
tical, buccal surfaces more gently sloping, Pro- 
tolophid formed by metacristid descending 
buccally from metacunid apex 10 lingual flank oi 
protoconid. Thick parucristid running aniero- 



lingually from apex of protoconid lo anlenuc 
margin Short, broad anterior cingulid anterior to 
anterior face of protolophid, hounded buccally by 
panicristid. lingually by anteriorly directed piv- 
metacristid. Broad prccingul id .sloping steeply 
vefltrally buccal lo paracristid. Sharply-dclli>cci 
poslmciacri.siid curving buccally from meiaconid 
ajx;x, desoending to narrow inierlophid valley. 
Anicnorly oriented precntocristid separated fri>m 
ventral end of post metacristid by nmrow cleft. 
Crislid obliqua very thick, descending an- 
leiolingually front apex of hypoconid lo inier- 
lophid valley, then inclining buccally n> apex of 
protoconid. Paracristid and cristid oblitiua form 
coininuous longitudinal ndge extending irom an- 
terior margin to hypoconid. Hypiilophid fomicd 
by buccal crest from the entoconid descending 
from entoconid apex and running buccally to 
meet lingual flank of hypoconid. Posthypociisiid 
descending lingually from hypoconid apex, 
crossing lingually posterior to buccal crest from 
entoconid to posterior of entoconid below apex. 

M^ larger and .squarer than Mj with prolt)lophid 
and hypolophid of about equal length and pro- 
toconid and liypdconid in alignment. Similar (o 
Mt but anterior cingulid longer and broader, prc- 
cingulid shorter and interlophid valley broader. 

Mji worn in trigonid region but lalonid rela- 
tively unworn . Crown very similar lo M: but niosi 
structures more clearly dellned. Cristid obliqua 
massive in interlophid region, not much lower 
than apices of buccal cuspids. Lwigual side of 
interlophid valley more open with greater sepoTii- 
lion of ventral ends of postmciacristid and pre 
cntocrisiid. Posthypocrislid sharply dellned 
crossing posterior suriacc from hypoconid apex 
to short, almost vertical (losicntocristid descend- 
ing Irom apc\ of entoconid. Deep, narrow trench 
between crest of postliypocrislid and buccal cresi 
from entoconid anterior lo il. 

M.4 unworn. Hyp(»lophid shorlcr than pro- 
tolophid. Cristid (thiiqua originating on aii- 
lerobuccal face ol hypoconid. beltnv ape>;. 
Preentocnstid and postmeiacrislid separated only 
by narrow cicll in interlophid vullcy. 
Posthypocrislid crest r»)unded, meeting cr^- 
toconid much closer to its apex than on anterior 
molars. Buccal crest from entoconid shorter atiii 
less sharply defined. 

DESCRIPTION OF PARATYPES. 

DENTARY FRAGMENTS. Horizontal ramus 
riLM as deep in juveniles as in adulis. Posterior 
menial foramen varies from bcnealli Mj/Mj. (ho- 



284 



MEMOIRS OF THE QUEENSLAND MUSEUM 






FIG. 1 . QMF30390, Holotype, Nowulgeematrixsp. nov. A. buccal view; B, lingua! view; C, siereopair of occlusal 
view. Scales = 10mm. AR number is that of the Archer colleciion. University of New South Wales. 



lotype) to as far anterior as beneath hypolophid 
ofMi.QMF20080prc.scrvcs angular process and 
much of ascending ramus but lacks condyle and 
coronoid process. Lingual margin of angular pro- 
cess low. aligned with molar row. posterior mar- 
gin sloping ventrally towards lingual end. 
Pterygoid fossa triangular in dorsal view, buccal 
margin slightly undercutting base of ascending 
ramus. Mandibular foramen a narrow, vertically 
oriented ellipse, opening to very short posterior 
portion of inferior dental canal opening via mas- 
seteric foramen to masseteric fossa. Masscicnc 
foramen just visible when masseteric fossa 
viewed from buccal side. Molar row ventrally 
concave. Ascending ramus at about I \3'^ lo line 
of a straight edge laid on molar row. 

LOWER DENTITION- dP2 and dP3 preserved in 
QMF2()080 and 20063. detached dP3 available 
for QMF30392. dP: (Fig. 2A) short, blade-like, 
with bulbous base tapering anteriorly and poste- 
riorly. Occlusal margin straight, relatively hori- 



zontal, with 4 small cuspids anterior lo longer, 
posterior cuspid overhanging posterior base of 
tooth. Fifth cuspid incompletely differeniialed 
from large, posterior cuspid in QMF20080. 
Transcristids associated with each of 4 anterior 
cuspids. Posterior, buccal face of crown abutting 
anterior lingual face of dP3. 

dP3 (Fig. 2A) narrower anteriorly than posteri- 
oriy. dominated by massive, laterally compressed 
protoconid, the tallest cuspid on tooth. No distinct 
metaconid. Paracrislid anterolingually directed, 
descending lo paraconid on anterior margin. 
Cnstid descending steeply frt)m paraconid apex 
to crown base on buccal side of anterior margin. 
Cristid descending posterior face of protoconid to 
interlophid valley. Paraconid, paracristid and 
protoconid form blade-like crest complementing 
that of dPa. Entoconid taller, more angular than 
hypoconid. Cristid obliqua running anterolingu- 
ally from apex of hypoconid, crossing interlophid 
valley and ascending buccal ilank of protoconid. 
In QMF20069 and QMF30392 ashort, buccally- 



NEW BULUNGAMAYINE KANGAROOS 



^85 





FIG. 2. Paralypes of Nowidgee matrix sp. nov. A. stereopair of occlusal view of QMF20080, righi deniary 
fragmeni with dP:-?, Mi-.-^. (M4). B. stereopair of occlusal view of QMF30395. right maxillary fragment with 
P3, M1.4. Visible number is that of the Arclicr collection, Universitv of New South Wales. 



directed proioslylid crest joining proloconid apex 
10 a prominence (reduced protostylid), contacted 
by anterior end of cristid obliqua. Sharp pre- 
eniocrislid running to inlerlophid valley from en- 
loconid apex. Buccally-direcied crest from 
cnloconid descending steeply buccaliy from cn- 
toconid apex to about midline of loolh. 
Posthypocristid descending lingually from 
hypoconid apex loshorterpostcntocnslid ascend- 
ing to entoconid apex. 



P3 in QMF19937 and 30394 resembles holo- 
lype but with seventh minor cuspid, imperfectly 
differentiated from long posterior cuspid of oc- 
clusal crest. Seventh minor cuspid more clearly 
differentiated in P3's removed from crypts in 
QMF20080 and QMF30392. 

Except forQMF19961 in which anterior molars 
very worn, molar teeth in paralypes less worn 
than lho.se of holoiype. Molar morphology among 
paralypes very similar to holoiype. 



286 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 1 . Denial parameters for type specimens of Nowidgee mairi.x sp. nov. 



Number 


P2 

1 


P2 

w 


P2 
h 


P2 
ten 


P3 

1 


P3 
w 


P3 
h 


P3 
ten 


dP3 

1 


dP3 
aw 


dP3 
pw 


Ml 

1 


M1 
aw 


Ml 
pw 


M2 
1 


M2 
aw 


M2 
pw 


M3 

1 


M3 
aw 


M3 
pw 


M4 

1 


M4 
aw 


M4 
pw 


F30390 










6.0 


3.2 


3.6 


6 








3.9 


2.8 


2.96 


4.1 


3.1 


3.2 


4.1 


3.2 


3.0 


4.2 


3.1 


2.8 


F20080 


3.9 


3.4 


3.4 


4 










3.4 


2.2 


2.6 


3.8 


2.4 


3.0 


4.0 


2.5 


2.9 


3.7 


2.6 


3.0 








F20069 


3.3 


2.7 


3.0 


4 


6.4 


3.2 


3.5 


6 


3.4 


2.0 


2.3 


3.8 


2.7 


2.7 


4.2 


3.1 


2.8 














F30391 
























3.9 


2.6 


2.8 


3.8 


3.0 


3.0 














F19937 










6.4 


3.6 


4.5 


6 
































F22761 
























3.3 


1.6 


2,0 


3.6 


2.3 


2.3 














F30392 


















3.4 


2.2 


2.3 


3.6 


2.6 


2.5 


4.2 


2.5 


2.9 


4.2 


2.5 


2.9 








F19961 
























4.2 


2.5 


2.5 


4.2 


3.1 


3.1 


3.9 


2.8 


2.8 


3.8 


2.7 


2.5 


F20255 




































4.5 


3.4 


3.1 


4.1 


3.0 


2.8 


F19991 




































3.9 


3.1 


3.0 


4.1 


3.1 




F30393 


















3.3 


2.4 


2.5 


3.7 


2.5 


2.5 




















F30394 










7.1 


3.2 


3.8 


6 








4.0 


2.2 


2.6 


3.8 


2.9 


3..0 


4.0 


3.1 


3.0 


4.0 


2.9 


2.7 


MEAN 


3.6 


3.1 


3.2 


4 


6.5 


3.4 


3.9 


6 


3.4 


2.2 


2.4 


3.8 


2.4 


2.6 


4.0 


2.8 


2.9 


3.5 


3.0 


3.0 


4.0 


3.0 


2.7 


SD 


.4 


.5 


.3 





.5 


.3 


.5 





.1 


.2 


.2 


.3 


•4 


.3 


.2 


.3 


.3 


1.5 


.3 


.1 


.2 


.2 


.1 


F30395 










7.6 


- 


3.9 


6 








4.1 


3.8 


3.7 


4.0 


4.2 


4.0 


4.0 


4.1 


3.6 


3.7 


3.4 


2.7 



2) Maxillary Iragment. QMF30395 occludes 
extremely well with dentary fragment, 
QMF30394 found in close proximity. Preserves 
buccal surface of maxilla from diastemal region 
to masseteric process, including ventral margin 
of infraorbital foramen, suborbital shelf, alveolar 
process containing entire cheek tooth row and 
very narrow portion of palatine wing. Masseteric 
process of no more than a ventral prominence 
separated from alveolar process by short, narrow 
sulcus. Maxillary foramen of infraorbital canal at 
anterolingual corner of triangular suborbital shelf 
of maxilla, numerous smaller foramina within 
ventral margin of foramen. Infraorbital foramen 
dorsal to midpoint of P\ 

UPPER DENTITION. (Fig. 2B). Molar row 
slightly convex in lateral view, occlusal edge of 
P-^ aligned with buccal margin of molar row 
which curves slightly lingually anteriorly. Molar 
size increasing from M' to M^' M'^ markedly 
smaller than M-^. 

P^ almost twice length of M', lingual margin 
damaged, buccal tnargin convex for 2/3 length, 
becoming concave for remainder. Occlusal mar- 
gin anteropostcriorly straight, on midline of 
tooth. Six small cuspules on margin, succeeded 
by larger, posterior cuspule which has strong 
lingual ridge associated with its ba.se. 



M' with straight anterior and buccal margins 
and convex lingual and posterior margins. Ante- 
rior width greater than posterior width but pro- 
toloph and metaloph of about equal length. Low 
crowned with lingual cusps more massive and 
more rounded than buccal counterparts. Buccal 
cusps closer to lateral margin of the tooth: buccal 
surfaces of crown almost vertical, lingual sur- 
faces sloping. Narrow lingual cingulum reaching 
from anterior, lingual base of protocone to base 
of metaconule. Protoloph formed by strong 
paracrista directed lingually from paracone apex 
and which meets buccal tlank of proloconc below 
apex. Preparacrista runs anteriorly from paracone 
apex to anterior margin and is continuous with 
anterior margin of paracingulum bounded later- 
ally by preparacrista and anterobuccally inclined 
preprolocrista which meets anterior margin ante- 
rior to junction of paracrista with protocone. Very 
large slylar cusp C closing inlerloph valley on 
buccal side. Postparacrista and premetacrista 
reaching floor of interloph valley from respective 
cusp apices, but not united. Postprotocrisla 
strongly developed but worn in interloph region, 
contacting metaloph crest just buccal to apex of 
metaconule. Prominent neometaconule at about 
centre of metaloph with rounded crista running 
posteriorly for about half height of metaloph. 
Postmetaconule crista buccally inclined on pos- 



NBW BULL^NGAMAYIKE KANGAROOS 



2R7 



rwnr fi^^c of nieiaconule, crossing posieriorbase 
urmetaltiph as margin of strong mctacingulum^ 
ixmiiuUng iK»sU'norly direclcd postJiiclacrisla al 
bascormctaconc. 

M-cunsideraM> wider antct'iorly than posrcri- 
Orly. occlusal outline more hlunlly Iriaiigular. 
Crown differing from M' in lingujil ciii^uhini 
continuous with prccingulum extending anteri- 
orly across hasc of protoconc to anterior end of 
prcprotoerisiav siylar eusp C sHghily mure anle- 
rit>Hy posihoned on buccal I lank ol paracone and 
docs not completely close buccal end ofinterloph 
valley; neorneiacotuile and its crista tcs.sobvii»us. 
M* very similar to M- but lingual cingulum noi 
as sharply defined, siylar cusp C smaller, 
poslparacrista and prcmctacrista anile lo form 
continuous cenLrocrisia. M"* much smaller than 
anlerioi nmlars, Metaloph markedly shorter Ihan 
proluloph. Lingua] cingulum separated Imm prc- 
cingulum. ;ill cristae sharply defined. MeUuronulc 
lower; no ncomctaconulc or stylar cusp C 

REMARKS. Nowulgee matrix is similar in "«i^e 

lo Buhmgamaxa dclkaui hut has bunolophodont. 
rather than lt)phodt>nt molars. Its bunolophodont 
lower molars resemble ihoscof P/i/T/W mosaivus. 
but molar occlu.sal ouilinc ui A' matrix is mure 
rectangular, rather than square as in P. nwsaictis. 
P3 of /V. /nrt//V.r differs from thai t.»f /'. mosiucii'i 
in having 6-7 rather than S Iranscrislids and. while 
Having a bulbous base, lacks the disiiticl lateral 
cingulids of Z'. ntosaicus. ft differs from P. tno.uii' 
ctts in having an Ii which has bojh ventral and 
dors-al enamel llangcs and in having enamel 
which, while confined 10 |hc buccal surface, ex- 
tends over that surlacc rather than bemg confined 
to its ventral portion as in P. mosaicus and other 
poluroines. LowcrmuLu*s of A'. niafrixnTC smiiiar 
lo lower molars from the Tarkarooiot> Local 
Fauna assigned by Flannery and Rich (1^X6) to 
Cfiituardee, hui differ from them by being smaller 
in .size. F3 t»f A', matrix has 6-7 iranscnstids, 
apparently many fewer than the Vy iVoni ihe Tar- 
karooloo Gw«u;rt7t'i', in which the poslenor half. 
the only portion feeovered, has 6 iranscrislids. 

Amupg poiuToids a dorsal enamel llange on li 
is confined 10 Hypsiprimnodon, Pownms, 
bulungamayines (Flannery ct al., \^M) and 
Milliyowi Inrngcmdit} (Flannery et al* 1992). N. 
matrix diflcrs from HypsiprirmwJon by having 
permanent premolars which are elongated with 
horizontal or concave occlusal margins rather 
Ihan plagiaulacoid withciinvex occlusal margins, 
|iv failure 10 ret am dP: after rhcerupnou of ^^ and 
hy havmg less dispiiruy between Ihe lengths of 



protolophid and hypolophid on Mi. N. matrix 
differs from Potoroiis by having lower molars in 
which the buccal cuspids are positioned lingual 
lo the adjaceni hneral margin wiib the result thai 
buccal crown walls axe not as sleep as ifi Pator- 
mts. N. matrix differs from the similarly strongly 
bunolophodi>nl early Pliocene Milliyowi bun- 
yandii} in having a strongly developed stylar cusp 
C on M' (abscnl m M. btrnjiantij) and in lacking 
branches oi the transcristids of P3. 

The resemblance of the Ii dorsal enamel llangc 
In that uluiacropttdids IS suggestive of a similarly 
macropodid-like cutting action during occlusion 
of up(K*r ;ind lower incisors, an unusual feamre in 
an animal with bunolophodont molars similario 
Ihose of omnivorous poloroids in which incisucs 
perform a more forcipulate function. 

The postcnwenirally inclmed buccinator sul- 
cus ill /v. matrix was icnned ihe 'labial groove' 
by Slirton ( 1%3) who noted it in Protemtwdon 
and oihcr macropodids. Woodburnc (1967) re- 
ported a similar siructure in Hadronomus 
jHwiirid^i. Where such a sulcus occurs among 
macropodids and other poloroids il is usually 
closer to ^umS parallels the alveolar margin. 

The reduced cuspid on the buccal flank ofllfc 
large, central cuspid of the dP^ trigonid which fs 
linked by ndges tu the ape\ of that cuspid ami to 
the crisiid oblii)ua, is interpreted herein as a re- 
duced protoslylid since ii ot:curs in the corre- 
sponding position and hears the same relationship 
lo the crisiid obliqua as do similar structures on 
Ml of other species, i.e., the pnUoslylid crest of 
Wumroo dnyamayi Cooke, (his volume, ihc dis- 
crete protoslylid o\ Nambaroo salfavna Flannery 
& Rich, I9K6. and the protoslylid of fti/((t'<Y'"*^<?r- 
ous prisctts. The dominant ingtmid cuspid tin* 
gually adjaceni to the protoslylid on dP.^ of .V. 
matrix and from which the paracristid arises is 
thus the piotocomd and Ihe mctac{>nid has been 
lost. The loss of Ihe metaci»nid uf dPi m^y he, as 
suggested by Ride ( 1993), the result of a need In 
supplement the shearing crest of dP.^ which is 
shorter ih;in the permanent premolar in this spis 
cics. 

Apan from ilie drscreie proioslylid rather than 
aproiostylar ridge, the holoiype looth of 7' pris^ 
cus, designated as Mi (their M2) by Flannery Si 
Rich (19S6), bears strong sinnlaiilies to dP^ ii) 
paralypcs ol M matrix. Undescribed Riversleigh 
bulungamayines also have a proiosiylat ridge on 
dP^ and a postcrobuccally inclined protolophid 
similar lo P. prisctts. Since Ihe lailcr characiet 
dr-cs not occur oh nviJai iccih of plcsionioi7ihic 
spescfcs such ?is V. fPWix which have otherwise 



288 



MEMOIRS OF THE QUEENSLAND MUSEUM 



similar bunolophodont molars, it is suggested that 
the hololype tooth of P. prisons is dP3 rather than 
Ml. If this is the case, P. prisciis must still be 
regarded as more plesiomorjihic than N. matrix in 
view of the discrete protoslylid on this tooth, but 
other differences in this tooth, or in molars re- 
ferred to this species, are here regarded as insuf- 
ficient to warrant the erection of a new subfamily. 
Subfamilial affinities of the species remain un- 
certain: its bunolophodont molar morphology 
suggests it may represent either a plesiomorphic 
potoroine or bulungamayine. However, the dis- 
crete protoslylid on the holoiype {dP3) is 
plesiomoqihic and the species may prove to be 
basal to both these taxa. 

Lower molars in N. matrix are suitable to be 
ancestral to B. delicara. Lophids in N. matrix arc 
clearly formed by transverse cristids extending 
buccally from the lingual cuspids. The posterior 
cingulid is enclosed by the poslhypocristid which 
sweeps lingually posterior to the hypolophid and 
low on the crown before linking to the postento- 
cristid on the posterior o[ the entoconid. In B. 
delicata the protolophid is formed in a manner 
similar to that of N. matrix but joins the pro- 
toconid closer to its apex. The posthypocrislid is 
more elevated on the crown, more transversely 
oriented and links to the entoconid much closer 
to the entoconid apex. The buccally oriented crest 
from the entoconid is reduced in length and in 
prominence, the posthypocrislid having formed a 
neomorphic hypolophid. 

In the low-crowned upper molars of A', matrix, 
lophs are formed by crislae extending lingually 
from the buccal cusps, upper and lower molars 
showing reversed symmetry in this respect. Lon- 
gitudinal crests, notably the pre- and postproto- 
crista are emphasised, as they arc in Gumardee 
pasciiali. Strong longitudinal cristae characterise 
bunolophodont upper molars as defined by 
Flannery et al. (1984) who suggested that these 
might work in a different way to lophodoni mo- 
lars in which transverse rather than longitudinal 
cutting crests are emphasised. 

In some undescribed plesiomorphic 
Riversleigh balbarines (pers. obs.) stylar cusps C 
and D or their stylar crests are present in ihc 
interloph region. A', matrix retains only stylar 
cusp C and lacks any trace of stylar cusp D. While 
both balbarines and bulungamayincs arc likely to 
be derived from bunolophodont ancestors, the 
absence o\' stylar cusps other than C in what is an 
extremely plesiomorphic liulungamayinc. sug- 
gests thai loss of other stylar cusps had already 
occurred in the bulungamayine ancestor which 



must in this aspect at least, be more derived than 
that of balbarines. 

Ganguroo gen. nov. 

TYPE SPECIES. Ganguroo bilamina sp. nov. 

DIAGNOSIS. Bulungamayincs with lower molars 
which are completely bilophodont, lacking any trace of 
a buccal ly-directed crest originating from the en- 
toconid and anterior to the hypolophid. 

REMARKS. Ganguroo gen. nov. differs from all 
potoroines, hypsiprimnodonlines and pro- 
pleopines by having bilophodont lower molars. 
It differs from al! macropodines and slhenurines 
by having a combination of: low-crowned mo- 
lars; tlnely-ridged, elongate premolars; a deeply 
penetrating masseteric canal conlluent over its 
length with the inferior dental canal. It differs 
from all balbarines by having the elongate, finely- 
ridged premolars referred to above, lacking a 
transversely compressed trigonid on Mj and in 
lacking a posterior cingulid on lower molars. 

ETYMOLOGY. Waanyi (as spoken by Ivy Slinkcn, 
formerly oi Riversleigh Station) gangu, grandfather 
and Voo' is a common Australian diminutive for kan- 
garoo. 

Ganguroo bilamina sp. nov. 
(Fig.3,Table2) 

DIAGNOSIS. As for the genus 

MATERIAL EXAMINED. Holotype QMF19915 
from Wayne's Wok, Hal's Hill' D-Siie Plateau. 
ParaiypesQMFI959l, 18810, 19814, 19835.30398, 
30399 from Wayne's Wok Site; QMF19868, 19870. 
19966, 30400 from Camel Sputum Site. Godthelp's 
Hill: QMF19642, 20293. 30396. 30397 from Upper 
Site. Godthelp's Hill; QMFI9988 lYom Mike's Me- 
nagerie Site, Godthelp's Hill; QMF23777 from Bites 
Anlennary Site,eastem part of D Site Plateau. All 
System B, Miocene sites (Archer et al., 1989). 

ETYMOLOGY. Latin lamina, blade; refers to the 
bilophodont lower molars. 

DESCRIPTION OF HOLOTYPE. Left dentary 
including horizontal ratnus, most of angular pro- 
cess and part of ascending ramus. Ii, P;^and Mi -4 
preserved. Ventral margin o[ horizontal ramus 
strongly convex, deepest below protolophid of 
IVI3. distinct digastric prominence on the ventral 
margin at this point. Diastema relatively shtirt. 
less than 20% of length of cheek tooth row. 
Slender 1 1 almost horizontal with dorsal occlusal 



NEWBLILUNGAMAYINEKANGARCIOS 



iK9 



ina/gin well below plahe of cheek tooth row. 
Alveolus for very small h on dorsal margin of 
diaslcma jusl posterior to margin ol li alveokis. 
Merual foramen close U* dorsal oiargin ofUia- 
slemM below anienor margin of ^*3. 2 y/cry small 
posterior mental foramina more posicriorly^ I 
below hypulophid o( Mj. the other bekiw pav 
toUiphitl of M-^. Very shallow sulcus lor diiafh- 
meni of biiccioatnr muscle slr>ping diagonally 
vcmrally on buccal surface below M1-M2. As- 
cending ramus ai about 105° lo line of a straight 
edge laid across high points of occlusal surfiices 
uf check tooth row. Since tooth row coivcijve 
clorsally, such a line contacts posterior orP3 aiuJ 
hvpolophid of M^. Buccal margm of masseieric 
fossa straight with tlat area for aiiachmeni of parts 
of superlicial layer of masseler extending ;^n- 
lerovenirally. Masseteric canal and inferior den- 
tal canal confluent anterior lo large masseteric 
foramen. Diameter of foramen and ;uitcrior con- 
sinclion of common canal suggest insenion of 
deep layer of masseier unlikely to have rcaichcd 
(liuch mt)rc anicriorly than M^. Posterior to mas- 
seteric foramen inferior dental camil very short: 
nuisseteric foiamen almosi overlapped by man- 
diliular foramen. Lingual miirgm ol angular pro- 
cess aligned with molar rov^'. Wide, shallow basin 
of pterygoid fossa overhung buccall) by rcmain- 
ittg anterior portion of ascending ramus. Mandib- 
ular symphysis decreases in height paslciiorty. 
extends to level of posterior margin of P3, 

Dcviition Cheek tooth row aniefopt>sieri<>rly 
straight; P3 ilcNcd slightly buccally oulof align- 
ineni. In lateral view occlusal margin of P^ above 
thaiofmolars. Molars low crowned. bilophoi)*»m- 
no trace of imy buccally directed crest associated 
with the cntoconid. Molar si/e increases from Mi 
lo M3; M4 is smaller than Mr 

li long and slender with low dorsal and ventral 
enamel flanges. Dorsal and ventnil n>argins al- 
most hofi/onlal U)\ most of length bclore latter 
converges on lormcr at anterior end. f^namei 
confined to buccal surface. Cross section circular 
cK)5C lo alveolar margm, triangular anterimrly, 
resulting from development of rounded, longitu- 
dinal ridge central lo lingual Mirface. 

Vs elongate, hlade-like with mostly hori/x>nlal 
occlusal margin serrated over most of length un- 
lerior K» large. j>osieiior cuspid taller than rest of 
tooth. Serrations formed by 6 mi nor cuspids, each 
with associated transciisiids. most posienor ol 
these least distinct and shortest. In fxxlusal view 
crt>wn base lapenng anteriorly and posteriorly to 
iToundcd nxargijis. Buccal margm Lonvcx. linguiil 
l?l%irgin r^laiively straight. Lingual base of cnmn 



bulbous adjacent 10 central region of occlusal 
blade, forming poorly-defined, rounded Ungual 
cingulid. Occlusal margm folliming midline hut 
posterior cuspid tlexcd lingually. Crisiid ck* 
scending anieriotiy from apex of most antcriiw 
cuspid lu crown base and posterior myrgm delin* 
oaicd by Nimilai hut shorter crisiid descending 
from posierjurcuspid. 

Mj ^ubrcciangularin (xclusal outline, narrower 
anieriorly than posteriorly. Lingual walls of 
crown subvcrticaf buccal side sloping n)OfC 
gently. Tooth worn, more wc^Ji on buccal slile. 
Wear facet on pi"isierior of metaconld anJ 
breaches in enamel of protoeonid. hypoconid and 
cntoconid. Melaconid UiUer, more anguUir thcUi 
protoconid which has been nsduced itt height by 
weai Sharply defined eresi of proiolophid de- 
scending buccally fmmape.x uf melaconid to i^wlt 
of pmtocotud. Sliort pa/acrislid mnning anteri- 
orly from base uf pa>toconid to anterior margin. 
Steeply descending cnamc! ridgcon anicrior mar- 
gin buccal to paracristid. forming margin of slvin 
precingulid. I3road anteriorcrngulid enclosed be- 
tween paraciisiid and prcmciacristid which runs 
between melaconid apex and anieriot margin, 
Sharp postmctacrisiid descending posteriorly 
from metaconid apex lo interlophid valley, meet- 
ing similarly wcll-deHncd precntocnstid running 
anteriorly from apex of the enloconid. Crisiid 
oblicjua worn, running anterolingually on anicrior 
face of hypoconul betore turning anteriorly 
across interlophid valley, meeting posienor face 
of proiolophid lingually adjaccnl to protoconid 
base. Enloconid taller than hypoconid which Ls 
more worn. Crest of hypolophid crossing from 
posterior of hypnconid apex to meet entoc<inid 
apexccntrahy Postentocrisiiddcscendmg poste- 
rior face of enloconid but no other ornameniaiion 
n I posterior lace of hypulophid. M: similar in 
outline lo M» but slightly larger and less worn. 
Metaconid taller than protoconid but hy|>oconid 
and enloconid subequal in lioighi, Postmcta- 
crisiid and prccmocristid not uniting in inter- 
lophid valley. M,t largest of molars, unworn, with 
all cuspids about equal height: lingual cus[>id.s 
with more angulai aj^iccs than buccal cuspids. Mi 
damaged, lacking protoconid and most of anterior 
margin. Hypoconid taJlcr than enloconid; no 
poslent;<ais(id 

DESCRIPTION OFPARA'n'PES. Condyle piv- 
servcd in QMFI98t4, Ilj«70. 19810 and :W96. 
In QMF 19870 and I98I4ks trans vciwlyclnngaic 
with rounded posterior margin. In QMF3IJ396 
condyle has mote circular outline. Thm ol* 



290 



MEMOIRS OF THE QUEENSLAND MUSEUM 






FIG. 3. QMF19915, Holotype of Ganguroo bilamina sp, nov. A, buccal view; B, lingual view; C, slereopairof 
occlusal view. Scales = 10mm. 



QMF19810 slightly damaged lingually but sim- 
ilar to, although somewhat smaller than, that of 
QMF19870 and QMF19814. Differences in 
shape possibly age related since QMF30396 is 
from a subadult animal, indicated by in- 
completely erupted P3. Height of condyle above 
plane of molar row varies from 7mm in QMF 
19870 and QMF30396 to 1 1mm in QMF 1 98 10, 
variation possibly being size related. Angle of 
ascending ramus relative to plane of molar row 
varies 120° tARI2517)-108° (QMF18814). 

Digastric process on ventral margin of horizon- 
tal ramus apparently variable within species: 
QMF19814 level of development comparable to 
that of holotype, but other paratypes show lesser 
or no such development. Number of posterior 
mental foramina also variable: most paratypes 
have only one such foramen, consistently located 
below M2, but none present in QMF30398 while 
two present in QMF30400 and QMF 19966. 
QMF 19988 has number of smaller foramina ac- 
cessory to mandibular foramen and also has sul- 



cus for vessels of inferior dental canal on lingual 
wall of masseteric fossa. Posterior portion of 
inferior dental canal between masseteric foramen 
and mandibular foramen longer in AR 1 25 1 7 than 
in holotype and most other paratypes. QMF30400 
and QMF19988 have direct opening via single 
foramen from pterygoid fossa into masseteric 
fossa with no intervening canal (the condition 
usual among extant macropodoids). 

Damage to ventral margin of horizontal ramus 
reveals extent of anterior insertion of masseter, in 
QMF 1 9868 and QMF20293 it reaches level of 
M: hypolophid, but only to level of M^ 
hypolophid in QMF30397. 

DENTITION. dP2 and dP3 in QMFl 9835, 23777. 
dP2 small, blade-like with rounded anterior and 
posterior margins, strongly convex buccal mar- 
gin and straight lingual margin. Occlusal crest 
serrated over much of length anterior to large 
posterior cuspid overhanging posterior base of 
crown. QMF19835 has 3 small cuspids in ser- 
rated region, each with associated transcristids; 4 



NEW BULUNGAMAYINE KANGAROOS 



291 



TABLE 2. Dental parameters for type specimens of Ganguroo b'damina sp. nov. 



Number 


1 


li 
w 


P2 

1 


P2 

mw 


P2 

h 


Pa 

ten 


P3 

1 


P3 

mw 


P3 

h 


P3 

ten 


dP3 

I 


dPj 
aw 


dP3 
pw 


Mi 

1 


Ml 
aw 


Ml 
pw 


M2 


M2 

aw 


M2 

pw 


M3 

1 


Ma 
aw 


Ma 
pw 


M4 

1 


M4 

aw 


M4 

pw 


F19915 


11.6 


2.2 










5.9 


3.3 


3.9 


6 








3.7 


2.4 


2.6 


3.7 


2.7 


2.8 


3.7 


2.6 


2.6 


3.8 


- 


2.6 


F30400 


































3.9 


2.8 


2.9 


3.7 


2,8 


2.8 


3.8 


2.8 


2.7 


^IZlll 






4.2 


2.5 


3.1 


3 










3.5 


2.0 


3.3 


3.6 


2.8 


3.0 




















F19868 


10.9 


2.1 










5.9 


3.1 


3.7 


5 














3.8 


2.9 


2.8 


3.8 


2.9 


2.9 








F19870 






















4.2 


2.6 


2.8 


4.0 


2.8 


3.0 


3.8 


3.1 


3.1 


4.0 


2.9 


2.8 








F19966 














5.9 


3,1 


3,9 


6 


3.4 


2.6 


2.8 


3.8 


2.8 


3.0 


3.9 


3.1 


2.9 


3.9 


3.0 


2,7 








F19988 






































2.8 


3.8 


2,8 


2.8 


3.6 


2.9 


2.5 


F30396 




1.7 










5.3 


2.4 




6 








3.4 


2.6 


2.6 


3.7 


2.7 


2.7 


3.9 


2.9 


2.7 








F30397 














6.3 


3.1 


3.7 


7 








3.5 


2.3 


2.5 


3.9 


2.7 


2.8 


3.9 


2.9 


2.8 


3.9 


2.7 


2.5 


F 19642 














5.6 


2.3 


3.9 


6 








3.5 


2.1 


2.4 




















F20293 


































4.1 


3.0 


3.0 


3.9 


3.0 


2.9 


3.9 


2.9 


2.9 


F30399 














6.2 


2.8 


3.6 


7 








3.5 


2.2 


2.7 


3.5 


2.5 


2.9 


3.6 


2.7 


2.8 


3.8 


2.7 


2.7 


F30398 














6.1 


3.0 


3.5 


6 








3.5 


2.6 


2.7 


3.6 


3.0 


2.9 


3.8 


3.0 


2.8 


3.5 


2.8 


2.3 


F 19591 








































3.9 


3.2 


2.7 








F19B10 




























3.6 


2.6 


2.6 


3.8 


2.8 


2.8 


3.8 


2.7 


2.9 


3.7 


2.8 


- 


F19814 














5.9 


3.2 


3.7 


6 








3,6 


2.5 


2.8 


3.7 


2.6 


3.0 














F19835 






3.4 


3.0 


3.6 


4 


5.7 


2.8 


4.0 


5 


2.9 


2.3 


2.3 


3.1 


2.5 


2.7 




















MEAN 


11.3 


2.0 


3.8 


2.8 


3.4 


3.5 


5.9 


2.9 


3.8 


6 


3.5 


2.4 


2.8 


3.6 


2.6 


2.7 


3.7 


2.9 


2.9 


3.8 


3.0 


2.8 


3.7 


2.9 


2.6 


SD 


4.9 


.3 


.6 


.4 


.4 


.7 


.3 


.3 


.2 


.7 


.5 


.3 


.4 


.2 


.4 


.2 


.4 


.3 


.1 


.3 


.3 


.1 


.3 


.4 


.2 



such cuspids in QMF23777. Anterior and poste- 
rior margins of crown delineated by vertical 
cristids descending from ends of occlusal crest. 
Occlusal crest runs slightly lingual to midline, 
lingual surface of crown more steeply inclined 
than buccal. 

dP3 better preserved in QMF23777 and used as 
basis for description below. Crown base roughly 
rectangular in occlusal outline, narrowing some- 
what anterioriy. Protolophid extremely laterally 
compressed, inclined posterolingually, domi- 
nated by tall protoconid with thick, rounded pro- 
tostylid cresl descending its buccal flank. 
Metaconid cannot be distinguished from pro- 
toconid. Prominent paracristid (less so in 
QMFI9835) runs anteriorly to tail paraconid 
(shorter in QMF19385) on anterior margin. 
Paraconid, paracristid and protolophid form 
blade-like crest complementing that of dP2. Ver- 
tical cristid descends from posterior margin of 
protolophid cresl to interiophid valley and is con- 
tacted by anteriorly directed preentocristid in 
QMF19835, but not in this specimen. 
Hypolophid transversely oriented, concave in 
posterior view. Cristid obliqua runs anterolingu- 
ally on anterior face of hypoconid, turning ante- 
riorly across interiophid valley and contacting 
proiostylid crest. No ornamentation on posterior 
face of hypolophid. 



P3 in most paratypes closely resembles thai of 
holotype but QMF30399 has 7 minor cuspids 
rather than 6. 

Molar morphology very similar to that of the 
holotype although variable postenlocristid be- 
tween different specimens and between different 
teeth in single specimens. 

DISCUSSION 

The horizontal orientation of Ii is similar to that 
in macropodines in which there is considerable 
ventral Hexion of the rostrum, necessary to bring 
upper and lower incisors into occlusion and there 
would presumably be a coiresponding tlexion of 
the rostrum in this species. dP3 is very similar to 
that of N. matrix but is more derived in that the 
reduced protoslylid of N. matrix is here further 
reduced to a protostylid crest. Molars in this 
species are more derived than in either N. matrix, 
B. delicaia or Wabularoo naughtonibecsiusQ they 
are lophodont. 

N, matrix, B. delicata and G. bilamina represent 
stages in an evolutionary sequence in which a 
bunolophodonl, omnivorous ancestral form is 
changed to that of a lophodont herbivore (Fig. 4). 
Hypolophid morphology is particularly informa- 
tive in this respect. As discussed earlier, a 
neomorphic hypolophid has been developed in B. 
delicata by elevation of the posthypocristid on 



292 



MEMOIRS OF THE QUEENSLAND MUSEUM 



the crown and directing the posihypocristid more 
transversely. Hypolophid morphology in W. 
naughtoni closely resembles that of B. delicata. 
The bunolophodont origin of this morphology is 
indicated by the reduced buccal crest from the 
entoconid anterior to the new hypolophid, repre- 
senting the remnant of the original hypolophid 
crest. No trace of this crest is evident in G. 
hilamina, the neomorphic hypolophid crest being 
formed entirely by the elevated, transverse 
posthypocristid as indicated by the presence of a 
postentocristid on the posterior face of the en- 
toconid. Loss of the buccally-directed crest from 
the entoconid represents a subtle change in mor- 
phology between N. matrix and G. bilamina but 
a highly significant apomorphy. 

The evolutionary series represented by these 
bulungamayine tax a demonstrates that 
lophodonty evolved independently twice among 
Oligocene-Miocene kangaroos - once in 
balbarines in a process which seems to have been 
essentially that proposed by Flannery & Rich 
(1986) and once among bulungamayines using 
the mechanism proposed above. While Flannery 
(1989) suggested that balbarines were ancestral 
to macropodines and sthenurines, the similarity 
of premolar and molar morphology of derived 
bulungamayines such as G. bilamina to that of the 
later Miocene macropodids from Alcoota is 
greater than that of more derived balbarines such 
as Balbaroo in which on Mi there is still consid- 
erable lateral compression of the protolophid and 
little development of the anterior cingulid. The 
premolar of balbarines is also much shorter than 
that of bulungamayines and the plesiomorphic 
Alcoota macropodids (Cooke, 1997). 

Lower molar morphology of G. bilamina has 
strong similarities to that of the much larger 
Hadronomiis puckridgi Woodburne, 1 967 from 
Alcoota which Murray (1991) regarded as a 
plesiomorphic sthenurine. Both species are low- 
crowned and bilophodont, have long anterior 
cingulids, have Mi protolophids which are not 
laterally compressed and lack posterior cingulids. 
although Hadronomus has a bulbous base to the 
hypolophid. Hadronomus also has an elongate 
premolar, resembling in that respect the premolar 
of bulungamayines, but that of Hadronomus is 
more coarsely serrated than that in any of the 
known bulungamayine species and bears well 
developed lingual and buccal cingula, not present 
in bulungamayines. However, paratypes of A^. 
matrix show variable differentiation of minor 
cuspids and transcristids on P3, indicating some 
lability in degree of serration of the occlusal 



margin of this tooth in bulungamayines. The bul- 
bous base of the bulungamayine P3 could serve 
as an adequate precursor of lateral cingula (a 
lingua! cingulum is poorly developed on P3 of G. 
bilamina). The premolar of all known balbarines 
is in contrast a shorter, more plagiaulacoid tt)oth. 

Similarities also exist between dental morphol- 
ogy in G. bilamina and in Dorcopsoides fossilis. 
also from Alcoota. While this species was origi- 
nally included within Potoroidae. Bartholomai 
(1978) placed it in Macropodinae. Both species 
have elongate premolars. Lateral cingula are 
lacking in P3 of Dorcopsoides while a lingual 
cingulum is poorly developed in that of G. 
bilamina and there are again differences in serra- 
tion and transcristids between the two species. 

dP3 in N. matrix and B. bilamina has some 
similarity with that of Dorcopsoides in that the 
metaconid is reduced or absent in each. 
Woodburne (1967) also noted the 'fused pro- 
toconid and metaconid' of dP^ in Dorcopsoides 
and 'a short posterolabial crest ... which turns 
abruptly posteriorly before descending into the 
transverse valley and continues posterolabially 
up the anterior face of the hypoconid'. This crest 
may be homologous with the prolostylid crest 
which is linked to the cristid obliqua of dP3 in N. 
matrix and G. bilamina but which is also present 
on dP3 in undescribed Riversleigh balbarines ref- 
erable to Nambaroo and in which there is also 
considerable abbreviation of the protolophid 
(pers. obs.). Ride (1971) suggested that close 
proximity of the protoconid and metaconid on 
dP3 is plesiomorphic for macropodoids (his 
macropodids), and the protostylid or its reduced 
form of a protostylid crest in both potoroids and 
macropodids suggests that this character is simi- 
larly plesiomorphic. 

While lower molar morphologies in G. 
bilamina and Dorcopsoides are similar in many 
respects, they differ markedly in that 
Dorcopsoides has a well-developed posterior 
cingulid, absent in all bulungamayines but pres- 
ent in balbarines. Derivation of Dorcopsoides 
Irom a bulungamayine ancestor would require 
development of a neomorphic posterior cingulid, 
such development possibly indicated by the 
swollen hypolophid base of Hadronomus. Evolu- 
tion from a balbarine ancestor would require 
modification to both the anterior cingulid and 
compressed protolophid of Mi, but modification 
of pre-existing structures is a more usual evolu- 
tionary phenomenon than the development of 
new structures. This notwithstanding, dental 
morphology in bulungamayines is such that, on 



NEW BULUNGAMAYJNE KANGAROOS 



293 



med 




pec 



phc 




Henk Godthclp for encourage- 
meni and assistance, Anna Gil- 
lespie. Steph Williams and 
others ai the University of New 
South Wales for preparation. 



pho 



FIG. 4, Development of lophodomy in buiungam:iyincs, illustrated by RM|, 
A. Nowidgee matrix , B, equivalent to B. dtlicafcL C, C bilamma. Abbre- 
viaiions: Pr=proioconid, Mc=TnciacoTiid, mcd=mctacristid, 
H Y=hypoconid, Ec=enioconid. ecd=buccai cresi from entoconid. 
phc=pusihypoc'ristid, pec=posientocTisiid, co=L'ristid oblicjua. 



the grounds of parsimony, they, rather than 
balbarines, must be preferred as the group most 
closely ancestral to macropodids. 

In the hypothesis of molar evolution within 
Fiulun^umayinae advanced herein, there is a tran- 
sition from a poioroid-like molar in basal species 
to a macropodid-like molar in derived species. 
Such a transiiional sequence within the group 
may explain the differing views i^'( familial affin- 
ity of bulungamayines (Case, 1984; Woodburne, 
1984; Rannery ci a!.. 1984). At the time their 
respective views were advanced, only 2 bulunga- 
mayinc species, B, delicata and VV. mm^hwnl. 
had been described. Molar morphology in both 
those species i.s intermediate in the transitional 
sequence and it is not surprising thai both 
macropodid and potoroid aflmities could be ar- 
gued on the basis of these species. 

If. as seems likely from the evidence providetl 
herein, that bulungamayines are directly ances- 
tral to macropodids, then monophylyof Bulunga- 
mayinae cannot be staled with certainty. Further 
doubts also arise concerning monophyly of 
Macropodidae. 

ACKNOWLEDGEMENTS 

Research grants from the Australian Research 
Council and the University of New South Wales 
have been the primary mechanism tor providing 
the material studied. Additional support for the 
Riverslcigh project has come Irom the National 
Estates Program grants, the Australian Geograph- 
ical Society, The Australian Museum. The 
Riversleigh Society, ICl Piy Lid. Century Zinc 
Limited, the Ml. Lsa Shire and private donors. 

I thank the Director and slafl of (he Queensland 
Museum for facilities and assistance, Jeff Wright 
for photographic assistance, Mike Archer ajid 



LITERATURE CITED 

APLJN. K.P. & ARCHER. M 19S7. Recent advances 
in marsupial sysiemalics with a new syncretic 
ela.ssiticalion. Pp xv-lxxii in Archer, M (cd) 'Pos- 
sums and Opossums: Studies in Evolution". (Sur- 
rey Beaiiy 8c Son.s: Sydney). 

ARCHER, M. 197Q. Wabularoo naughtom gQW, qX ^y 
nov, an enigmatic kangaroo (Marsupiiiliij) Imm 
the middle Tertiary' Ciiri Creek Limestone of 
nonbwcsiem Queensland. Results of the Ray IL 
Lemlcv Expediiions, piin 4. Memoirs of the 
Queensland Museum 19: 299-307. 

ARCHER, M. 1984. The Australian marsupial ladia- 
tion. Pp. 633-808 In Archer. M, i^ Clayton. G 
(eds) 'Vencbraic Zoogeugrapliy and Evuluiion ui 
Australasia, i Hesperian Press: Perth, Western 
Aaslralia). 

ARCHER. M. & BARTHOLOMAL A. 1978. Tcniary 
mammals of Au-stmlia, a synoptic revievV- AI- 
cheringa'2: 1-19. 

ARCHER, \4- & FLANNERY.T.R 19S5. Revision of 
the extinct gigantic rat kangaroos (Potoroidac: 
Marsupialiaj, with adescription of a new Miocene 
genus and species and a new Pleistocene specicii 
oi' Fropk'opnx, Journal of Palconlulogv 59. 1131- 
1149. 

ARCHER. M., GODTHELP, H , HAND, SJ ^ 
MEGIRIAN. D. 1989. Kossil mammals ol 
Riverslcigh, norlhwesiem Queensland: prelimi- 
nary overview of biosiraiigraphy. correlaiion and 
environmental chance Australian Zoologist 
23l2); 29-65 

BARTHOLOMAI, A. 1 978-The lossil kangaroos. Aus- 
tralian Mammalogy 2: 15-22. 

CASE, J, A 1984 A new genus of Poloroinae 
(Marsupialia: Macropodidae) from the Miocene 
Ngapakaldi Local Fauna. South Australia, and a 
dcfiniiion odhe Poioroinac. Journal of Paleontol- 
ogy 58(4): 1074-1086. 



294 



MEMOIRS OF THE QL^EENSLAND MUSEUM 



COOKE, B.N. 1992. Priniilive mucropodids from 
Riversleigh, northwesiern Queenskmcl. AJ- 
chcnngci 16:201-217. 
1997. Two ticw balbarine kangaroos and lower 
molar evolution wiihin ihc suhfamilv. Memoirs 
ofthc Queensland Museum 41 : 269-280- 

FLANNERY. T.F. 1989. Phylogeny of the 
MacTopodoideu: a study in convergence. Pp I -46. 
In Grigg, C Jarman, P i)t Hume, I. (eds) "Kanga- 
roos, Wallabies and Rat-Kangaroos'- (Surrey 
Beatty & Sons Sydney )- 

FLANNERY, T. & ARCHER. M 19S7a. 
Hxpsipn'iiuiOLJoti hartholaifuui ( Potoroidac: 
Marsupialia). a new species from the Miocene 
Dwamamor Local Fauna and a rcasscssmeni of 
the phylogenelic position of H. waschauts. Pp 
749-758. In Archer, M. (ed). Possums and opos- 
sums; studies in evolution. (Surrey Bealiy & Sonj;: 
Sydney ). 
i987b. Benongia moycsi,\x new and plcsiomorphic 
kangaroo (Marsupialia; Potoroidae) from 
Miocene .sediments of northwesiern Queensland. 
Pp 759 -767. In Archer. M. (ed) 'Possums and 
Opossums: Studies in Evolution'. (Surrey Bcaity 
& Sons. S>dnev). 

FLANNERY, T.F. & RICH, T.H.V. 1986. 
Macropodoids from the Middle Miocene Namba 
Fomvalion, South Australia, and the homology ot* 
some denial struclures in kangaroos. Journal of 
Palconiologv 60(2): 4 1 S-447. 

FL.\N'NERY,T.F. ARCHER, M & PLANE, M 19S2. 
Middle Miocene kangaroos (Macropodoidca; 
Marsupialia) from Ihree localities in northern Aus- 
inslia, with a description of two new subfamilies. 
Bureau of Mineral Resources Journal of Austra- 
lian Geology and Geophysics 7: 287-302. 
1984. Phylogenetic relationships and a reconsidera- 
tion oi' higher level systcmallcs within the 
Potoroidae (Marsupialia). Journal ol Paleontol- 
ogy 58(4): 1 087- 1 097, 

FLANNERY.T.F.,RICH.T-H.,TURNBULL.W'.D.& 
LUNDELIUS- E.L., 1992. The Macropodoidca 
(Marsupialia) o( the early Pliocene Hamilton 
Local Fauna, Vicwria. Australia. FiekUuna Geol. 
25: 1-37. 

FLOWER. WTL 1867. On the deveiopmcnl and suc- 
cession of Icelh m the Marsupialia. Philosophical 



Transactions of the Royal Society oi' London B 
157:631-641- 

LUCKE1T, W.P 1993. An ontogenetic assessment of 
denial homologies in Iherinn mammals, Pp 182- 
204. In S/alay, F,S, Novacek, M.J. &. McKenna, 
M.C. (eds) 'Mammal Phylogeny*. (Springer- Ver- 
lag: New York). 

MURRAY, P.F. 1991. The sthenurinc afFinily o\' the 
Late Miocene kangaroo, Ihidronomas puckrid^i 
Woodbume (Marsupialia. Macropodidae). Al- 
chcrinua 15:255-283. 

RIDE. W.aL. I97L On the fossil evidence of the 
evolution of the Macropodidae. Au.siralian Zool- 
ogist 16(1): 6-16. 
1993. Jackmahoneya gen. nov. and the genesis of 
the macropodiform molar. Memoirs of the Asso- 
ciation of Australasian Palaeontologists 15:441- 
459. 

STIRTON. R.A. 1963. A review of the macropodid 
genus Protemnodon. University oi California 
Publications in the Geological Sciences 44:97- 
162. 

SZALAY, F.S- 1969 Mixodectidae. Micrusyopidac. 
and the Insectivore-Primaie transition. Bulleiin o\ 
the American Museum of Natural History 14(). 
193-330. 

TEDFORD. R.H. & WOODBURNE, M.O. 1987. The 
ilariidac. a new family of vombaliform marsupials 
from Miocene strata o'i South Australia and an 
evaluation of the homology of molar cusps in the 
Diprotodoniia. Pp 401-418. In Archer. M. (ed) 
'Possums and Opossums: Studies in Evolution'- 
(Surrev Beatlv & Sons: Svdne-y). 

WOODBURNE, M.O. 1967. The Alcooia Fauna Cen- 
tral Australia: an integrated palaeonlological and 
geological study, Bureau fj\\' Mineral Resources. 
Geology and Geophysics Bulletin 87. 
1 984. Wakiewakie lawsoni, a new genus and species 
of Potoroinae (Marsupialia; Macropodidae) of 
medial Miocene age. South Australia. Journal o\ 
Paleonlologv 58(4): 1062-1073 

WOODBURNE, MO., MACFADDEN. B.J.. CASE, 
J.A . SPRINGER. MS. PLEDGE. N . POWER, 
J.D.. WOODBURNE. J.M. ik SPRINGER, K.B. 
1993. Land mammal bioslratigraphy and niag- 
netostratigraphy ofthc EtadunnaForniaiion (late 
Oligocene) of South Australia. Journal of Verte- 
brate Paleontology 14: 483-515. 



BIOSTRATIGRAPHIC IMPLICATIONS OF FOSSIL KANGAROOS 
AT RIVERSLOIGH. NORTHWESTERN QUEENSLAND 

B N COOKE 

Cooke. B,N, 1997 06 30; Biostraligraphic implications of fossil kangaroos ai Rivi^rslctgii, 
norihwcslcrn Queensaind Memoirs ofifw Qu^vuslatuf Afiiseiim 41(2^): 295-302. Brisbane. 
ISSN 0079-8835. 

Kangaroos form an imponani component of the faunal assemblages of Riversleigh and most 
other Australian Cainozoic im<;i| deposits, Attempts to use fossil kangaroos to determine llic 
relative ages o( Riversleigh sites suggests that overall faunal composition may he a more 
uselul guide to relative age than presence or absence of parliculur species. Marked changes 
in kangaroo faunal composition occur between Riversleigh System B and C sites with the 
nppareni cximction of most balbarine species and rise to dominance by lophodont 
buluMgamayine species. This change coiTclates with climatic decime following mid-Miocene 
climatic optima. Approximate age equivalence is suggested between Riversleigh System B 
sites and faunal zones B -l-C (Woodbume et al., 1993) of the Eiadunna Formation, South 
Ausiralia. Riversleigh's System A sites are older. The more derived, lophodont 
buliwlgamayincs of Riversleigh System C are considered potentially ancestral to 
picsiomorphic macropodids such as IhuUvnomas from the AlciKHa Formation. Kangaroos 
support an age span for prc-Plioccne deposits at Riversleigh thai extend from the \\\\.it- 
Oligoccnc to late middle Miocene and possibly early late Miocene. [H Kangaroos, 
Oli^oct'fitr, Miocene. Plioane, Riversleigh. 

fi.N, Cooke. School of Life Scieruv, Qtteenslamf Universitv ofTechrialogw Gi^O Box 2434. 
Brifihane, Queensland 400 L An^truliu: J4 Lhremher 1996. 



Tcdfard (I967)prc>vidcc1 Ihf first dciicripliun of 
kangaroos Imin Riversleigh. Since then the area 
has yieklecl many ihousands uf kangaroo fossiks 
representing at least 32 new species. 

Archer ( 1 979) desct ihej Wahnlaroo nnnghrnnt 
from D Sue. Flannery et al. (19H2) described 
Bulun^amaya delicato (erecting the 
Bulungamayinae), W. naui=;hr(»Ni, Halharoo 
^rc\;oriensis from Riversleigh. B. camfiekiensis 
IroniBuIlock Creek. Norlhcrn Territory (middle 
{o Late Mii>ccnc) and a single specimen ol 
Balharoo from Kangaroo Well. The species of 
Ballmroo were placed in a new niacropodid 
Balbarinac. They also described the poloroine 
Cumaniee pascuali and macropodid Calaiuvla 
ti'ssellufu from Riversleigh. 

Archer & Flannery (1985) described the 
Riversleigh proplcopine Ekaitadcta ima. Flann- 
ery 6l Archer |l9K7a) described Hypsiprytnri' 
oilon Intrtluflofnaii and (19S7b) lU'lkfni*iii 
nioyesii, the only pre-Pliocene representalives of* 
these genera Cooke ( 1 992) described the bulbar 
ine Gunawamaya with G. ucris, G. aediiulia and 
G. ornata. 

The number (if Riversleigh fossil kangaroo 
specimens, ranging from isolated teeth, isolated 
posicraniul elements, ntaxillary nnddcntary frag- 
ments to cotnpleie sktdls is alnn>st tiverwhelm- 
ing. I have concentrated on the more complete 



remains available. These indicate a further 21 
now species, a number of which are described in 
this vt)lume. Prorem/wdon sp. has been recorded 
from the Pliocene Rackham's Roust Site and 
Macropus agilis has been recorded from the 
Pleistocene Teiracc Site (M. Archer pers. com.). 

This brmgs the total Riversleigh macropodoid 
fauna to 34, including the Poioroinac, Hypsi- 
prymnodontinae, Propleopinae, Balbarinac. 
Bulungamayinae and Maeropodinae. 

Archcr et al. ( 1 989) recognised mt>re than IIK) 
local faunas from the Riversleigh fossil area, with 
age^ estimated to range from Oligocenc-Miocenc 
to HoKicene. ( Discoveries since 1 9H9 have raised 
that number to more than 150 sites with faunal 
assemblages). They grouped the Oligocenc- 
Miocene sites into three 'Systems' designated 
A-C. with System A sites regarded as oldest and 
System C sites as youngest. 

Mcgirian ( 1992) treated the eniire sequence of 
fossil iferous Limestone overlying the Thornton ia 
Limestone as comprising the Carl Creek Litne- 
slone. He used the 'Systems' of Aivher et ul. 
(1989) in a purely biostraligraphic sense and has 
later (1994) challenged the use of the Sy.stcm 
concept c»n the groimds that the temiinology has 
been unsaiisfaciorily detlned and that there is no 
precedent lor such usage in lithoslratigraf^hic niv 
mcnelaiurc. 



296 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE ]. Sites of occurrence and numbers of specimens of each identified pre-PIiocene Riversleigh 
macropodoid species. QL=Quantum Leap; WH=White Hunter; DU=Dunsinane; COA=Cleft of Ages; 
BA=Bitesantennary; DT=Dirk's Towers; OUT=Outasite; WW=Wayne's Wok; BO=Boid; CS=Camel Sputum; 
IN=Inabeyance; MM=Mike's Menagerie; UP=Upper; CR=Creaser's Ramparts; N'sG=Neviiie's Garden; 
LM=Last Minute; FF=Fireside Favourites; H'sH=Henk's Hollow; TT=Two Trees; DO=Dome; J'sC=Jim's 
Carousel; ENC=Encore; ?=uncertain position within the System sequence of Archer et al. (1989). 



SPECIES 


Syst. A 






?A/B 






Syst. B 


?B 

intr. 


Syst. C 


?mid 
Mio 


Uncert 






Lower 


Higher 


LowC 


MidC 


Higher C 


? 




D 


G 


Q 

L 


W 
H 


D 
U 


C 

A 


B 
A 


D 

T 


R 
S 
O 




u 

T 


w 

w 


B 



C 
S 


N 


M 
M 


u 

p 


c 

R 


N's 

G 


G 

A 
G 


L 
M 


low 


High 
-er 


F 
F 


H-5 

H 


T 
T 


D 

o 


J'sC 


E 
N 
C 


jen. Wan. sp. 1 






































m 














Wr 




1 


Qen.Wan. sp.2 








































1 








"3.."=. 








I 


3u.delicata 


i 










Mitiii m 


q^ 






9 




^~ 


.i 




















/Vab. nauqhtoni 
Gan. bilamina 


H 




.-~— 
















m 




^3 




.t. 


#. 














■ — " 












Gan. sp.2 




































3 


! 








3 






2 




Set. moyesii 


















































^^ 








^. ima 






































^■^ 




















H. bartholomaii 


HI 




































it 




















Now. matrix 








1 












"M^ 




m 


























\low.sp.2 

3a/. qreqoiiensis 




■ 












m 




-fadn^mi 




m 




Si 




■ 


















Bal. sp.2 


















SB 




































Bal. sp,3 


























































Sal. sp.4 














































t 


.2*H 










Gal. lessellata 
























































Gan. acris 
























m 
































Gan. aediculis 










1 














































Gan. omata 


, 










1 









■ 










™ 
























3an. sp.4 
Nur. dayamayi 


— 














— 














H 


— 






















Wur. sp.2 












































Nut. sp.3 


















n 
































^iam. couperi 


















n 
































^am. sp.2 




















■M 
































Nam. sp.3 


















m^ 
































Mam. sp.4 
Nam. sp.5 


— 




id- 








M 




-Ik. 










H 






















Nam. sp.6 
Nam. sp.7 


— 




















M 








H^" 






















Vam. sp.B 1 




1^^^ 


















n 




1 

















Archer et a) . ( 1 989) Archer, Hand and Godthelp 
(1991) and Archer et al. (1994) interpreted the 
older Riversleigh local faunas, those from sites in 
Systems A-C, as representing rainforest commu- 
nities. This interpretation was made on the basis 
of characteristics of those faunas such as: high 
species diversity; presence of complex feeding 
guilds of small, sympatric mammals; high num- 
bers of obligate leaf eaters in single local faunas, 
indicating high tree species diversity; presence of 
high numbers of known rainforest taxa; high 
numbers of browsing marsupials but absence of 
grazers. 

Megirian (1992) disputed the hypothesis of a 
rainforest palaeohabitat for the older Riversleigh 
local faunas on the grounds of his interpretations 
of drier, even semi-arid environmental conditions 
prevailing during the genesis of the fossiliferous 
limestones. He suggested instead that the high 
species diversity evident in the local faunas re- 
sulted from a combination of resident rainforest 
communities inhabiting rainforest refugia con- 
fined to permanent water bodies, and the use of 



such permanent water by animals drawn from 
more distant, mesically adapted communities. 

None of the Pre-Pliocene kangaroos from 
Riversleigh exhibit dental adaptation for a graz- 
ing habit which might be expected if they were 
drawn from a more arid environment. Regardless 
of their palaeohabitat, kangaroos have a wide- 
spread occurrence and are abundant among the 
many local faunas now known from Riversleigh. 
They should prove to be important in attempts to 
determine relative ages of those assemblages. 
This paper presents the results of a preliminary 
attempt to use kangaroo fossils to assess relative 
ages of sites within Riversleigh, assess the age of 
Riversleigh deposits relative to those of other 
Australian Tertiary sites which have yielded 
comparable kangaroo fossils, and to correlate 
changes apparent within the macropodoid fauna 
of Riversleigh with Tertiary climatic and geo- 
logic events. As part of that attempt, the Systems 
concept of Archer et al. (1989) has been used as 
an hypothesis which can be tested using the dis- 



BIOSTRATIGRAPHIC IMPLICATIONS OF FOSSIL KANGAROOS 



297 



Iribution of fossil kangaroo species in sites pre- 
viously assigned to each of systems A, B and C. 

INTER-SYSTEM COMPARISONS AND 
OTHER RELATIONSHIPS AT RIVERSLEIGH 

Pre-Pliocene Riversleigh sites that have 
yielded identified macropodoid species, together 
with the number of specimens of each, are shown 
in Table 1 . 

Column headings in this Table follow Archer 
et al. (1994) who listed sites known to that date, 
indicating those confidently assigned to Systems 
A-C, provided indications of possible System 
affinities of some sites, and estimated ages of 
other sites whose faunal composition differs from 
those allocated to these Systems. As indicated in 
Table 1 and discussed further below, evidence 
drawn from the distribution of kangaroo fossils is 
not in complete agreement with ages suggested 
by Archer et al. ( 1 994) for some Ri versleigh sites. 
Site G of Flannery et al. ( 1 982) is included within 
System A on this table since it is described by 
those authors as lying within the Carl Creek 
Limestone. 

Late Oligocene (Archer et al., 1995; Myers & 
Archer, this volume) System A may have been a 
lime of 'icehouse conditions' with relatively low 
temperature, rainfall and biodiversity (Frakes et 
al., 1987). 

Five macropodoid species have now been iden- 
tified in System A sites. Of these, Balharoo 
gregoriensis and Wabularoo naughtoni are 
known also from System B and Bidimgamaya 
delicata occurs in sites in all 3 Systems. 
Gumardee pascuali and Galanarla tessellata are 
so far known only from System A. Unfortunately 
molar teeth preserved in the holotypes of the last 
two species are badly worn and/or damaged, re- 
ducing their usefulness for biostatigraphic analy- 
sis. However, lower molars of Galanarla 
tessellata exhibit a well-developed posterior 
cingulid linked to a postentocristid, a feature typ- 
ical of balbarines (see Cooke this volume), but 
not present among bulungamayines. 

G. tessellata is here assigned to the Balbarinac. 
As noted by Flannery et al. (1982), Gumardee 
pascuali is of comparable size to Wabularoo 
naughtoni, a common variable species within 
Riversleigh's Systems A and B. G. pascuali is 
considered to fall within the range of variation 
observed among specimens of W. naughtoni. 

Nambaroo, Wururoo {Cooke, this volume) and 
Ganawamaya, Balbaroo sp.2, Nowidgee matrix 
(Cooke, this volume) and Ganguroo bilamina 



(Cooke, this volume) all occur in System B, but 
not in System A or C. These taxa in newly dis- 
covered sites of unknown relative age may there- 
fore be suggestive of an age comparable to other 
System B sites. Occurrence of B. delicata in all 
three systems and of W. naughtoni and B. 
gregoriensis in Systems A and B suggests that 
caution is indicated before declaring any of the 
above taxa, so far found only in System B, as 
definitive indicators of that System. Balbaroo 
gregoriensis, for instance, is more derived in 
molar morphology than any Nambaroo and it 
seems likely that representatives of the laUer 
plesiomorphic balbarine and perhaps of others, 
such as Wururoo and Ganawamaya, may also 
ultimately be found in System A. If System B 
sites are correctly interpreted as early Miocene in 
age (Archer etal., 1994, 1 995), remains found in 
those sites may have accumulated during *green- 
house conditions' with high temperatures, rain- 
fall and biodiversity (Frakes et al,, 1987). 

The macropodoid fauna of System C is more 
distinctive than those of Systems A and B, con- 
taining representatives of 5 subfamilies: 4 species 
of Bulungamayinae and 1 eachof Hypsiprymno- 
dontinae, Potoroinae, Propleopinae and 
Balbarinae. Of the 8 macropodoid species known 
from System C, Bulungamaya delicata is the only 
oneoccuring in other Systems. Of the remainder. 
Wan (which includes the *Gag Site macropodine' 
of tannery, 1 989) and Ganguroo sp.2 are among 
the most highly derived bulungamayines and 
Balbaroo sp.4 is more derived in molar morphol- 
ogy than any other known balbarine species. 
Presence of these species in particular in any 
given site may suggest but not define an age 
comparable to, or perhaps younger than that of 
System C sites. Archer et al. (1994, 1995) sug- 
gested that System C is middle Miocene. If this 
is so, this interval was also characterised by 
'greenhouse conditions* (Frakes el al., 1987). 

Overall macropodid faunal composition, rather 
than presence of particular species, may provide 
a more reliable guide in assessing relative ages of 
Riversleigh sites. If Galanarla tessellata is ac- 
cepted as a balbarine and Gumardee pascuali as 
a bulungamayine, these subfamilies are repre- 
sented in System A by 5 species, roughly equally 
divided among between the two: 3 bulunga- 
mayines versus 2 balbarines, with 7 identified 
specimens from each subfamily. System A de- 
posits may thus be characterised by roughly equal 
diversity and abundance of balbarine and 
bulungamayine species. 



i^n 



MF.MOIRS WTHF-Ql'EBNSLAND MUSEUM 



The itiv'c'sUy of maCTupud*jiil spixics is great- 
est in abs>(j[ii(c tcTms in Syslcm B. Balbarincs and 
bulungamayincs arc boihpresenl. Willi balbarines 
tlominating in terms of lutmbers of sptx^ics ( 13 
versus 4), If numbers of idenlificd specimens arc 
lakcn as a crude guide to abundanec within spe- 
cies, biikmgamayincs appear to have been more 
abundant, the 3 species of lophodont bulunga- 
mayines being represenied by a lolal ul 102 spec- 
imens, ctmipared to a total of 32 balbarinc 
specimens from 12 species. System B deposits 
may thus be characterised by high species diver- 
sity of halbannes with accompanying low species 
abundance, and by a lower species divcrsiiy of 
bulungamayines bin a relatively higher abun- 
dance of members of iho&c species. 

Sublamilial diversity is greatest in System C\ 
but Potoroinac, HypsipryiViiKKloiuinitt; an»Ji Pro- 
pleopmae are each represented by single species. 
Among the lophudont ipecie^. buJungamayines 
have ^aiticd the ascendancy <ner balbarincs in 
eenms of numbers of .s-pceka% [4 versus I j at^d in 
relative abundiince ( f 6 idennried ^pe^imens ver- 
sus.^). 

Syrfem C may thu.s be ehanicieriscd by domi- 
fiiinee of lophodont bulungamayines. low inci- 
dence of balbarincs and possible hypsiprymno 
doniines. propleopines and poloruincs, although 
undescribed propleopinc remains arc known 
from sites such as Dirk's Towers which is prob- 
ably equivalent in age to system B. 

I vower absolute diversity in System A depuftits 
may result from ecMiler, drier climatic eondiiir>n.s, 
exaccrbincd by the lower nutTiher ol System A 
sites (2 1 yielding ideniiHed macnopodoid specie'^, 
compared lo the number of System B sues ( 10) 
yieUUng such species. MactT>pijdoid diversity 
wiihin System A 3S thus likely 10 have been higher 
than that currently fcmiwn. 

System B dcpasiis are suggested to Ivive accu- 
mulated in pools or shallow laiccs and System C 
deposits in deep pools, shallow pools or emergent 
.accreting surfacets, or cave oulwashes (Aaher el 
al., 1994). Depositional cnvironnxinis are thus 
more comparable for Systetns B and C . The prob- 
abilibes of accumulatii^g renu^ins of terrestrial 
mammals are also likely to be comparable for 
these systems. DilTcrenccs in overall 
macropodoid fauna! composiutm between these 
two Systems are therefore more likely to be a true 
rcllccimn of condiiions prevailitlg duritlg the 
limes of ilept.xsilion of these Systems. 

There is a striking chitnge in macropodoid fau- 
nal composiiu>n l>ciween System^ B and C. With 
the exception of a single, highly-derived lopho- 



dont. Balbaroo sp.4^ browsing btilbimnei are ap- 
parently absent faMii System C, as »s WimthnHt 
naii^htoni which persists through Sysicms A and 
B. tiulnngiwmya deikam persists only in .sites 
low in the sequence ofSysietnCdeposib. System 
B omnivorcs. .such as Nowidgve. arc ncploccd in 
System C by Betuvigio fnoyesii and Hypsipnw- 
noilon bartholomoii. Compared \o System B. 
System C macropodoid assemblages are depau- 
perate m numbers ol species and dominated by 
larger, derived, lophodont bulungamayines 
whose long premolars and general molar mor- 
phology bear strong similarities to iho.se ol* 
plesiomorphic macropodids. The System B 
macropodoid assemblages have a high species 
diversity^ particularly among balbatines. High 
faunal diversity is a eharacieristic of rainfnrcsi 
habitats, suggested by Archer et al. ( 1989) to be 
the habitat for older Riverslcigh local faunas (in- 
cluding those from System C), The decline in 
overall macropodoid diversity evident within 
System C and the dominance of larger. lopho- 
dont hulungamayine species suggests that 
rainforest habitat indicated by Archer ei al. 
(1989) may have been in decline during accumu- 
laliun of System C. That this was not a sudden 
event is mdicaled by the persistence ol the pre- 
sumably rainforest adapted B. dt^licLita into the 
lower levels ol System C al Gag Site, the occur- 
rence at this same Mte of Hvp\tprytnno(I{tn. mod- 
em representatives of which arc confined U) 
rajnturesiinnonhcm Queensland, and the occur- 
rence there of a high diversitv of possutT^ species 
(Archercial., 1991). 

Aaher el al. ( 1994. 1995) eslimaled the age ol* 
System B as early Miocene and System C as 
middle Miocene, McGowran & Li (1994) corre- 
late Lhe planktonic foraminifera record of soulii- 
ctn Australia, oceanic d"*0 levels atid sea level 
tluctuations and indicate generally warmer, wet- 
ter cUmatic conditions during lhe Oligocenc- 
carly Miocene, wiitt 3 waim iind moist climahC 
optima occurring during the Miocene. Two of 
these occur during the early Miocene iluring the 
.lanjukian and Longfordian stages respectively, 
and a double-peaked optimum occurred during 
the early miildle MiocenL*, corresponding with 
Batesfordian and Balcomliian stages, I6-I5ma.. 
b'ol lowing the latest ol these climatic optima 
there is a general and world wide decline towards 
a cooler, drier climatic regime associated with a 
'reverse greenhouse' effect. In northern Australia 
the eflects of this decline would have been exac 
erbiiled by the middle Miocene uplift of the New 
Guinea Highlands. Archer et al. ( 1989) have sug- 



BIOSTRATIGRAPHIC IMPLICATIONS OF FOSSIL KANGAROOS 



299 



gesled these newly upthrust mountains could 
have created a 'rain shadow' effect across north- 
ern Australia which may have been one of the 
most important factors causing the decline of 
central and northern Australian rainforests. The 
climatic decline following the early middle 
Miocene climatic optimum coincides well with 
the early middle Miocene estimate of Archer et 
al. (1994, 1995) for the System B/C boundary. It 
is therefore likely that the decline in macropodoid 
diversity at the System B/C boundary is a reflec- 
tion of the combined effects of the geological and 
climatic phenomena outlined above. 

A number of Riversleigh sites are of uncertain 
stratigraphic relationship. The local biostrati- 
graphic implications of the macropodoid taxa 
which have been found in some of these are 
discussed below. 

The low stratigraphic position of White Hunter 
Site and its unusual faunal assemblage make it 
uncertain whether it belongs to System A or B. 
Of its 7 identified macropodid taxa found, 5 are 
unique to the site and, of themselves, do little to 
settle the question either way. Of the remainder, 
the plesiomorphic balbarine, Nambaroo sp.8, is 
found elsewhere only at Dunsinane Site whose 
relative age is also uncertain (but see Arena, this 
volume). Nowidgee matrix, is found only within 
System B at sites in both lower and higher levels 
of the sequence. The occurrence of this species 
and the overall composition of the macropodoid 
fauna of White Hunter Site — dominated by 
plesiomorphic balbarines with 2 plesiomorphic 
bulungamayincs, suggests that the site is possibly 
a basal member of the System B sequence. The 2 
species of Nambaroo found in this site are ex- 
tremely plesiomorphic balbarines (Cooke, this 
volume) and the species oi Nowidgee found there 
are similarly plesiomorphic bulungamayincs 
(Cooke, this volume). Occurrence of such 
plesiomorphic species in the one site sugesis that 
White Hunter may be even older, perhaps belong- 
ing to System A (Creaser, this volume). The latter 
interpretation is supported by Myers & Archer 
(this volume) who report the occurrence at White 
Hunter Site of Kuterintja ngama, an ilariid con- 
specific with one in the Mammalon Hill Local 
Fauna of central Australia that is dated as 24myo 
(late Oligocene) by magnetoslratigraphy. 

The occurrence of N. sp.8 at Dunsinane Site 
complicates rather than clarifies understanding of 
this already enigmatic site in which are preserved 
plant material, insects and fossil bone. Dunsinane 
occurs in an area close to the boundary of the 
Tertiary limestone and Precambrian quartzite. 



The occurrence of this plesiomorphic balbarine 
suggests the site may be equivalent in age to 
White Hunter Site, but there are several reasons 
for caution. No other vertebrate remains support- 
ing this age determination have so far been iden- 
tified from this site. All vertebrate remains here 
are fragmented and poorly preserved and may 
well have been re-deposited after previous re- 
working. 

Bitcsantennary Site and Dirk's Towers Site are 
both intrusive deposits on the D Site Plateau, 
regarded as probably equivalent to System B 
assemblages (Archer et al., 1994). Only a single 
macropodoid, Ganguroo bilamina, has been 
identified from Bitcsantennary Site, bui this spe- 
cies is otherwise known only from System B. 
Dirk's Towers Site has 3 macropodoids, includ- 
ing Bulungamaya delicata. known from Systems 
A, B and C. Balbaroo gregoriensis is known 
from both Systems A and B. However, 
Nambaroo sp.5, is known only from System B. 
On balance, macropodoid fauna of Dirk's Towers 
Site supports a System B age. 

Archer ei al. (1994) suggested thai the faunal 
assemblage of Quantum Leap warrants its likely 
inclusion in System A. Only 3 macropodoid spe- 
cies are so far known from this site. Bulungamaya 
delicata is uninformative since it occurs in all 3 
Systems. However, 2 species of Nambaroo are 
known from the site, both species otherwise 
known from System B sites or those likely to be 
of equivalent age, e.g.. Dirk's Towers and 
Neville's Garden Sites. The macropodoid fauna 
of this site therefore suggests a closer affinity 
with System B rather than System A. 

Neville's Garden Site is a possible cave out- 
wash deposit considered to be equivalent in age 
to System B (Archer et al., 1994). Macropodoids 
of this site include the ubiquitous B. delicata. W. 
naughtoni and Balbaroo gregoriensis, Ihc latter 
two occurring in systems A and B, and 2 species 
of Nambaroo known from both upper and lower 
levels of System B. The occurrence of the latter 
species and the typical System B composition of 
the Neville's Garden macropodoid fauna support 
a System B age for this site. 

Dome Site, Jim's Carousel Site and Cleft of 
Ages Site have all been suggested to be younger 
than System C sites (Archer et al., 1994). 
Gen.Wan. sp.l (the 'Gag Site macropodine') is 
the only macropodoid so far identified from 
Dome Site. Il is known elsewhere only from Gag 
Site in System C and Encore Site of possible late 
Miocene age (Archeretal.,1 994). Gan^Mroosp. 2 
is the only macropodoid so far known from Jim's 



JOo 



MEMOIRS OF THE Ql'EENSLAND MUSEUM 



Cvousel Site- This species is also Vnnwn rrom 
Gag and lasi Mmuie Siies, low in ihe sequence of 
System C deposits, and from Hcnk\s Hollow Site 
in the higher levels of that hequent-e. 

Rolh species arc highly derived hulunga- 
mayincs which Tirsl iippcMr in Syslcni C, hui ihcre 
is no reason lu suggest thai Ihcy may nol have 

E!fsistcd beyond System C. Tticir presence in 
oine and Jim's Carousel Sites iherefory cannoi 
contirm or deny the younger age posuilaicd for 
these sites. 

The single macropodoid. Wururoo sp.2, from 
Clel'i of Ages is a plesiomorpiiic balhririne other- 
wise known only from System B. lis presence 
therefore suggests system B. This is in conflict 
with wombat teeth in this site, not known from 
any of the older Riversleigh siltis, and the *gencr- 
ally more modem* (presumably 'more derived' i 
appearance oloiherremains( Archer el af. 1989). 

Encore Site has been suggested to be younger 
still, possibly late Miocene { Aa^her el af., 1994) 
Encore has 2 highly derived bulungamayine spe- 
cies, g^n.Wan., spp. 1&2. Both species <ire known 
from Sysicn) C. hut there is no reason to suggest 
that these moderately robust, lophodonl sjxjcies 
might iK)t persist beyond the age of ihiU system. 
Their pfcscncc does not preclude the age csii- 
DiiUed for this Mte. 

MACROPODOID CORRELATES 

BETWEEN RIVERSLEIGH AND OTVIER 

AUSTRALIAN TERTI.ARY FOSSIL AREAS 

OF COMPARABLE AGE 

Afchorccal, (1 989) suggested ihw Riversleigh 
System A units may fall 'somewhere Iwtwecjn the 
Ditjinianka and Kutjumarpu and Tarkarooloo 
LFs of South Australia' a view supported by 
Wixidbumc et al. (1993 ). The latter authors main- 
tained that the base of the mamrnal-beanng se- 
quence in the Etadunna, Namba and Wipajiri 
Formations j>redaies that of ihe Riverslci^jh suc- 
cession. They noted in their < lowest) faunal zone 
A at Lake Palankarinna Kyecma mahoneyi^ 
chiimed to be the most plesiomorphic potoroid so 
far found. Forma! description of this species has 
yet to be published and its level of evolutionary 
development therefore cannot be compared with 
any of the plesiomorphic Riversleigh potoroids. 

Woodburne et al. (1993) also report 2 new 
species nl Nambaroo — species A and B, both 
occurring in faunal zone C from the Ngapakaldi 
Local Fauna of the Etadunna formation, with 
species B also present in the Ngama Local Fauna 
in faunai /one D. Both species are described as 



imine primitive than Nambaroo saltavm and N. 
tarrmyen from the Tarkarooloo Local Fauna, 
provisionally equaled with /one D of the 
Etadunna Fotmation at Lake Palankarinna. 
Archer ct al. (1989) correlated Riversleigh Sys- 
tem B Local Faunas with the Tarkarooloo and 
Kutjumarpu Local Faunas Woodburne ct al. 
( 1993) considered the Kutjumanni U>c«il Fauna 
of the Wipajiri Formation to represent the upper- 
most faunai unit in the eastern Lake Eyre Basin, 
and indicated a possible latest Oligocenc age for 
this Local Fauna. Mi morphology in Nonihoroo 
sp.8 from While Hunter Site is more plesio- 
morphic than thaiof any of the A'fmi/?^/m/> species 
in the Tark:»roo!oo Local Fauna. It retains a 
hypoconulid. has a straight or only ^^lighily 
curved paracrisiid, a short, low anterior cingulid. 
an underHleveloped precingulid and a diagonal 
poslhypocrislid on the posterior face of the 
hypolophid. Using the level of evolutionary de- 
velopment argumenl of Woodburne ei ;d (1993), 
the White Hunter assemblage would be older than 
either the Lake Tarkarooloo or Ktiljumarpu Local 
Fatinas. possibly of equivalent age to their /one 
B+C, estimated by them to be between 25.5- 
25.0myo. If White Himler Site is a member of 
System B, System .A sites at Riversleigh would 
ihercfone be older, possibly equivalent in age to 
zone A of Ihc Etadunna formation, i.e., > 
25.5myo. 

\V\xKihiunne ei al ( 1993 ) re(X)rtcd a now gcnas 
and species of macropodid (their mucropodine 
Gen. F sp. A,) from /one C of ihc Etadunna 
Fi)nniUion 

This species was described as more apo- 
morphic in Mi irigonid nK)q)holog^ than species 
of Na/nharoo. having a 'reduced protostylid*. It 
was considered l\v them to be potentially ances- 
tral to two new species ot Balbaroo from the 
Kutjumarpu Local Fauna. Given this infof?naiion 
the new genus must be a balbarine comparable lo 
Wnruron (Cooke, this volume) fn>m System B. 
The comparable levels of development of these 
genera provide furthcrevidcncc forcquaiing Sys- 
leni B sites with /one B+C of the Etadunna for- 
mation. The overall high diversity of balbarine 
species a^jxnted from the various /ones of the 
Etadunna Formation and from the Kutjumarpu 
Local Fauna (5 species of Nambaroo, 2 of 
HaiharoiK 3 of the balbarine genus P. and a spe- 
cies of a macropodine Ipossibly balbarine?| 
genus W) is, as has been noted above, typical of 
the high balbarine diversity oi System B <nt 
Riversleigh and lends lurlher suppon to a late 
Oligocenc age for Riversleigh Syslent B, its basal 



B10STRATIGRAPHIC IMPLICATIONS OF FOSSIL KANGAROOS 



m 



units bGW^ probably equivulenl in age K> ihe 
Diljiinanka and Ngapakakli Local Faunas nf the 
liladunna F(innalion. 

At the opposiic end o\' the linic scale, derived, 
)i)phodonlbulungarTmyincssuchasgcn.V\''t7«.,OL*- 
curring in Riversleigh System C and pONsibly 
younger sites, have been suggesicd elsewhere 
(Cooke, this voiume) to be likely anlecedenis of 
plesiomorphic macropodids such as HoJrttnomas 
puckrldgi from the late Miocene Alcoota Local 
Fauna. S'oungcsl Miocene sites at Rivetieigh arc 
Iherclbre probably (.>lder than liv: Alcooia Local 
Fauna. BalbaroospA has lower molar morphol- 
ogy that is more derived ihan that ol B. 
fumfteldensis fron) the late middle Miocene Bui- 
Kx:k Creek Local Fauna. B. sp.4 occurs in Ihc 
highest levels of Sysieii'» C which may llicrelV>re 
approximate ihe age of ihe Bullock Creek Local 
Fauna. 

Kangaroo fos&ils so tar t^ecovcrcd from 
Riversleigh thus suppon an age span ("or prc- 
Piiocenc deposits extending from laic Ohgoconc 
lo at least late middle Miocene. For some 
Riversleigh sites, e.g.. Dunsinane and Cleli ol 
Ages Sites, they suggest ages older than tiiose 
previously indicated by Archer et al. (10*M). 

ACKNOWLEDGF.MF.NTS 

Research grants provided from Ihe Australian 
Research Council and the University of New 
South Wales have been the primary mechanism 
for providing the research maierial evamincd in 
this study. 

Additional support Jbr the Riversleigh project 
hits come from the Naiional Fsiates Program 
grants, the Australian Cleographical Society. The 
Australian Museum. The Riversleigh .Society, 
ICI Ply Ltd, Century Zinc Limited, the Ml Isa 
Shire and private donors. 

LITERATURE CITED 

ARCHER..M. 1979. Wuhukuvo nai4,^htoni gon. ci sp. 
nov . an cnignulic kangamu (Marsupialiu) Iruin 
the middle Tertiary Carl Creek Linicsionc of 
norlhwcMcm Queensland. Results ot ihe K;iy E. 
Lctnlcy Expedilitms. pari 4. MemoirN ut llic 
QuecnsKiml Museum I'.J; 299-307, 

ARCHER, M iSc FLANNERY,T.r. I9N5. Revision of 
Iho cstinct gigamic rat kangiiaxis (Poiomiidut;' 
Marsupialia). with udeseriplion of a new Miocenc 
gCTH'S and species ynd a new Pleisioccnc species 
of ProplcopLis. Journal of Paleontology 59: 1131- 
1149 

ARCHER. M.. GODTHfiLP. H.. HAND. S J & 
MEGIRIAN, D \^m I'lfwiil munmiuls nt 



Riversleigh, nonhwcslem Queensland: prelimi- 
iiiiry overview of bioslratigraphy, correlation and 
environmental change. Australian Zoologist 
:5(2): 29-65. 

ARCffER. M , HAND. SJ & GODTHELP. H. Pr^^l 
Riversleigh. (Reed: Sydney). 

ARCHER. M. et aJ„ 1994. l.isi of the pfincipal 
Riversleigh local faunas and their tnlcmrclcd rel- 
ative ages. AbslracLs; The Riversleigli Sympo- 
sium J9'>4 (Supplement): 28-31, 

ARCHER. M.. HAND. S.J & GODTHELP. \l 1995. 
Tertiary cnvironmeninl and biolic change in Aus- 
tralia. Pp. 77-90. In Vreba. E.S.. Dcnton.G.H.. 
Panridge. T.C- & Buckle. L.H. (cd.s). Palaeoeli- 
maie and evolution, with emphasis on human 
origins (Yale University Pre.ss: New HaveiO. 

ARENA. R. 1997. The palaeoniology and gcolo^?^ ol' 
Dunsinanc Site, Riversleigh Memoirs of ihe 
Queensland Museum 41. 171-179. 

COOKE, B,N. 1992. Pnmltive macropodids from 
Riversleigh, northweslern Queensland. Al- 
chcmigal '6: 201-217. 
1997 New Miocene hulungamayinc kangaroos 
(Marsupialia. Poioroidae) Irom Riversleigh. 
norlhweslern Queensland. Menu>irs ol ihe 
Queensland Museum 41 : 269- 2S0 
1997. Two new hatbanne kangaroos and lower 
molar cvoluiiou within the subfamily Memoirs 
of the Queensland Museum 4 1 . 2S 1 -294. 

CREA>SER. P 1997. Oligu-Mioceiie .sediments of 
Riversleigh; ihe potential significance of ii>pogni- 
phv. Memoirs ot the Queensland Museum 41: 
303-3J4. 

rt.ANNERY, T.f. 1989, Phytogeny of ihc 
MacropixJoidca. a study in convergence. Pp 1-46. 
In Grigg, O. Jarman. P He Hume, I. (eds). Kanga- 
roos, wallabies and rat-kongaroos (Surrey Bcuily 
& Sons. Sydnevj. 

FLANNERY. t" ^ ARCHER, M. I9g7a. 
Hyitsipnmiuulo/i hartholontaii (Poloroidae: 
M\\rsupialia). a new species from ihe MiiKriic 
DwariKiinor Local Fauna and a reassessment of 
the phylogenclic p<-isiiion of H. moscliutuY. fp 
749 758. In Archer. M. (ed.), Possums and opos- 
sums: studies incvolution. (Surrey BeaUy & Sonjw: 
Sydney). 
I987h. Hi'tUm^Ui moyt-ii, n new and plesiomorphic 
kangaroo (Mar.supialia: Poioroidac) from 
Miocene sedimenisornorlhwcslem Queensland. 
Pp 759-7t)7. In Archer. M {ed ), Possum.s and 
opossums: studies in evolution. (Surrey Bearty & 
Sons: Sydney) 

FLANNERY. T.F. & RICH TH V I9H6. 
Macrtipodoids trom the middle Mkxcne Namba 
Fomiaiion. South Au.siralia. and ihc homology nf 
sonic dental siiuciures in kangiuoos, JfHim*il af 
P;ileonlolog>60(2):4lS-447. 

FLANNERY, TF.. ARCHFR, M. & PLANE. M. I9K2. 
Middle Miocene kangaroos (Mjcropodoidej*: 
Mlirsopiahal IVott) three localities m noiiheni Au.s- 
traJia, wiih a dcsenplion of two new siihfamiliis. 



302 MEMOIRS OF THE QUEENSLAND MUSEUM 



Bureau of Mineral Resources Journal of Austra- MYERS, T.J. & ARCHER, M. 1997. Kuterintja ngama 

lian Geology and Geophysics 7: 287-302. (Marsupialia, llariidae): a revised and extended 

FRAKES, L.A., McGOWRAN, B. & BOWLER, J. M., systematic analysis based on material from the late 

1987. Evolution of Australian environments. Pp. Oligocene of Riversleigh, northwestern Queens- 

1-16. In Dyne, G.R. & Bolton, D.W. (eds). Fauna land. Memoirs of the Queensland Museum 41: 

of Australia. Vol. 1 A: general Articles. (Australian 379-392. 

Government Publishing Service: Canberra). WOODBURNE, M.O. 1967. The Alcoota Fauna cen- 

McGOWRAN, B., & LI, Q., 1994. The Miocene oscil- t^"^' Australia: an integrated palaeontological and 

lation in southern Australia. Rec. S. Aust. Mus. geological study. Bureau of Mmeral Resources, 

27(2)- 197 212 Geology and Geophysics Bulletin 87. 

x.or^.o /kt r^ inn^ . ■ r u x.- WOODBURNE, M.O., MACFADDEN, B.J., CASE, 

MEGIRIAN, D. 1992. Interpretation of the Miocene j^^ SPRINGER, M.S., PLEDGE, N., POWER, 

Carl Creek Limestone northwestern Queensland. y^ WOODBURNE, J.M. & SPRINGER, K.B. 

Ihe Beagle. 9: 219-248. j993 ^and mammal biostratigraphy and mag- 

1994. Why the "Systems" terminology used at netostratigraphy of the Etadunna Formation (late 

Riversleigh should be abandoned. Abstracts; The Oligocene) of South Australia. Journal of Verte- 

Riversleigh Symposium, 1994: 17. brate Paleontology 14:483-515. 



OLIGOCENE-MIOCENE SEDIMENTS OF RFVERSLEIGH: 
THE POTENTIAL SIGNIFICANCE OF TOPOGR.\PHY 

PHILCREASER 

Crease r, P iO*-)? 06 30, Oligoccnc-Mioccnc sedimeni.s of Riverslcigh: the polcniial signifi- 
cance of ^opograph>^Affmo//•A•(7///If'^wfmv/i3/;rf^fM5^«f?Ml (2): 303-3 14TSSN 007 

Although faunal assemblages pmvide ihe best indication of reiaiive ages and environments 
of dcposilion in Tertiary and Quatemiiry .sediments of the Riversteigh region, geological 
evidence provides additional significant information about the prehistory of the area. Data 
prcscmcd herein on topographic heights of siies in ;ireas of horizonially-bedded sediments 
lead to an hypothesis of cyclical scdimenialion. At least 3 cycles of Oligocenc- Miocene 
sedimentation consist of 4 siagcb; 1 ) uplift and/or lowering of the waierlable: 2) erosion and 
development of a karst landscape; 3) subsidence iind/or raising of the water table, and 4) 
Nedimtmt accuraulaiion williin the karsi lerrain and in .surrounding shallow basins. 

Ptiil Creasef, 3 Paroo Place, KuhmACT26t7, Australia: received 3 February 1997. 



This paper uses data (wm Archer el al. (1989. 
1994) and Megirian (1992). palaeontology and 
preliminary mapping using phutogrammclnc 
base maps al 1:2,000, 1 : 15,000 and 1:20,000 lu 
inierprei stratigraphy and palaeogeography of 
Oligoccne-Miocene sediments at Riversleigh. I 
focus on Oligocene-Miocene sediments on D Site 
and Gag Plateaus dated primarily through 
biocorrelation with magnelostratigraphically 
dated deposits in South Australia (Woodbumc ci 
al.. 1994). 

Archer el al. (1989) and Megirian (1992) fo- 
cussed on D Site and Gag Plateaux, particularly 
Codlhelp's Hill and Hal's Hill areas and the 
northern Gag Plateau. I concentrate herein on the 
northern D Site Plateau and the southern Gag 
Plateau. I also builds on Archer el al.'s (1989") 
observation on the Gag Plateau that it is possible 
to correlate widely separated exposures of flat- 
lying sediments. In areas where the sediments are 
faulled or areas on Ihe margins of microbasins 
where the beds are dipping, correlalion is limited. 

All sites have been plotted onto ba.sc maps 
lodged with the Vertebrate Palaecmtology Labo- 
ratory, University of New South Wales. Queens- 
land Museum and the Queensland National Parks 
and Wildlife Service. I provide new infomialion 
on relative U >pographic heights which contributes 
to geological understanding of the region. 

1 consider geographic sections of each plateau 
and look al the geology of each section (Fig. 1) 
noting the range of sediment types, presumed age 
of the sediments based on palaeoniological evi- 
dence and. where appropriate, topographic 
heights. 

The other area considered is the 'Mesas'. Iso- 
laied erosiunul remnants of Tertiary sediment U 



of the Riversleigh/Lawn Hill road. A number of 

these sites appear similar in lilhology and stratig- 
raphy to sites in the northern section of the D Site 
Plateau. 

1 recognise 3 sedimentary sequences (Verdon 
Creek. Godlhelp's Hill and Gag Plateau). Tlw 
Verdon Creek sequence is best represented in the- 
northern section of D Site Plateau and consists uf 
mainly System A sites ( Archer et al., 1989). Tl>c 
Godthclp's Hill sequence occurs in the central 
section of D Site Plateau with System B sites. The 
Gag Plateau sequence is best represented in the 
northern section of Gag Plateau and contain.s 
mainly System C sites but may also include Sys- 
tem A sites. Recent fieldwork has indicated a Ittck 
oi unilbrmily and continuity of the basal scdi- 
itionis in Ihe northern section of Gag Plateau. 

D SITE PLATEAU 

The northern section includes Neville's Gar- 
den/Burnt Oriering area and the major gully sy.s- 
tern to the south of this area with sites such as 
Quanturu Leap. Gillespic\s Gully and MIM. li 
also includes sites on the caslcin edge of the 
plateau (LSO and Dirk's Towers) as well as the 
sites on the western edge (BIB). The southern 
boundary is at or about Syp's Siberia Site to the 
north of Godthelp's Hill. 

The central section includes Godihelp's Hill, 
Hal's Hill and other sites in the valley to the south 
cast oflhese hills including White Hunter, ABKS, 
Sticky Beak and W^^yne's Wok. 

The southern section includes sites south of 
Hal's Hill commencing with the Biggies Flics 
Again Site, SM and TOTE Sites and Bone Reef, 
Jeaneite's Amphitheatre and Chinatown Sites. 



304 



MEMOIRS OF THE QUEENSLAND MUSEUM 



D SITE PLATEAU 



GAG PLATEAU 



North Section Central Section 



North Section South Section 



VERDON CREEK 
SEQUENCE 



GAG PLATEAU 
SEQUENCE 



System C sediments 



Uplift and Erosion 



Sediments overlying 
D Site level 



Sediments overlying 
D Site level 
equivalents 



GODTHELP' SKILL 

SEQUENCE 
(System B sediments) 



D Site level 

(System A) with 

cave and related 

deposits 



System A 
sediments 



D Site level equivalents 
(System A) 



System A sediments 
(inc. White Hunter) 



System C sediments 



Basal sediments. 
? System A 



Sequence of cave 

deposits, fissure fills 

and 

stratigiaphic/shallow 

pool deposits. 

There is at least one 

period of uplift and 

erosioa 

Sediments range in age 

from ?System A to 

System C 



Overlying calcarenites 

(Possible System A 

sedin>ents) 



Basal sediments with 
White Hunter equivaleni 



FIG. I. Generalised Riversleigh stratigraphy based on geological observations, biocorrclations and topographic 
heights. 



NORTHERN SECTION (VERDON CREEK 
SEQUENCE).This sequence is best seen in the 
area of Bitesanlennary, Burnt Ofiering and 
Nevilie'sGarden Sites where it consists ofa basal 
conglomerate, overlain by areniles and calcaren- 
ites, up to 20rn thick. A 3m homogeneous lime- 



stone, the D Site Limestone, overlies these sedi- 
ments. Cutting into and lying on the D Site Lime- 
stone are a series of cave deposits and possibly 
related tufa deposits. A further series of calcaren- 
ites are the highest units of this sequence. 
The basal conglomerate consist of a series of 



OLIGO-MIOCENE SEDIMENTS OF RIVERSLEIGH 



305 



180m 



175m 



172m 



170m 



165m 



162m 



155m 



152m 



Black Coffee (??Possible System C) 



D Site level (Damaged Digit etc) 
also Bitesantemiaiy, Neville's Garden 



Burnt Offering (BO), Upper BO, VIP, Dirk's 
Towers, Judith Horiz'is, Gillespie's Gully 



Quantum Leap, Steph's Small Reward, MIM 



?LSO 



IT, PA 



Low Lion, Phil's Fangs, KJO 



Carbonate cemented chert conglomerate 



I Judy's Jumping Joint i 

! (??SystemB) | 

I Equivalent D Site ] 

! level* ' 

[ Wayne's Wok" | 

! Greaser's Ramparts ' 

I?Sticky Beaic, BoFe's 1 

[Bonanza \ 

r , 

I I 

' I 

I ?White Hunter I 

I 1 

I I 

\ THiatus I 

• I 

' I 



FIG. 2. Verdon Creek Sequence, System A with possible Systems B and C, 'type' section in northern section of 
D Site Plateau . Heights are in metres above sea level. Dashed boxes indicate relative positions of sites from 
the Hal's Hill area in the central section. 



massive or normally graded, matrix-supported 
breccias and conglomerates and clast-supported 
cobble and pebble conglomerates (Megirian, 
1992). Archer et al. (1989) questioned whether 
this conglomerate is contemporaneous through- 
out the region, suggesting that it could represent 
different, non-contemporaneous cycles of local 
weathering. The thickness of conglomerate var- 
ies considerably throughout the D Site Plateau. 

There are several fossiliferous levels in the 
overlying arenites and calcarenites (Fig. 2) dom- 
inated by turtles, crocodiles, large birds and rare 
marsupials (usually diprotodontids). 



The lowest level at the northern end of this 
section includes Low Lion and Phil's Fang Sites. 
A distinctive 25m wide fossiliferous horizon at 
the Low Lion level is at the same level as KJO 
Site. The level is dominated by large bone frag- 
ments similar in colour and preservation to fossils 
from Low Lion Site. 

Above this level at the northern extremity of the 
Plateau are the IT and PA Sites which yielded jaw 
fragments of Yalkaparidon. These two adjacent 
sites, which contain small terrestrial assem- 
blages, are the only ones known from the lower 
part of the sequence in this area. Although there 



306 



MEMOIRS OF THE QUEENSLAND MUSEUM 



are other fossiliferous assemblages at about this 
level, they tend to consist of well- worn fragments 
of aquatic vertebrates. 

The next fossiliferous level includes LSO Site. 
Above this are sites in the major gully in the 
northern part of this section including Quantum 
Leap, MIM and Sleph's Small Reward Sites. 
Cooke (1997) considered the Quantum Leap Site 
kangaroos to be most similar to others from Sys- 
tem A or B assemblages; its stratigraphic position 
and sedimentology suggest that it is a System A 
assemblage. 

Above these sites but below the D Site Lime- 
stone level, is a higher more widespread fossilif- 
erous level which includes Burnt Offering, Upper 
Burnt Offering, VIP, Judith Horizontalis, Punky 
Brewster, Gillespie's Gully and Dirk's Towers 
Sites. These appear to be slratigraphically con- 
trolled and do not represent a later incised de- 
posit. While they are at the same topographic 
level and appear to be horizontally bedded, they 
may be of different ages. 

Black ( 1 997a) suggested that Burnt Offering is 
a System A site. Cooke ( 1 997) suggested it might 
be a System B site; the macropodid fauna indi- 
cates that it could be either System A or B. Black 
{i997a) considered that Upper Burnt Offering is 
a System B site but Neohelos n. sp. 1 is only found 
in System A deposits. Black (1997a) considered 
VIP to be a System A site on the basis of a 
plesiomorphic zygomaturine. As yet, there is in- 
sufficient data from the Judith Horizontalis, 
Punky Brewster and Gillespie's Gully Sites to 
allocate these and no clear evidence from Dirk's 
Towers Site as to whether it is System A or B. 

D Site Limestone is a distinctive marker bed 
that outcrops over much of the D Site Plateau and 
is characterised by a fossil assemblage of mainly 
large vertebrates dominated by mekosuchine 
crocodiles, dromomithids and diprotodontoids. 
However, the stratigraphy of D Site (Tedford, 
1 967) and of the ridge to the north of D Site, are 
less readily interpreted because of extensive scree 
slopes. 

Archer el al. (1989) equated the D Site Lime- 
stone and its fossil assemblages with their System 
A. However, given further research since the mid 
1980s and recognition of several fossiliferous 
levels well below this marker bed, it is recom- 
mended here that System A be expanded in con- 
cept to include all of the lower sediments of the 
Verdon Creek sequence. 

A complex series of cave and possibly also tufa 
deposits have been etched into the D Site Lime- 
stone. This suggests a period of uplift or lowering 



of the water table following deposition of the D 
Site Limestone, followed by karst weathering of 
the limestone to form caves and other sedi- 
ment/fossil traps, and then infilling of these 
microbasins. The best known cave sites are 
Microsile and Biiesanlcnnary Site with the best 
example of a tufa deposit being Neville's Garden 
Site with sediment and fossils accumulating at 
and beyond the entrance to a cave. However, 
Neville's Garden Site may represent a strat- 
igraphically controlled site which can be corre- 
lated with Dirk's Towers or Judith Horizontalis 
Sites from this section or possibly Wayne's Wok 
and Greaser's Ramparts Sites from the central 
section of this Plateau, Black (1997) and Cooke 
(1997) assign this site to System B on its 
diprotodontoids and macropodoids respectively. 

A thin series of calcareniles overlies the D Site 
Limestone and the cave and tufa deposits. How- 
ever, fossils from these sediments are not com- 
mon and only one site in this section. Black 
Coffee Site (above Gillespie's Gully), has been 
sampled. Us faunal assemblage has yet to be 
analysed in detail. 

Topographically, the base of this sequence at 
the northern end of the Plateau is at 152m with 
the lowest fossiliferous level at about 1 55 m (Low 
Lion and KJO Sites). The base of the D Site 
Limestone is at 1 75m. The highest point on the D 
Site Plateau is at 202.7 m. Above Neville's Gar- 
den Site, the highest point is 192m which would 
give a thickness of at least 40m. 

CENTRAL SECTION .The central section con- 
tains the discreet, richly-fossilifcrous Godthelp's 
Hill sequence which is separated both 
stratigraphically and topographically from other 
sediments in this section which are similar in 
lithology and stratigraphy to the Verdon Creek 
sequence. The Godthelp's Hill sequence may be 
the equivalent of the cave and tufa deposits of the 
Verdon Creek sequence. The other sediments 
can probably be equated to the other Verdon 
Creek sequence sediments. 

GODTHELP'S HILL SEQUENCE. Because 
they arc separated, possibly due to faulting 
(Megirian, 1 992), from the main sequence it is not 
clear whether the tufa deposits on Godthclp' s Hill 
are the equivalent of the cave and related tufa 
deposits of the Verdon Creek sequence. Although 
faunal assemblages indicate a similar age, the 
Godthelp's Hill sediments, which have been re- 
garded as System B (Archer et al., 1989), are 
regarded as a distinct sequence (Fig. 3). The 



OLIGO-MIOCENE SEDIMENTS OF RIVERSLEIGH 



307 



GODTHELP'S HILL SEQUENCE 



VERDON CREEK SEQUENCE 



Highest 



Mid/High 



Mid 



View Delightful 

Panorama 

Ten Bags 

Boid 

Boid Site East 



Helicopter 
Mike's Potato Patch 



CAVE DEPOSITS 

Bitesantennaiy 
Microsile 

CAVE/POOL DEPOSITS 

Neville's Garden 



All occur at the D Site Level 



Upper 

CS 

Inabeyance 

Mike's Menagerie 



Mid/Low 



RSO 

RV 

Outasites 

Souvenir 



Others 



G Spot, DDDD, Victor's Vacuum, Paul Willis, Dredge's Ledge 



FIG. 3. Godthelp's Hill Sequence, System B with possible System B equivalents from the Verdon Creek 

Sequence. 



308 



MEMOIRS OF THE QUEENSLAND MUSEUM 



200m 



198m 



I97m 



196m 



194m 



192m 



188m 



179m 



Jaw Junction 



Henk's Hollow, Bob's Boulders, Grimes Site 



No Name, Arthur's Seat, Golden Steph, Phalanger 



Crusty Meat Pie, Skull, Neville's Riches, 
Fireside Favourites, H?ny's Hun^ 



Flowstone Level: Bat sites: Bat Smear, Gotham City (E & 

N), First Drop, Sticky Wicket. Other sites at or about this 

level: Sue's Diprotodontid, GOH, Bird Bone, Group, Main, 

Quentin's Quany, Courtney's Cache, Two Trees 



Ringtail, Melody's Maze 



Gag, LD'94, Jim's Jaw, Last Minute 



Don Camillo 



FIG. 4. Gag Plateau Sequence, System C, 'type' section in northern section of Gag Plateau based at Don Camillo 
Site. Heights are in metres above sea level. Sites to be plotted: Archie's Absence, Archie's Parlour, Bemie's 
Bedford, Kangaroo Jaw, Lockwood's Link, and Bruty and the Beast. 



OLIGO-MiOCENE SEDIMENTS OFRIVERSLEIGH 



309 



thickness of ihe sediments on Godthelp's Hill has 
been estimated to be 7m (Archer et al., 1989) to 
1 2m (Megirian, 1 992). Detailed photogrammetry 
of the area indicates that the estimate of 12m is 
more accurate. 

OTHER SEDIMENTS IN THIS SECTION. 

These sediments are around Hal's Hill to the 
south of Godthelp's Hill. Hiatus Site, the lowest 
in this area, is just above Precambrian sediments 
on the northern side of Hal's Hill. Although there 
is some doubt about its position because of fault- 
ing, it appears to be a base level. Black (1997a) 
notes that Silvabestius michaelbirti from Hiatus 
South Site is the most plesiomorphic zygomatur- 
ine known. Stratigraphic and topographic posi- 
tion and faunal assemblage suggest that White 
Hunter Site on the south side of Hal's Hill is very 
low in the sequence. However, its lithology dif- 
fers Irom that of Hiatus Site and the basal sedi- 
ments in the northern section of the Plateau. 
Black (1997a) indicates that Hiatus South Site 
belongs to System A. Cooke (1997) considers it 
a System ?A/B Site because several macropodoid 
species from White Hunter Site are also found in 
other undoubted System A and B sediments. 
However, there are also 5 unique macropodoid 
species from this site. Myers & Archer (1997) 
indicate White Hunter Site is the only one at 
Riversleigh that contains ilariids. Taken together, 
these suggest that White Hunter either represents 
adistinct interval of time or, if contemporaneous, 
a different ecosystem. I suggest that White 
Hunter Site is a basal System A deposit. 

There are no distinctive fossiliferous levels im- 
mediately above either Hiatus or White Hunter 
Sites. The next site up section may be Sticky Beak 
Site which is a lower level than the D Site Lime- 
stone equivalent, approximately at the same level 
as Boles' Bonanza Site. Black (1997a) suggests 
that Sticky Beak Site belongs to System A. 

The next level up is immediately below the D 
Site Limestone level equivalent. In this section, 
Wayne's Wok and Greaser's Ramparts Sites are 
at this level and may be correlated with Dirk's 
Towers and Judith Horizontalis Sites from the 
northern section. Both Black (1997a) and Cooke 
(1997) consider Wayne's Wok Site to be low in 
System B. However, this site contains a number 
of species that are found in Systems A and B. The 
age of Greaser's Ramparts Site is also unclear and 
more palaeontological information is needed. 
Black (pers. comm.) recognised a phascolarctid 
from this site, that is the most primitive from the 
Australian Tertiary. 



In this section, the only D Site Limestone 

equivalent site is Neville's Pancake Site (with a 
plesiomorphic meiolaniid turtle; Gaffney et al., 
1 992). Fig Tree Site, with the most plesiomorphic 
zygomaturine Nimbadon\ Hand et al., 1993 is 
straligraphically below Neville's Pancake Site. 
Both sites are NE of Hal's Hill. 

Above the D Site Limestone level, in the over- 
lying calcarenites, only Judy's Jumping Joint site 
has been sampled. This site is a localised con- 
glomerate found on the crest of Hal's Hill and 
belongs to System B. However, because the rela- 
tionships are not clear, it is not possible to deter- 
mine at present whether these calcarenites 
represent System A or B. 

SOUTHERN SECTION. There has been rela- 
tively limited exploration in this section of the 
Plateau with preliminary fieldwork indicating a 
thin series of sediments. Sites such as Bone Reef 
and Jeanelte's Amphitheatre are as yet largely 
unassessed. They are dominated by large animals 
more or less of the same kind (but far more 
abundant) that characterise the D Site Limestone 
at, for example. Site D. Black (1997a) considers 
these two sites System A deposits. Immediately 
below these are fossiliferous sediments. Two 
other sites collected from this region are China- 
town, which produced a System B assemblage, 
and a possible cave deposit with a rich bat fauna. 
SM and TOTE Sites, in the northern part of this 
section, are dominated by large vertebrates. 

GAG PLATEAU 

The northern section includes the vast range of 
sites at the northern end of the Plateau including 
Golden Steph, GOH, LD94 and First Drop. The 
central .section is relatively barren apart from 
Wang Site and this relative lack of sites is its 
defining feature. The southern section includes 
AL90, COA, Dunsinane, Dome, JC, Encore and 
others. The northern boundary of this rich south- 
em section is at Peter the Pilot Site. 

NORTHERN SECTION (GAG PLATEAU SE- 
QUENCE). Based on a composite section starting 
with Don Camillo Site at the base (near the north- 
ern point of this section), up through Gag Site to 
Jaw Junction Site at the top, together with equiv- 
alent sites at the appropriate levels at the northern 
end of this section, the Gag Plateau sequence was 
considered (Archer et al., 1989) to consist of 
fossiliferous basal sediments overlain by 
calcarenites and a series of tufa and 'deep water 



110 



MEMOIRS OF THE QLEENSLAND MUSEUM 



poor iltrposiis which, in soiiw ca&cs, coniain di- 
verse faunal assemblages. However, this se- 
quence contains a complex vanely of basal 
scdimenis. At Don Camillo sue. which was con- 
sidered ihe straiigraphic cquivalcni of the D Site 
Limestone, significantly diflerenl iithologies are 
present. At the eastern end of this section is a 
rossilifen)uscongloineraie and ai the western end 
a vertical sequence of richly lossiliferous sedi- 
ments. Overlying the Prccambiian is a thin se- 
quence of arenaceous sediments overlain by 
fossiliforous calcareniies which in turn arc over- 
lain by (?)wcaihercd lateritic sediments. These 
arc overlain by the 'lypical' calcareniies which in 
some pUures have rich vertebrate assemblages. 
However, noneofihese sediments are apparently 
continuous across the nonhcrn section of the Gag 
Plateau. Some of these basal sediments may be 
lateral equivalents of System A sediments in Ihe 
Verdon Creek sequence 

While the basal sediments vary considerably, 
there is apparently more uniformity higher m Ihe 
sequence including the tufaand 'deep water poof 
deposits of Archer et al. ( 1 98^). The lufa deposits 
Such as Gag, Henk's Hollow and Golden Sleph 
Sites, arc dominated by ]ciTcsiriai faunas. In con- 
trast, ihe 'deep wuier pool' deposits, such its 
Crasty Meat Pie. Qu^^nlin s Quiury. Bi>b's Boul- 
ders and Ringtail Sites arc dominated by aquatic 
faunas. 

Don Camillo Site is at approximately 179m 
with the highest point al 2fJlni (Jaw Junction 
Site), giving a ftiai;imum thickness of 22m as- 
suming the beds arc hori/.ontal. Gag, Last Minute 
and LD'y4 Sites are all al lS8m with the erosional 
break recognised by Archer et al. (1989) and 
Mcgirian (1992) at 194m. The only sites with 
signilkanl bat accumulations (Gotham City, 
Sticky Wicket, fiat Smear and Tirst Drop Sites) 
arc found at this level (Fig. 4j. 

CENTRAL SECTION. Only Wang Site is known 
from this section, with no clear 'dividing line* 
between the noriherii and soulhern sections. 
However. sedin>enLs in the southern section differ 
in the type and extent ul their lithologies and 
faunas, 

SOUTHERN SECTION. The straiigraphy of this 
section is complex. Unlike the northern section of 
the D Site Plateau or the upper northern section 
of the Gag Plareau, the series and types of sedi- 
mcnis m the southern section are appatently not 
limited in lateral extent, often significanlly differ- 
ent in age and dn not appear U) he hori/onlally 



bedded. This makes it p<?inicularly difYlcuIt io 

coiTclatc this section with others (Fig. 5). Pal- 
aeonlologicaJ evidence from this section allows 
some correlations. 

The oldest recognised sediments are at Dunsin- 
ime Sile (and equivalents) with plant and animal 
fossils. These sediments arc tiveriam by less los- 
siliferous calcareniies into which are incised 
richly fossiliferous cave and fissure fill deposits. 

Dunsinanc, Sue's Rocky Road. Custard Tart 
and Bcrnie's Cooking Pol Sites may represent ihe 
oldest from the Gag Plateau, possibly System A, 
on Ihe basis of correhilion of mammals with 
W^hite Hunter Siie (Arena, 1997; Cooke, 1997). 

Fossils are nut common in the overlying 
calcareniies at Anna's Horribilis, Two Gloves, 
Anton's Pi\ic, Arachnid Ridge and DonU Ask 
Me Sites, all of which have yel lu be studied in 
detail but are probably System A sites. Faunas 
and lithologies indicate cave deposits at Dome, 
AL'90, Peter the Pilot. Crcaser's Crouch and 
Angela's Bat Pale Sites, Fissure till deposits such 
as COA and Keith's Chocky Block Sites arc 
easily recognisable because of their lithologies. 
Jim's Carousel and Encore Sites could represent 
lufa deposits incised into a pre-existing Tertiary 
limestone. 

While the lithologies and environment of depo- 
sition oi' many oi" these sites are similar, it is clear 
thai ihey represent a wide range of ages- Black 
1 1997a) suggests that AL*90, Jim's Carousel and 
Dome Sites may all be System C deposits. Black 
( 1 997a) suggests^ thai COA Site may be a System 
A Stic, but Cooke (1997) suggests it is either 
System A or B. A. Gillespie (pers. comm.) con- 
siders it to be System B. Bncore Sile despite being 
lilhologically very similar lo v>iher sites in the 
area, is early late Miocene, System C fArcher cL 
al.. 1994; Black. 1997b). 



liNVIRONMENTS OF DEPOSITION AND 
PALAF.OC.EOGRAPHY 

The Tcrtiairy lin)csione dcjitisits of the Gregory 
River area arc freshwater fluvio-lacuslrine depos- 
its. Archer el al. (I9S9) recognised a complex 
scries of lacu.sirine, alluvial, travertine and cave 
deposits while Megirian (1992) has documented 
alluvial, tufa and karst fades. 

I agree with these views antl proix>sc thai a 
cycle of .sedimentation/eitjsion thai involves: I, 
uplift and or lowering of the water table; 2, ero- 
sion and development of a karSL landscape; 3, 



OLIGO-MIOCENE SEDIMENTS OF RIVERSLEIGH 



m 



Central Section 



Wang Site 



Southern Section 



Cave Deposits 



Fissure fills 



Stratigraphic/Shallow Pool Deposits 



Peter the Pilot, 
Greaser's Crouch (CC) 

Upper CC 

Nicole*s Boulders W 

AL'90 

Captain Androgen 

Jolly Roger 

Bernards Belted Belfiy 

Angela's Bat Pate 

BatEeik 
Anne's Bat Room 

Dome (N & S) 
Jeanette's Bat Stuff 



COA 
Keith's Chockv Block 



Arachnid Ridge* 

Nicole's Boulders 

JC Sites 

Not JC8 

Anna's Horribilis* 

Two Gloves* 

Encore 

Angela's Sinkhole 

Don't Ask Me* 

Anton's Pixie* 



Dunsinanc, Custard Tart, Sue's Rocky Road, 

Bemie's Cooking Pot 

Equivalent to White Hunter Site on D Site 

Plateau 



FIG. 5- Gag Plateau Sequence, ceniral and southern sections. The age of these sites is iinclear(apart perhaps t'rom 
Dunsinane Site and its equivalents) although there is evidence that there is a range of different ages represented. 
Deposits marked * appear to represent faunas from the caicareniies which overlie the Dunsmane Site and its 
equivaleni.s. 



subsidence and or raising of the water table; 4, 
sediment accumulation within the karst terrane 
and surrounding basins. This 4-slage cycle oc- 
curred at least 3 limes during Oligo-Miocenc time 
at Rivcrsleigh. Each new cycle may have been 
initiated by mmor tectonic activity that may have 
been responsible for changes in the 
hydrogcologic system. Megirian (1992) noted 
faulting on Godthclp's Hill which may have been 
responsible for deposition of the basal conglom- 
erates as a debris flow. 

Following initial observation by B. Cooke that 
Tertiary limestone on the mesas not uncommonly 
rest directly on Precambrian quarlzite, M. Archer 



demonstrated that there did not appear to be any- 
where on either the D Site Plateau or the Gag 
Plateau where Tertiary sediments directly over- 
lay the Cambrian limestones. In a number of 
places, however. Tertiary sediments can be found 
adjacent to Cambrian limestones (e.g. Microsite 
which is topographically situated between a high 
of Thornionia Limestone to the wesi and D Site 
Limestone to the east) which suggests that karst 
topography of Cambrian limestones may have 
controlled sedimentation palterns in the Tertiary. 
1 suggest that the basid iu-enaceous sediments 
and the ovcrlymg caicareniies were dcpt)siled in 
an alluvial fan/braided stream envirimmeni- 



312 



MEMOIRS OF THE QUEENSLAND MUSEUM 



C3ivcD the oulcmp paLicm. h is likdy Lhat tbc 
Mfe;iiiis (Itjwod in a tionheuxierly dirccTion. A> 
Mcgirian l !992) has noted, fossils arc not com- 
nvon in this lype of 4?nvin:>nineni. However, in 
Mime paas of this environment, possibly in 
swump areas or stagnuni water away from Ihe 
forest, assemblages of large aquatic and Icnrs- 
liial animal fossils accumulalcd. in other ureus 
freshwater limestone tufa pools developed in or 
at the edge of the rainforest hut these are rela- 
lively rare. The level eonlaining rich fatinas fron) 
sites such as Dirk's Towers. Greaser's Ranipaits. 
Judith Hori/onialis and Bumi OlTering Sites, is a 
notable example. These sites arc often found in 
giiilies today because they arc more easily weath- 
ered than the suirounding more resistant sedi- 
ments. There does not appear to beany breaks in 
this part of the set|uence and it is suggested that 
this sedimentation continued until (?)iectonic ac- 
livily or a raised water table led to the empond- 
meni of larger lacustrine bodies of walerin which 
accumulated the sedimenis (e g.. die DSiie Lime- 
stone) exposed at localities such as D Site. These 
sediments are fairly uniform in lithology upart 
from Ihe northeaslcm corner of the D Site Plateau 
(near the Lawn Hill roail and Vcrdon Creek) 
wlverc there is a high percentage ol quartz grains 
sug^es^ing a Precambrian source to the nooh- 
casl. Large vertebrates, including crocodiles and 
turtles, tire common in this limestone which is 
c^msidcrcd ui represent an open lake or swamp, 
at sotne- distance from the rainforest. This would 
cxjilain the scarcity of tcitcstrial faunas espc- 
ciaJly when compared to the shallow pool tufa 
(tcpr^iis (Archer ci at., 19H9)- 

Renewed tectonic uplift and/or a lowering of 
the water table resulted in renewed devclopmcnl 
uf a kiusi tandscai>e in the Tertiary as well as 
Cambrian linKstoncs. This enabled formation of 
cavesand Iheii subsequent filling with fossilifer- 
ous deposits fc.g. Micawite and Bitesantennary 
Site), ds wel I as deposits formed at cave entrances 
(e.g. Neville's Garden Site where fragments of 
speleothems (straws) and in sUu triivcrtine rills 
and small stalagmites have been foutid), 

Godthelp's Hill sequence is regarded as a sep- 
arate unit, which may ite equivalent in part to 
these cave deposits. Lithology of the sediments 
and the fossils from sites on Godthelp's Hill 
suggest tufa deposits ihiU iiccumulated over a 
period oi time. 

The uppermu-si scries of calcoxcnitcs in the 
Verdon Creek sequence sug^e^l a return to the 
alluvial braided vinfara facies. possibly following 
erosion of the karsi landscape. These s^dinfients 



may be the lateral equivalents of Ihe System C 

sediments at ihe northern end (»f the Gag Plateau, 
or alternatively they may be part oi' System A. 
System B faunal assemblage from the conglom- 
erate at Judy's Jumping Joint SiiedLK^s not enable 
iiiiy definite conclusions lo be drawn because the 
relationship of this Site to the surrounding sedi- 
ments is unclear. 

The basal sediments at the northern end of the 
Gag Plateau vary from fossilifcrous conglomer- 
ates to arenaceous sediments and weathered lai- 
eritic sediments. These sediments are overlain by 
culcarenites that suggest an alluvial braided 
stream facies. and a fossil-rich tufa and deep pcx)l 
(aquatic) deposits. Like similar deposits in the 
Verdon Creek sequence, the tufa deposits appear 
to have accumulated in and around shallow ter- 
restrial pools. There is no cleat pattern evident in 
the distribution of these two types of shallow 
water tula und deep water deposits, Thetts is also 
a significant eru^iimul break in the sequence that 
can be rxicognised in the field by flowstone and 
travertine deposits. Palaeonlological evidence 
also indicates a significant break between the 
upper and lower parts of the sequence in this arra 
and it may be no coincidence lhat the only sites 
in this area which contain significant bat accumu- 
lations are at tins level. This suggests another 
cycle of uplift and or lowering of the water lablev 
cave developmenis cuve fill, and erosion before 
sedimentation recommenced. 

In contrasi. there are apparently no similar se- 
quences of shallow water tufa or deep pool de- 
posits at the southern end of the Gag PJalcau- 
Isolatcd sites such as Jim's Carousel Site, how- 
ever, may be of this type. It is not clear if the 
sediments at the northern end are equivalent to 
the main sequence of sediments at the southern 
end of ihe Plateau, 

Mammals from Dunsinanc and related sites at 
the southern end ol the Plateau, which appeal to 
represent basal sediments, suggest correlation 
with While Hunter Site, a probable System A 
assemblage (Arena, 1^)96. 1997). These sedi- 
ments arc overlain by a scries of cakareniics 
which were probably deposited in an aJluvial 
j'an/braidcd stream cnvironmcni. Associated with 
the caicarenites are cave and fissure fill deposits 
which have been incised into the earlier Tertiary 
limestones indicating that a karst landscape had 
already been formed. The age of these cave and 
fissure fill deposits appears to range considerably 
with .some (e.g., Encore Site) being the youngest 
Oligo-Miocene sediments in the region. 



OLlGO-MIOCENE SEDIMENTS OF RIVERSLEIGH 



313 



PALAEOENVlRONMEN'i 



ACKNOWLEDGEMENTS 



On ihc basis of (he lussil fauna^i. Archer ei al. 
(1989. 1994) suggested ihat rainforest covered 
the region al least during Ihc early to middle 
Miocene. Archer (pcrs. conirn.. 19%) suggests 
that there is much less f.ninal evidence for raint"t>r- 
est being ubiquitous during the late Oligocenc. 
Other researchers support this view although 
there seems to be evidence Ihiil this rainfonesl was 
unlike an> found in Ausiialia today and thai the 
rainlbresls of New Caledonia or mid-montatic 
Papuii Now Guinea may be more smiilar. Boles 
( 1997) and While (1997) suggest that ihefc were 
jiomc patches oi open forest. Mcgirian (1992) 
suggested Ihc rainforest was a refugium confined 
to the proximity ol perennial, .spring-led streams. 
He concluded thai the palaeoclm»;ne was rela 
tivcly dry, perhaps semi-arid, despite the fact thai 
Ibc sediments were considered to be charactcris- 
Ijc nf humid alluvial fans. Archer et al. (1995) 
refuted this suggestion on faunal evidence. 
Creaser (1977) has also showed ihai ahhoiigh 
calclilhilcs, terrigenous clastic rocks in which 
carbonate fragments dtiminale, form mainly in 
alluvial fans in arid/scmi-arid andglacial/pcrigla- 
cial environnienls. they are also forming today on 
the Huon Terraces in Papua New Guinea where 
tcctonism and climate both appear to influence 
Ihe accumulation of the caklithilcs. The Huon 
Terraces arc tn a rainforest environment with 
2(X)0-2500mm of rain per annum. The rainfall has 
a marked peak in December to Februiiry, with the 
nearby mountain ranges forming a rainshadow, 
and a pronounced dry season al other times of the 
year. While th^rc are differences between the 
Huon TciTaccs and the Gregory River region, it 
is possible for a species-rich rairU'uresI lo exist »n 
an area of tectonic activity and produce the full 
range of sediments cvideni in the Gregory River 
basin. Although there is no evidence for seitson- 
aliiy in the early to middle Miocene sediments of 
Rjvcrsleigh, growih nngs in wood fragmtrnis 
(Wothofaiitts sp.) from Dunsinane Siic suggest 
seasonalily (Jane O'Bnen. pers. comm.) or at 
least episodic changes in growth rates. Unfortu- 
nately, the age and relative stratigraphic position 
of these fragiuenls is in doubt (Arena. 1997). 
They could be cither late Oligocenc or laic 
Miocene, both icehouse intervals (Frakcs & 
Macgowran. 1987) when rainforest is less likely 
to have characterised the regii^n. The plants oi 
Dunsinaiie Site may have grown on the edge of a 
forcM clearing, sum^arvding the Dunsinane b^-idy 
of water 



Many of the ideas and thoughts in this paperare 
the result of the observations and fieldwork of 
others, especially fellow wt)fkers at Riverslcigh 
and in particular, Michael Archer, Henk 
Godti^elp and Su/anne Hiind. Vital support for 
research al Rivcrsieigh has come from the Aus- 
tralian Research Grant Scheme; Ihe National Es- 
tate Grants Scheme (Queensland): Ihc University 
of New South Wales; the Commonwealth De- 
partment of Environment, Sports and Territories: 
the Queensland National Parks iind Wildlife Ser- 
vice; the Commonwealth World Heritage Unit; 
ICI Australia Ply Lid; the Australian Ideographic 
Society; the Queensland Museum; the Australian 
Museum; the Royal Zoological Society of New 
South Wales, the Linncan S<iciety of New Souih 
Wales; Ccniur>' Zinc Pty Ltd; Mount Isa MiiR'i 
F*iy Ltd; Surrey Beauty & Sons Ply Ltd; Ihe 
Riverslcigh StK'icty Inc.; and private supporters 
including Elaine cWk. Margaret Beavis, Martin 
Dickson. Sue & Jim Lavarack and Sue & Dtw 
Scott-Orr. Viial assistance in the field has con>e 
from many hundreds of volunteers as well us staff 
and postgraduate students of ihe Universiiy of 
New South Wales. 

LITERATURE CITED 

ARCHER, M.. GODTHELP, H.. HAND. S.J. A 
MEGIRIAN. D. I9S9. Fossil mammals of 
Riverslcigh. Nortli Wcsicm Queensland: prclimi- 
Tiary ovcr\'icvv of biostnUinraphy. conrlalion uiid 
cnvlfonmcmal cliac^gc. Australian Zoologist 25: 
29-65. 

ARCHEl^, M-. HAND, S I & GODTHELP. H. 1994. 
Ri^'erslesch. Second Edition (Reed: Sydney) 

ARCHER. M-.'HANO. S.I At CiODTHELP. H. 1495- 
Tertiarv enviionntcnial and biotic change in Aus- 
iralia. "pp. 77 tn) In Vrha, E.S., Demon, GM,, 
Piirtridge. TC. & Buivklc. L.H. (eds), Paleocli- 
malc and cvoliiiion, willi cmphasi.s on huniai) 
origins. (Yale University Press New Haven). 

ARENA, K. |y06. Dunsinauc Site: the case of the 
Tertiary time cnpsulos. Rivcrsieigh NtXesSO; 4-6. 

ARENA, K. 1^97. The palaeontology and geology oi 
Dvinsimme -Site. Memoir?; of the Queciwland Mu^ 
seum^t: 171-17^). 

BLACK, K. 1997a. Diversity and biostraligraphy of the 
Diprotodonloidea ol^ Riverslcigh, nortii western 
Queensland Memoirs of the Queensland Mu.seum 
41: 187-192. 

BLACK. K. I^97b. A new .species of Palorchcstuiie 
(Marsupiiilia) from Ihe laie middle to early late 
Miocene Encore Local Riuna, Riverslcigh, norlti 
western Queen.sUuid. Memoirs oftlu'OuecnslanJ 
Muscuni4I- IS1-I85 



314 



MEMOIRS OF THE QUEENSLAND MUSEUM 



BOLES, W. 1997. Riversleigh birds as pal- 
aeoenvironmental indicators. Memoirs of the 
Queensland Museum 41 : 241-246. 

COOKE, B.N. 1997. Biostratigraphic implications of 
Riversleigh fossil kangaroos. Memoirs of the 
Queensland Museum 41 : 295-302. 

CREASER, P.H. 1977. Lithogenesis and diagnostic 
features of recent and ancient terrigenous lime- 
stones (calclithites). Unpubl. M.Sc. Thesis, ANU. 

FRAKES, L.A., MCGOWRAN, B. & BOWLER, J.M. 
1987. Evolution of Australian environments. Pp. 
1-16. In Dyne, G.R. & Walton, D.W. (eds), Fauna 
of Australia, Vol. lA, General Articles. (Austra- 
lian Government Publishing Service: Canberra). 

GAFFNEY, E.S., ARCHER, M. & WHITE, A. 1992. 
Warkalania, a new meiolaniid turtle from the 
Tertiary Riversleigh deposits of Queensland. The 
Beagle 9: 35-47. 

HAND, S.J., ARCHER, M., GODTHELP, H., RICH, 
T.H. & PLEDGE, N.S. 1993. Nimbadon, a new 
genus and three new species of Tertiary 
zygomaturines (Marsupialia: Diprotodontidae) 
from northern Australia, with a reassessment of 
Neohelos. Memoirs of the Queensland Museum 
33: 193-210. 

MEGIRIAN, D. 1992. Interpretation of the Miocene 



Carl Creek Limestone, northwestern Queensland. 
The Beagle 9: 219-248. 

MEGIRIAN, D. 1994. Approaches to marsupial 
biochronology in Australia and New Guinea. Al- 
cheringa 18: 259-274. 

MYERS, T. & ARCHER, M. 1997. Kuterintja nganm 
(Marsupialia, Ilariidae): a revised systematic anal- 
ysis based on material from the late Oligocene of 
Riversleigh, northwestern Queensland. Memoirs 
of the Queensland Museum 4 1 : 379-392. 

TEDFORD, R.H. 1967 Fossil mammals from the Carl 
Creek Limestone, northwestern Queensland. Bul- 
letin of the Bureau of Mineral Resources, Geology 
and Geophysics, Australia 92: 217-236 

WHITE, A., 1997. Cainozoic turtle assemblages from 
Riversleigh, northwestern Queensland. Memoirs 
of the Queenland Museum 41:41 3-42 1 . 

WOODBURNE, M.O., MCFADDEN, B.J., CASE, 
J. A., SPRINGER, M.S., PLEDGE, N.S., 
POWER, J.D., WOODBURNE, J.M. & 
SPRINGER, K.B. 1994. Land mammal 
biostratigraphy and magnetostratigraphy of the 
Etadunna Formation (Late Oligocene) of South 
Australia. Journal of Vertebrate Paleontology 13: 
483-515. 



PALORCHESTESAZAEL {MAMMALIA. PALORCHESTIDAE) FROM THE LATE 
PLEISTOCENE TERRACE SITE LOCAL FAUNA. RIVERSLEIGH. NORTHWESTERN 

QUEENSLAND 

A.C. DAVIS AND M. ARCHER 



Davis, AC & Archer, M., \^91 0(^30. PalorchesU-s aznel M2mnra\\d.?i\\orchtsi\6ac)imm 
the laic Pleistocene Terrace Site Local Fauna, Riversleigh, nonhvesiern Queensland. jWfw- 
oinofthf Queensland Museum 41(2): 315-320. Brisbane. ISSN 0079-8S35. 

A niaxilla of Pr//r?/r/7fA7f i/3.:fle/ IS described (rom gravel deposits at Terrace Site, Riversleigh 
Station. A rudiocarbon dalcol 23.900 +4 1 00 -2700 years BP is reported from the fossiliierous 
unit ai Terrace Site supporting previous interpretations of a fate Pleistocene age for the 
Terrace Site Local Fauna. 

Anj?e!a Davis, Geology Department, Australian National University; Canberra ACT 0200, 
Australia {Present address: Western Australian Museum, Francis Street, Perth. Western 
Australia 6000, Atistralia) : Michael Archer, School of Biological Sciettce, University of New 
South Wales, Svdnex, New South Wales 2052, Australia; received-^ December 1996. 



Terrace Site occurs in unconsolidated tluviatilc 
sediments on ihc eastern bank of the Gregory 
River 5 kin downstream from the Lawn Hill road 
crossing on Riversleigh Station, NW Queens- 
land. These deposits were interpreted as 
Pleistocene gravels resting on Tertiary and Cam- 
brian limestones (Archer ex al.. I9S9, 1994). The 
Teirace Site Local Fauna is listed in Archer ei al. 
(1994). 

Material is deposited in the Australian Museum 
(AMF), the Natural History Museum, I-ondon 
(BM). Museum ofVictoria(NMVP). Queensland 
Museum (QMF), South Australian Museum 
(S.\M), Department of Geology. James Cook 
University (P). Molar number follows Luckeu 
( 1993); premolar number follows Flower ( 1 867); 
molar crown morphology follows Archer 
(1984). 

STRATIGRAPHY AND AGE 

Terrace Site has u 3m high cross-section 
through hori/.onla! and lenticular beds in an up- 
wardly fining sequence. The basal sediments are 
poorly sorted, light grey sands and gravels with 
abundant mussel shell fragments and most of the 
venebrates including QMF308H2 The section 
grades upwards into finer sands, silts and clays 
with finer shell fragments. Charcoal particles up 
to 5mm occur throughout in small lenses or iso- 
lated fragments. The charcoal occurring in lenses, 
maxilla and most other specimens being un- 
abraded and the recovery of articulated material, 
suggests that al least part of the fauna and associ- 
ated charcoal is a primary accumulation. 

Based on the mammal fauna the age was inter- 



preted as possibly late Pleistocene (Archer el 
al ., 1 994). Charcoal from the basal bone-rich layer 
that coniained QMF3()882 gave a conventional 
radiocarbon date of 23.900 +4100 -2700 BP 
(ANU-7620). Although the standard eiTors arc 
high, the range within two standard errors con- 
firms the Late Pleistocene age. 

SYSTEMATICS 

Order DIPROTODONTIA Owen, 1866 
Suborder VOMBATIFORMES Woodburnc. 

1984 

Family PALORCHESTIDAE Tate, 1948 sens. 

Archer & Bartholomai, 1978 

Palorchestes Owen, 1873 

TYPE SPECIES Pahrclwstes azael Qwcu, 1873. 

PalorchestesazaelOwcn, 1873 
(Fig. 1: Table!) 

MATERIAL. QMF30882, a left maxillary fragment 
with near complete M^'^, Ihc fragmented alveolus of 
P3. and a portion of the palate and jugaJ. 

DESCRIPTION. Upper molars high-cnnvncd, 
bilophodont, trapezoidal in occlusal view, with 
the prololoph wider than the metaioph. Lophs 
broad at their bases, narrow toward the apices, 
slightly crescentic. Broad anterior and posterior 
cingula on each tooth, extending around the tooth 
forming narrow lingual and buccal cingula. 
Molar enamel crcnulated and rough on anterior, 
posterior and intcr-loph surfaces, but smooth on 
lateral surfaces. 



316 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. I. Palorchestes azael Owen, 1873, QMF30882 from Terrace Site, maxillary fragment with M*-M\ in 
occlusal view. a.c.=anieriorcingulum; p.c =posteriorcingulum; l.c.=lingual cingulum; f.].=forelink;m.l =mid- 
link; h.l.=hindlink; La.l.=lingual accessory link; pr=protocone; pa=paracone; me=melacone; mcl=metaconuie; 
pri=protoloph; mel=meialoph;p.s.=palatal sinus. 



PALORCHESTESAZ\EL?ROSi RIVERSLEIGH 



317 



Tabic I. Dimensions (mm) of upper cheek leeih of O^^F30882 and comparaiive samples otP. azaei (BM463I6. 
A1VIF452. PIS6593, AMF7272, QMF7074. QMF772). P pamfs (from Woods 1958). P painn (from 
Woodlmrne. \%l)mdP. selesriae (from Mackness, 1995). The dimensions of ihc ho!oiypc of A tizMt/equjI 
the minimum vahies of ihe observed ranges. e = estiniale; H=holoiype 







QMF 

308R2 




/' am 


=/ I 


P pan'iis 




P- pan 


ii 


P 

.sdtLSiuift 

(H) 


N 


MEAN 


RA.NGE 


N 


MHAN 


RANGE 


N 


MEAN 


RANGE 


p' 


L 


17.0c 


9 


20.8 


18,7-23.7 


1 


16.8 




4 


13.9 


13,8-14,4 


. 


M' 


L 


28.8 


6 


26.3 


23.5-28.7 


4 


19.6 


17.8-22.1 


4 


17.4 


16.5-18.2 


22.6 


AW 


24. 1 e 


6 


23.0 


19.3-27.7 


4 


15.5 


14.2-17.3, 4 


M.2 


13,6-144 


166 


PW 


2? .5 


6 


21.9 


I7.1-23-.0 


3 


_ 


■ 3 


13.5 


13 2- 15-8 


16.9 


M- 


L 


Z92c 


5 


27.2 


24.9-29.2 


3 


20,1 


18.7-20.0 ; 3 


18,1 


1 7-8- 18.1 




AW 


27.2 


5 


24-5 


21-5-26.8 


1 


16.3 


15.0-17.5 


T 


15.4 


14.8-15-p' - 1 


PW 


24.7 


5 


22.3 


19.9-24.7 


3 


- 




3 


13.6 


I3.M4.3 




M^ 


L 


31,5 


4 


27.4 


25.3-31.5 


4 


21 1) 


19.6-22,3 


4 


1 8,5 


1&.0-J9.4 




A^^ 


26,2 


3 


22.6 


19.726.0 


3 


16.6 


15.9-17,5 


3 


15.3 


I5.0-I6.3 


- 


PW 


24.7 


3 


20-5 


18.1-23. 1 


4 


- 


' 4 


13.3 


I26-14-I ! 


m'-m^ 


L 


85.2 


3 


79.5 


74.0-86 4 


- 1 


i - 


■ 


- 


- 



P^ missing- Partly preserved alveolu.s suggest- 
ing P^ as subiriangular, ahoul 17 mm long. 

M' nearly complete, missing only a small part 
of the enamel of the buccal side of ihe protoloph, 
Willi 2 high forclinks joining the piotoloph to a 
broad anleriui cmgulum. Lingual lorelink offsei 
diagonally, joining the anterior cmgulum from 
ihc protoconc at ihe midline of the looih. Deep 
pits between, and on both sides of, the forclinks. 
A high double-midlink between the lophs. On Ihe 
lingual side of ihe double-midlink is un accessory 
link forming a deep lingual pit. Lingual accessory 
link high and well-developed on M' PoslcrioT 
cingulum compressed by molar crowding Jfom 
M*, an irregular folded posterior face of the 
mctaloph formed from a thickened cresl 
(hindlink) descending posteriorly from ihc 
meUtconule. 

M- larger than M'. missing the anierolingual 
face of the protoloph. with a single weak lorelink. 
and midlink a single (cf. double in M') high link 
following the midline of the tooth. Lingual acces- 
sory link V-shaped in lateral view, differing from 
the siraighter link of M' and slightly lower. An- 
lenor and posterior cingula well-OeveJoped, an- 
icriorcinguium pressed inioM' because of molar 
crowding. Hindlink present. 

M^ siriiilar in si/e and morphology to M% dif- 
ferences in length being atlribuled to molar 
crowding, Porclink. midlink and hindlink all sin- 
gle. Lingual accessory midlink more reduced 
than in M-*. 

Base of the jugal projecting perpendicuUir lo 
Ihc maxilla, its anterior edge originating at Ihe 
M'-M^ conuict beibre sloping gradually posteri- 



orly, with its posterior edge peqiendicular to the 
maxilla at the M-^-M-^ junction. A few fragments 
of the orbital surface preserved on the upper part 
of the specimen. A minute palatal sinus level wilh 
the posterior edge of P^ alveolus 1 2 mm Irom ihc 
lingual edge of the looih row. Only the left side 
of the palate preserved, exieiuling from ihc totilh 
row to neai- the midpalaial suture. 

COMPARISON. QMF30S82 compares well 
with Woods (1958) diagnosis for P. azaet and its 
molar dimensions (Tabic 1 ) lie within ihc ranges 
for P. azael and outside those of other species of 
Palorchestes. When compared wilh a cast of the 
holotype (BM463I6) and description by Owen 
( 1 873. pi. 82 fig. 1 ). Ihe Riversleigh specimen i;* 
larger in all dimensions but is otherwi.sc similar. 
The holotype is the smallcsi of ihe com()aralivc 
sample of A tiztu'l examined. 

QMF30882 is most similar lo AMF452, from 
Wellington Caves (Dun, 1893, pi. 16), in which 
the only notable difference is the lingual acccs- 
sopyiiiidlmk being only si ighily developed in M'. 
less so in M-and abscm in M-\ OMP3()S82 show% 
a similar gradient of development from wcll-dc- 
vclopcd anteriorly to simplest posteriorly. 
QMF772 {Woods. 1958, fig. I) also shcvws a 
lingtial accessory midlink gradient, and is inicr- 
mediale in development belween the othci iwo 
specimens. SAMP3I370 and 31371 (Pledge. 
1991, fig. 4) also show ihe lingual midlink. 
PI 86593 Iron) the Wyandotte LF (McNamara. 
1990^ does not show the midlink at all. and it is 
unclear m Llic worn hulolype BM4(i3l(i mA 
.\.\!F7272 Tl)e CMt>3me cxprcsskm of this cbiir- 



n& 



MEMOIRS OF THE QUEENSLAND MUSEUM 




■ sites wKh P. azMt 
D sites with P. spp. 
other than Pa/ae/ 



V 



r Riverelelgh (a) 1 

2, Wyandotte (o?) 2 

3, Bluff Downs (o) 1 
A Alcoola W 5 

5. Chinchilla Sands (P,X) 3 

6 Eastern Darling Downs - Macalister. Gowrle, 

KlngsCreek, Warra. St Ruth (flt,X,p) 11 

7 Cemen! Mills, Gore (af) 2 

8 Tea Tree Cave (X) 1 
9, Myall Creek (09 1 

10 Bow(P)1 

11. Cuddy Springs (X) 1 

12, Wellington Caves (b.P) 3 

1 3 Wee Jasper Caves (p) 1 

14 Grong Grong (a) 1 

15 Buchan Caves («,X) 4 

16 Gipp's Land, Tamtx) Rrver («) l 

17 Son^nto (a) 1 

18 Werribeefojl 

19. Hamltton Grange Bum {?■) 1 

20 near Keilor (o) 1 

21 - Terang {^ 1 

22, Spring Creek («) 1 

23, Strathdownie(a.P)2 

24, Goulden's Hole Cave (P) 2 
25 Henschke Fossil Cave (?«) 2 
26, Victona Cave (e^ 1 

27 Thebarton («) 1 

28. Tooiapinna Fauna, Warburton River ("?•) 1 

29. Curramulka («) 1 

30. Scotchtown Cave (X) 1 
31- Mowbray Swamp («) 1 
32 Pulbeena Swamp (a) 1 

uncertain locality (b,P,X] 26 specimens 



KEY TO SPECIES SYMBOLS 

W P. azaef, R cf, azaet 

Ip) P parvus, P. cf. parvus 

W P painei 

(a) P selestiae 

(X) P sp., P sp. nov, R sp. indet. 



FIG. 2. Distribution oi^ Palorchestes in Australia. The minimum number of individuals at each site (number atler 
the species symbol in the site listing) includes published and unpublished specimen^. 



acier in QMF30882 is within the inlraspeciilc 

range of variation. 

The large variation in molar size for P. azael 
(Table I), coupled with intraspecific variation 
support Woods' (1958) view thai it is a highly 
vaiiablc species. Molar morphology grades along 
the looth row from complex anterior molars with 
additional links to simple diprotodoniid-like lecih 
posteriorly. Sample sizes are too small to estimate 
coefficients of variation for molar dimensions, or 
identify sexual dimorphism. 

The Riversleigh specimen is larger than P. 
ponms from the early Pliocene Chinchilla Sand 
(Woods, 1958) (Table 1). Its molars are relatively 
broader resulting in squarer molars in occlusal 
view. The lack of a double hindlink on the M' as 
in QMF789 (Darling Downs;Woods, 1958. tlg.4) 
and NMVP48987 (Buchan Caves) also dis- 
linguishcs the two species. 

The Riversleigh specimen differs from P. 



selestiae, from the early Pliocene Bluff Downs 
Local Fauna (Muckness, 1995), in being about 
25% smaller and in the lingual forelink being in 
contact with ihc anterior cingulum rather than 
terminating in the cingular basin. 

It differs from P. painei, from the late Miocene 
at Alcoota (Woodbume, 19(i7) in being signifi- 
cantly larger (Table 1 ), with more complex molar 
moiphology including higher crowns, having a 
double forelink on MI. and in having a single 
midlinkon M2. 

DISCUSSION. Over 80 Plio-Pleistocenc Pal- 
orchestes specimens have been registered with 
museums from at least 32 open sites and cave 
deposits throughout Australia (Fig.2). P. azael, 
the most widely distributed species, is present in 
21 ofthe 32 sites. 

Few sites with P. azael have radiocarbon dales 
available. Published dales range from 



PALORCHESTESAZML FROM RIVERSUIGH 



m 



iy.800±390 BP at Spring Creek (Flannery & 
Con, 1984) to 54,200 +l],(>00 -4,500 DP a( 
Pulbeena Swamp (Banks ei i*l.. IQ7(>)iirtd 30.400 
+750 -700 BP at Wyandotte (McNamara, 1 990). 
Specimens at Weiliiiglun Caves. Naracoorlc 
Caves (Wells et al., 1984) and the Warburton 
River (Toolapinna Fauna; Tedford ci al., 1992} 
could be considerably older. 

The presence of 2 species of Palorrhestcs at 
Cement Mills (Bartholomai, 1977), Wellington 
Caves. Buchan Caves, and Slralhdownie (Mann- 
cry & Archer, 1 985 ) has been iiiierpretcd by some 
lu mean that they arc mixed Pliocene and 
Pleistocene assemblages. Allcmaiively F. patrux 
may have extended into the Pleistocene as sug- 
gested by Bartholomai (1977) or Pleistocene P. 
parvus may be incorrectly assigned. 

.•\Jthough widespread. P. azoei specimens arc 
runr at any one site, generally being represented 
by only one or two indiviiiaals and no vuher 
I'ocisils (at least 8 known sites) or a single speci- 
men of P. azael with associated faunas (at least 8 
sites). The low density may be an artefact of 
preservation, but could indicate that P. cKMel was 
a solitary animal (Flannery & Archer. 1985). 

Al Terrace Site P, azael is associated with the 
large to medium-si2eii herbivores in a fauna dom- 
inaied by riverine turtles, cR)codiles. water ratii 
and fish (Archer el al., 1994). The palaeoenviron- 
mcnt IS inicrpreted to have been siinil:ir to thai 
which characterises the area today (Archer el al., 
1994). The other northern Queensland assem- 
blage, the Wyandotte Local Fauna (McNamara. 
1990), contains a similar faunal assemblage 
More southern faunas are dominated by large 
browsing and gra/Jng manmiaJs (c.jj. Diprotoihn 
opuifuin and diverse Mcuntpus, Ptoiemnvdon 
and Sthcnurus spp.) in open sderophyll forest 
(Barlholumai. 1977), Eucalyptus woodland 
(Bankset al. 1976). low heath (Flanneiy & Gow. 
1984) and trees & shrubland (Dodson cl al., 
1993). P. azoel evidently loieiatcd a wide range 
ofelimaiie conditions and habitai types. 

ACKNOWLEDGEMENTS 

Wc acknowledge support from: Australian Re- 
search Grant Scheme. The University of NSW. 
the National E.'^tateGrants Scheme (Queensland), 
the World Heritage Unit in Canberra, the Depiu^t- 
ment of Envia>nmenl. Sports and Territories, (he 
Queensland National Parks and Wildlife Service 
(particularly Paul Sheehy). the Waanyi People 
and ihc Carpentaria Land Council, ICI Aiisiraliii, 
the Australian Geographic Sot-ieiy. ihe Qi>cen&- 



land Museum; the Australian Museum, Century 
Zinc (particularly Doug Fishburn), MIM, Ihc 
Mount Isa City Council. Surrey Beauy & Sons, 
the Rivcrslcigh Society; private supporters in- 
cluding Elaine Clark. Sue & Jim Lavarack. Si>e 
& Don Scotl-Orr. Margaret Beavis and Maain 
Dickson: research coTleagues notably Henk 
Godihelp, Suzanne Hand, Alan Bajthol(»mai, Phil 
Creaser Peier Murray, David Ride, Sue Solo- 
mon. Arthur White. Anna Gillespie. Virginia 
O'Donoghue, Cathy Nock, SypPrascuihsoukai>d 
Stephan Williams; and postgraduate studenlji 
working on Riversleigh fossil maicriids who have 
generously shared ihcir understanding including 
Bemie Cooke, Jeanelle Muirhead. Paul Wilbs 
and Anita Van der Mcer Panicular thanks arc 
owed to Sue and Jim Lavarack who spent many 
years slogging to and from Terrfice Site us leaders 
of the Tcrraeisis'. 

LITERATURE CITED 

ARCHER, M. 1 984. The Australian marsupial radia- 
tion. Pp. 633- 808. In Archer , M. <t Clayton, G 
(eds). Venebraie zoogeography and evolution In 
Australasia. (Hesperian Press iPcrtJi). 

ARCHER, M , GODTHELP, H., HAND, S.t. At 
MEGIRIAN, D. 1989. Fossil mommids of 
RivcTslcigh, northwestern Quccnslund. preliinir 
nao' ovcniew of biosiraiigraphy. conclaiion and 
environmental change. The Australian Ziwlogisi 
25. 29-6.5- 
1994, Rivcrsleigh. 3nded. (ReediSydney). 

RANKS, MR . COLHOUN. E.A. & VAN DEGEFi?, 
G. 197fi, Lute Quaccmary Pulorchesies lauti 
(Mammalia, Diprototlomidae) from nonhwestcm* 
Tasmaitia. Alchcringa I: 159-166. 

BARTHOLOMAI, A. 1977- The fossil vertebrate fauna 
from Pleistocene deposits at Cement Mills, Gore, 
southeastern Queensland- Memoirs of Oie 
Queensland Museum 18:41-51. 

DODSON, J . FliLLAGAR. R . FLRBY, J.. JONF-S, 
R. (te PROSSER, 1. 1993. Humans and rnegiilauna 
in a iaic Pleistocene cnvironmcni Inom Cuddic 
Springs, north wcsiern New .Stmih Wales. An:hac- 
ology m Oceania 28. 94-99. 

DUN. W.A. 189:^ On palatal lemaiuii ol PuhivheHes 
(iuiel. Owen. Irom the Wcllingu»n Ciwvs hone 
deposit. Records of the Gcolaglcal Survey of New 
South Wales 3: 12-124. 

FLANNERY, T.b. ^ ARCHER. M. 1985. A//cn/;«r'Siri 
Owen. 1874. Large and small p<ilorchcstids Pp 
234-239. In Ricli, P.V. & Van Tots. G. (cds). 
Kadimakara exlincl vcrtehratcs of Auslralia. (Pi- 
oneer Desi^M) Studio: Lilydale. Victoriii). 

FLANNERY. TF, & GOTT. B. I9S4. The Spn'ng 
Creek Locality, souihwcsicm Victoria, a laie sur- 
viving mcgafaunal assemblage, llie Aui^raiiiin 
Zo.>logJ5t 21: 385-422 



320 



MEMOIRS OF THE QUEENSLAND MUSEUM 



FLOWER, W.H. 1867. On the development and suc- 
cession of teeth in the Marsupialia. Philosophical 
Transactions of the RoyalSocieiy of London L57: 
631-641. 

LUCKETT, W.P. 1993. An ontogenetic assessment of 
dental homologies in therian mammals. Pp. 182- 
204. In Szalay. F.S., Novacek, M.J. & McKenna, 
M.C.(eds). Mammal phylogeny; Mesozoic differ- 
entiation, multituberculaies, monotremes, early 
therians, and marsupials. (Springer- Verlag, New 
York). 

MACKNESS. B. 1995. Palorchestes selesiiae, a new 
species of palorchestid marsupial from the early 
Pliocene Bluff Downs Local Fauna, northeastern 
Queensland. Memoirs ofthc Queensland Museum 
38: 603-609. 

MCNAMARA, G.C. 1990. The Wyandotte Local 
Fauna: a new, dated. Pleistocene vertebrate fauna 
from northern Queensland. Memoirs of the 
Queensland Museum 28: 285-297. 

OWEN, R. 1873. On the fossil mammals of Australia 
part IX. Family Macropodidae; Genera 
Macropus. Pachysiagon, Leptosiagon. Pro- 
copiodon and Palorchesies. Philosophical Trans- 



actions of the Royal Society of London 164: 783- 
803. 

PLEDGE, N.S. 1991. Occurrences of /'a/orc/ie.srfs spe- 
cies (Marsupialia: Palorchesiidac) in South Aus- 
tralia. Records of the South Australian Museum 
25: 161-174. 

TEDFORD, R.H., WELLS, R.T. & BARGHOORN. 
S.F. 1 992. Tirari Formation and contained faunas. 
Pliocene of the Lake Eyre Basin, South Au.stralia. 
The Beagle, Records of the Northern Territory 
Museumof Arts and Sciences 7; 173-194. 

WELLS, R.T., MORIARTY, K. & WILLIAMS. 
D.L.G. 1984. The fossil vertebrate deposits of 
Victoria Fossil Cave, Naracoorte: an introduction 
to the geology and fauna. The Australian Zoolo- 
gist 21: 305-333 

WOODBURNE, M.O. 1967. The Alcoola Fauna, Cen- 
tral Australia. Bulletin of the Bureau of Mineral 
Resources Geology and Geophysics Australia 87: 
1-187. 

WOODS, J.T. 1958. The extinct marsupial genus Pal- 
orchestes Owen. Memoirs of the Queensland Mu- 
seum 13: 177-193. 



PRISCILEO ROSKELLYAE SP. NOV. (THYLACOLEONIDAE, MARSUPIALIA) FROM 
THE OLIGOCENE-MIOCENE OF RIVERSLEIGH, NORTHWESTERN QUEENSLAND 

ANNA GILLESPIE 

Gillespie, A., 1 997 06 30. Priscileo roskellyae sp. nov. (Thylacoleonidae, Marsupialia) from 
the Oligocene- Miocene of Riversleigh, northwestern Queensland Memoirs of the Queens- 
land Museum 41{2): 321-327. Brisbane. ISSN 0079-8835. 

Upper dentition of the marsupial lion Priscileo roskellyae sp. nov. from the Upper Site Local 
Fauna of Riversleigh provides the first detailed information about upper dentition of the 
genus. The upper adult dental formula is 11-3, CI, PI -3, Ml -4. This species is smaller than 
P. pitikantensis and is the most plesiomorphic thylacoleonid. Relative to Wakaleo, P3 is less 
bowed and molars are square and have a metaconule at their posterior margin. 
\Z\Thylacoleonidae, Wakaleoninae. Priscileo, Riversleigh, Oligocene, Miocene. 

Anna Gillespie, School of Biological Science, University of New South Wales, New South 
Wales 2052, Australia; received 15 February J997. 



The marsupial lion genus Priscileo Rauscher, 
1987 contains only P. pitikantensis Rauscher, 
1 987, from the late Oligocene Ngapakaldi Local 
Fauna, S. Aust which is known only from a max- 
illary fragment, a few teeth, and a number of post 
cranial elements. Additional Priscileo material 
has been recovered from the Oligocene-Miocene 
of Riversleigh, northwestern Queensland. This 
material includes a near complete skull from the 
Upper Site Local Fauna representing Priscileo 
roskellyae sp. nov. 

Dental terminology for molars and the last pre- 
molar follows Luckett ( 1 993) who has shown that 
a molariform dP3 (Ml of Archer, 1978) in mar- 
supials is replaced by P3. Accordingly, the re- 
maining molars represent Ml -4. However, 
homology of the other premolars follows Flower 
(1 867). Material is housed in the Commonwealth 
Palaeontological Collection, Bureau of Mineral 
Resources, Canberra (CPC), Northern Territory 
Museum (NTM), Queensland Museum (QMF), 
South Australian Museum (SAMP), Museum of 
Palaeontology, University of California, Berke- 
ley (UCMPB). 

SYSTEMATICS 

Superorder MARSUPIALIA Illiger, 1811 

Order DIPROTODONTIA Owen, 1 866 

Family THYLACOLEONIDAE Gill, 1872 

Priscileo Rauscher, 1987 

TYPE SPECIES. Priscileo pitikantensis Rauscher, 

1987. 

DIAGNOSIS. Small; P3 length usually less than 
12mm; Ml and M2 relatively square in basal 



outline, with a posterolingual metaconule; ante- 
rior root of the zygomatic arch projecting an- 
terolaterally dorsal to M2-3. 

Priscileo roskellyae sp. nov. 
(Figs 1-3) 

MATERIAL. Holotype QMF23453. a skull with left 
and right P3, Ml-2, alveoli for left and right 12-3, CI, 
Pl-2, M3-4, and partial alveoli for the left and right II 
from early Miocene Upper Site, Godthelp Hill, 
Riversleigh. 

ETYMOLOGY. For the former Australian Minister of 
Arts, Sport, the Environment, Tourism and Territories, 
the Hon. Ros Kelly, who provided significant support 
for the Riversleigh Project. 

DIAGNOSIS (by comparison with the type and 
only other species). Smaller; P3 approximately 
2/3 as long; metaconule on Ml and M2; alveolus 
of PI closer to the P2 alveolus than to the canine 
alveolus; lingual root on Ml smaller, not intrud- 
ing as far medially into the palate. 

DESCRIPTION. Upper dentition. Formula 11-3, 
CI, PI -3, Ml -4. Alveoli for II large and incom- 
plete. Alveolus for 12 smallest of incisor alveoli. 
Alveolus for CI ovoid, larger than alveolus for 
13, smaller than that for II . Canine alveolus closer 
to 13 alveolus than to PI alveolus. Two small 
alveoli for two single-rooted pretnolars between 
the canine and P3. CI and PI alveoli separated by 
an approximately 3 mm. P2 alveolus close behind 
that for PI , abutting the anterior base of P3. 

P3. As in Wakaleo but 25-66% smaller. Posterior 
portion only slightly broader than the anterior 



322 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 1. Priscileo roskellyae sp. nov.. holotype, QMF23453, partial skull, from Upper Site, Godthelp Hil 
Riversleigh. Ventral view slereopair, showing upper dentition. 



portion (unlike Wakaleo where the posterior is 
much broader). Longitudinal blade slightly in- 
wardly-curved in contrast to the distinctive in- 
wardly-curved blade of most Wakaleo. 
Relatively uniform width with gently curved lon- 
gitudinal blade giving P3 rectangular shape. Lon- 
gitudinal blade running between 2 major cusps. 
In buccal view, the shearing blade of P3 W- 
shaped, the longitudinal blade foiming the rise in 
the middle, and the anterior and posterior blades 
ascending from the major cusps at each end. The 
anterior cusp is slightly higher than the posterior 
(as in W. alcootaensis). In W. vanderleueri the 
cusps are approximately equal in height. Three 
vertical blades, one anterior, one lingual, and one 
buccal, ascending from the anterior cusp to the 
base of the crown. Anterior blade curving lin- 
gually as it ascends, bending slightly posteriorly 
before merging with the base of the crown. Lin- 
gual curvature of blade producing a small vertical 
lip towards the base of the blade. Similar lip in 
some Riversleigh Wakaleo. Posterior to this blade 
lingual face of the crown curving concavely 
forming an anterolingual basin. Lingual blade 



curving slightly anteriorly as it ascends and 
merges with the base of the crown. Many T. 
carnifex and T. crassidentatiis also with lingual 
blade in contrast to W. alcootaensis and W. 
vanderleueri in which it is absent. Posterior to the 
lingual blade the lingual flank of P3 formiing a 
sharp depression extending from the base to the 
crown. Lingual flank following the curve of the 
posterior root, curving convexly to the posterior 
border. Buccal blade of the anterior cusp ascend- 
ing in a slight posterior direction, merging with 
the crown midway up the tooth. Short buccal 
blade also ascending from the posterior cusp, 
merging with the crown midway up the tooth, not 
as prominent as the anterior buccal blade. Similar 
anterior and posterior buccal blades in Wakaleo. 
Posterior blade running posterolateral ly from the 
posterior cusp to the posterior margin of the tooth, 
an extension of the longitudinal blade, contiguous 
with the preparacrisla of Ml, in P3 of Wakaleo 
but absent in Thyiacoleo. Short oblique blade at 
termination of posterior blade at the posterior 
margin of P3, ascending anterolaterally on the 
buccal flank, merging midway with the base of 



PRISCILEO ROSKELLYAE SP. NOV. FROM RIVERSLEIGH 



323 



TABLE 1. Dimensions ofuppercheekleeih of species 
of Priscileo and Wakaleo. Data from Clemens & 
Plane ( 1 974), Archer & Rich (1982), Rauscher ( 1 987) 
and Murray & Megirian ( 1 990). Measurements for P. 
pitikaniensis (except P3 ) were taken from from a caste 
of UCMP88448. a=a!veolus measurement. 





Length (mm) | 




P3 


Ml 


M2 


M3 


M4 


P3/M1 1 


P. roskellvae \\ 


QMF23453 


8.2 


5.8 


4.3 


3.3a 


2.9a 


i.4. 


P. pitikamensis \ 


UCMP88448 


1 1.0a 


6.5a 


5.3 


3.7a 


3.5a 


1.69 1 


U'. aicootaensis \\ 


NTMPI 


23.3 


14.7 


7.2 


1 


- 


1.58 1 


VV. vanderleueri \\ 


CPC26604 


17.9 


11.2 


7.0 


5.0 


- 


1.59 1 


Riversleigh Wakaleo 1 


QMF24680 


12.9a 


8.9a 


6.3 


4.9a 


4.3a 


1.44 


QMF23446 


12.6 


9.6 


6.0 


5.2 


- 


1.31 


QMR3443 


12.2 


9.4 


6.0 


4.5 


3.5a 


1.30 



the crown, lacking in Thylacoleo and W. ai- 
cootaensis. VV. vanderleueri with a small blade in 
a similar position, differing by commencing a 
short distance before the end of the posterior 
blade, more vertically oriented. On the buccal 
flank of P3, a broad valley running between the 
anterior and posterior buccal blades, much 
broader, in Wakaleo. 

Molars. Left and right Ml and M2 relatively 
unworn. Alveoli for M3 and M4 indicating 3 
roots for each; 2 equal anterior roots, with slightly 
larger posterior one. In Wakaleo vanderleueri 
anterior roots of M3 larger. Molar gradient steep, 
similar to P. pitikaniensis and Wakaleo. M 1 and 
M2 square, unlike the triangular molars in 
Wakaleo. Molar morphology similar to Wakaleo. 
Ml wider anterioriy than posterioriy. Paracone 
highest cusp. Small blade ascending anterioriy 
from the paracone to the anterior edge, joining 
siylar cusp B, contiguous with blade ascending 
from the posterior cusp of P3. Postparacrisla run- 
ning posteriorly, meeting the ascending pre- 
metacrisla, forming a notch midway along the 
straight centrocrista. Metacone well-developed. 
Postmetacrista ascending posteriorly from the 
metacone to the posterior margin of M 1 . Buttress- 
ing the steep lingual face of the paracone a short 
crescent-shaped preparaconulecrisia straighten- 
ing medially, terminating at a paraconulc. Short 
postparaconulccrista running posteriorly into the 



trigon basin. VV. vanderleueri and Riversleigh 
Wakaleo with much straighter and longer blade 
ascending lingually from the paracone. Prc- 
prolocrista arising from the anterior edge of the 
trigon basin medial to the paraconule, running 
postcromedially to the proiucone. From the pro- 
tocone a postprotocrista running posteriorly to a 
metaconule. Postmetaconulecrista curving 
posterolaterally from the metaconule. ascending 
to merge with the posterior margin of the 
postmetacrista. Short crescent-shaped ridge as- 
cending lingual face of metacone, terminating 
midway between metacone and metaconule. Ml 
of VV. vanderleueri and Riversleigh Wakaleo whh 
similar ridge. Blades joining the 4 major cusps 
forming margins of a deep, square trigon basin. 
Within the basin, fine enamel crenulations radiate 
outwards. Similar crenulatcd trigon basins occur 
in Wakaleo and T. crassidentatus. Buccal flank 
of M I , especially anterioriy. swollen resulting in 
a broad base for the siylar shelf. A stylar basin 
running from the posterobuccal face of the 
paracone to the posterobuccal margin of the 
metacone, becoming shallower posteriorly. Lat- 
eral wall of the basin lined with small vertical 
ridges. 

M2. M2 smaller than M 1 , lacking the distinctive 
broad siylar shelf and basin. Paracone highest 
cusp, more lateral than in M 1 . Short, semicircular 
preparacrista running anteromedially from the 
paracone. Small ridge buttressing lingual base of 
the paracone. Medial to this ridge a preprotocrista 
arising, running postcromedially to the pro- 
tocone. Protocone prominent, more anteriorly 
placed than in Ml. Protocone lingually bulbous 
as in P. pitikaniensis and Wakaleo. A narrow 
anterior shelf running from the base of the pre- 
paracrista to the protocone. Postprotocrista run- 
ning posterobuccally to a gently-rounded, 
posterior metaconule, producing squaring of the 
lingual outline. No metaconule on M2 of P. 
pitikaniensis but could be lost by damage to rear 
portion of the tooth. Short postmetaconulecrista 
running posterolaterally, merging with the poste- 
rior margin of the tooth. Metacone rounded and 
more posteriorly situated than in M I , with lingual 
blade running medially, connecting with a small 
blade running laterally from the metaconule, 
forming anterior border of a small, oval, basin at 
the posterior. Postparacrisla running 
posterolaterally to the buccal margin of the stylar 
shelf, converging with anlerolaterally orientated 
premetacrista. Deep, square, trigon basin be- 
tween the major cusps with fine enamel crenula- 



324 



MEMOIRS OF THE QUEENSLAND MUSEUM 



18.1 




FIG. 2. Prisciieo roskellyae sp. nov., balo(>pe, QMF23453, measurements (mm) of left tooih row.P^ on left, M' 
andM^. 



lions concentrated on the lateral and posterior 
walls. Crcnulatcd basins in M2 of P. paikantensi.s 
and Wakaleo. On the buccal margin of M2 a 
small, elongate, stylar basin parallel to the 
postparacnsla. terminating midway between the 
paracone and melacone. Similar basin in 
Riversleigh WakaU'ty. dii'tlcull to discern in the 
heavily worn M2 of W. vanderleueri. 

COMPARISON. P. roskellytje differs from all 
species of Wakaleo in: hein" smaller, the P3 of P. 
roskellyae is approximately 1/3 length of P3 of 
$V. akoQtaemis. Ml the length of P3 of W. 
vanderleueri, and 2/3 the length of P3s of 
Riversleigh Wakaleo; having a lingual crest as- 
cend from the anterior cusp of P3; having the 
shearing blade of P3slraighter: having the poste- 
rior root of P3 only slightly enlarged; having the 
posterior half of P3 relatively square in basal 
outline; having MI and M2 relatively square in 
basal outline. P. roskellyae differs from Tfiy- 
lacoleo in: being smaller; P3 being approxi- 
mately 1/3 length of P3 of r. /?///( and 1/6 length 
of P3 of T. carnife.w having M3-(4; having M 1 +2 
relatively square in basal outline. 

DISCUSSION 

The diagnosis of Prisciieo is amended to include 
P. roskellyae. Rauscher (1987) distinguished 
Prisciieo from other thylacoleonids by M4, a 
P3/M I length ratio less than 1 .70, and M2 with a 



crenulate, anieroposieriorly broad trigon basin. 
Some of the new Riversleigh Wakaleo specimens 
have these features. Two Wakaleo specimens 
have an M4 (Table I ). and most have a relatively 
broad crenulaied basin on M2. All species of 
Wakaleohavc ^P3/M\ ratio of less than 1.70. The 
value for A roskellyae (1.41 ) falls midway wilhin 
this range. 

Rauscher (1987) distinguished Prisciieo from 
Wakaleo by the loss of PI or 2 and M4 in the 
jailer, Although some new Riversleigh Wakaleo 
specimens have these teeth, Ihe plesiomorphic 
features of P. roskellyae exhibits (signiUcanily 
smaller si/e, square molar shape, metaconule and 
relatively straight cutting blade on P3) require 
generic distinction. P. pitikantensis and P. 
roskellyae ^harc generic features of denial dimen- 
sions, shape c»f M2 and position of the anterior 
base of Ihe zygomatic arch. Spccillc distinction 
of P. roskellyae is based on the si/e difference 
between these two species.P. pitikantensis being 
33% larger. 

INTRAFAMILIAL RELATIONSHIPS 

Rauscher (1987) found no synapomorphies unit- 
ing Prisciieo and Wakaleo, or uniting Prisciieo 
and Tbxiacoleo. Analysis of Prisciieo and Thy- 
lacoleo included comparison of postcranial ma- 
terial and for a number of character-slates, 
Thylacoleo exhibited the plesiomorphic condi- 
tion while Prisciieo was derived. Rauscher con- 



PRISCtLEO ROSKELLYAE SP. NOV. FROM RIVERSLEIGH 



325 



prpac 




me 



mcl 



ac alb linb 



5mm 



FIG. 3. Priscileo raskellyae sp. nov.. hokHype. QMP'23453. left cheek dentition. lb=longJtudinal blade: ab-an- 
lerior blade: pb=posierior blade; ac— anterior cusp; pc=posienor cusp: abb^anierior buccal blade: linb=lingual 
blade; pbb=posieriar buccal blade; aIb=anterior lingual basin: pa=paracone; pacl=paraconulc; prpac=pre- 
paracrista; popuc=posiparacri&ia; prpaclc=preparaconulecris»la; popack-postparaconulecrisia; pr=proiocone: 
mcl=nielaconule; pomck-posttnetaconulecrista: rrie=metacone; ponic=posimelacrista; prmc=premetacrista: 
5tB=siy!ar cusp B: trb=trigone basin. 



eluded that Thylacoleo\ priniiiivc fcaiures were 
secondarily derived and not a retained 
plesiomorphic condition. Because Wakaleo and 
Thylacoleo share the synapomorphy of loss of 
M-i, Rauschcr ( 1 987 ) considered them to be sister 
groups. Priscileo was regarded as the sister group 
of a Wakaleo/Thylacoleo clade. 

Rauscher (1987) suggested that loss of the 
melaconulc could he a diagnostic feature for the 
Thylacolconidae based on the irilubcrcular upper 
molars of Wakoleo and Priscileo and the second- 
arily quadritubercular Ml of Thylacoleo. How- 
ever, Ml and M2 of P. raskellyae have a 
metaconulc and are basically square in outline. 
The metaconule on these molars, especially on 
M2, is posteriorly positioned and its seeming 
absence from M2 of P, pirikantetisis may be a 
result of damage which is evident at the rear 
margin of this tooth. Consequently, loss of the 
metaconule can no longer be considered a syn- 
apomorphy for the family- 
Murray el al. (1987) placed Wakaleo and Thy- 
lacoleo in separate subfamilies: the Waka- 
leoninac includes Wakaleo; and the 



Thylacoleoninae which includes Thylacoleo iXJid 
possibly Priscileo. Wakaleonines were regarded 
to differ from ihylacoleonines in absence of PI 
and formation of a tympanic wmg composed of 
alisphenoid and squamosal contributions. Fea- 
tures distinguishing Ihylacoleonines from 
wakaleonines include PI. squamosal contribu- 
tion to the tympanic wing and frontal-squamosal 
contact on the lateral cranial wait. The new thy- 
lacoleonid specimens from Riverslcigh indicate 
that, in terms of dental morphology. Priscileo 
evhibits no features that prevent it from being 
ancestral to Wakaleo and Thylacoleo. The dental 
features of Priscileo, including small P3 and 
molar si?e, square (nearly bunodont) molar 
shape, metaconule. and lull premolar and molar 
complement are almt)si certainly plesiomorphic 
features within the family. 

However, these same features clarify some 
questions about relationships of the family with 
the Order Diproiodontia. ll was once commonly 
believed that ihylacoleonids evolved from a 
phalangerid-like diprotodontian which had 
quadritubercular upper molars including a hyper- 



326 



MEMOIRS or THE QUEENSLAND MUSEUM 



LrophicdmclacoTiulc(Krcll(, l872;Uroom. 1898; 
Bcnslcy, 1903; Ride, 1964: Archer, 1976). 
Archer ^ Rich (19821 hypoihcsised th;il ihe 
Iriluhcrcular shape of the molars of VV. nlaforneih 
jf'y wca' secondarily derived from an anee5.lral 
quadrituhcrcular shape through suppression of 
the niciaconule. Il has been suggesied ihal the 
iriani^ular moiurs of Wakalvo are plesiomorphic 
tor ihe family (Murray eta!., 1987). The primitive 
dental features of A* rnskellyae, especially Ihe 
meiaconule aud square molar shape, provide sup- 
port for Archer i<c Rich Uy«2). 

Priscileo and Wdkdh'o have been collected 
from Riversleigh's System B sites indicating 
overkipping ol early Miocene thylacoleonid lin- 
eages. Temixirai overlapping of species of T/n^ 
lacoleo (/". crassitietitattts <ind T. hilli) also 
occurred in the early Pliocene {Archer Sl Daw- 
son, 1982). In each case there was a distinct si/c 
difference in the lineages involved. In tcrmsolPJ 
length. System B spt^^imens of Waktilcr^ arc ap- 
proximately 1.5 limes larger than P. roskcllyae. 
Similarlv. P3 of */" crassiderttattis is iv^ice |he 
length of ihauooih in T. //////(Pledge, 1977). It is 
possible (hat si/e di|Terer)ces of this magnitude 
among sympatric lhylaci>lconids were an iinporl- 
iint factor in reducing compeiuion. Morphologi- 
cal studies of the limbs of P. pitikantensis suggest 
il was arboreal ( Rauscher, 1 9H7 ). Wakuleo. being 
larger Uiui nodouhi heavier, may have been n'lt^rc 
terrestrial. Murray ^ Megirian (199G) also inti- 
mated a icrresirial cxistchce fcir Wakaleo based 
on the wrist joint ant! heavily-worn dentition 
uhich they consider may indicate a .scavenging 
mode of life. 

ACKNOWLEDGEMENTS 

1 thank Michael Archer and Hcnk Godthclp for 
constructive comments and Siephan Williams, 
Karen Black and Jenni Brammall lor assistance 
with photography Support for research at 
Rivcrslcigh has coinc from the Australian Re- 
search Grant Scheme; liie National Csl;iie Grants 
Scheme (Queensland); the University of New 
^i>uth Wales; the Commonwealth I3epanmcnl of 
Environment. Sports, and Terrilones; the 
Queensland National Pinks and Wildlife Service; 
the Commonwealth Heniage I'nit; Id Australia; 
the Australian Geographic Society; the Queens- 
land Museum; the Australian Museum: the Royal 
Zoological Society of NSW the I jnnean Society 
of NSW; Century Zinc, Mount Isa Mines. Surrey 
Deaity *& Sons; the Riversleigh Society; and pri- 
vates supporters including Elaine Clark, Margaret 



Bcavis, Martin Dickson> Sue & Jim Eavarack and 
Sue & Don Scott -Orr. Invaluable assistance in the 
field has ci^mc from numerous volunteers, staff 
and postgraduate students of Ihe University of 
New South Wales. 

LITERATURE CITED 

ARC^III'.R, M. 1976. Phiiscolarclid origins and ihc po- 
Icnlial of ihc scleaodonl mular in tlie evolution of 
di()a»icxiont marsupials. Memoirs of ihc Queens- 
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ARCHER. M, & DAWSON. L. I9S2, Revision oi 
mnrsupiul lions of the genus Thyfctcolco Gei-vais 
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ARCHER M. & RICH. T.fl. I98I Results of ttic Ray 
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GITTLEMAN, J L. 198'). Carnivore group living: 
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PRJSCILEO ROSKELLYAE SP. NOV. FROM RIVERSLEIGH 



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J.L. (cd.). Carnivore behaviour, ecology and evo- 
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GUGGISBERCC.A.W. 1975. Wild cats of the world. 
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LUCKETT, W.P. 1993. An ontogenetic assessment of 
dental homologies in therian mammals. Pp. 182- 
204, In Szalay. F.. Novacek, M.J. & McKenna, 
M.C. (eds). Mammalian phylogeny. Mesozoic 
differentiation, multituberculates, monotremes. 
early iherians, and marsupials. (Springer-Verlag: 
New York). 

MURRAY. P., WELLS. R. & PLANE, M. 1987. The 
cranium of ihe Miocene thylacoleonid. Wakaleo 
vamlerleueri : click go the shears-a fresh bile at 
thylacoleonid systemalics. Pp. 433-466. In 
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MURRAY, P. & MEGIRiAN, D. 1990. Further obser- 
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occurrence and the distribution of Thy- 
lacoleonidae in South Australia. Records of the 
South Australian Museum 17: 277-283. 

RAUSCHER, B. 1987. hiscileo pitikantensis, a new 
genus and species of thylacoleonid marsupial 
(Marsupialia: Ttiylacoleonidae) from the Miocene 
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studies in evolution. (Surrey Beatty & Sons and 
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marsupials. Journal and Proceedings of the Royal 
Society of Western Australia 47:97- 131. 



ZYZOMYSRACKHAMI SP. NOV. (RODENTIA, MURIDAE) A ROCKRATFROM 

PLIOCENE RACKHAM'S ROOST SITE, RIVERSLEIGH. 

NORTHWESTERN QUEENSLAND 

H. GODTHELP 



Godthelp, H., 1997:06:30. Zyzomys rackhami sp. nov (Rodentia, Muridae) a rockrat from 
Pliocene Rackham's Roosi Site, Riversleigh. northwestern Queensland. Memoirs of the 
Queensland Museum 41(2): 329-333. Brisbane. ISSN 0079-8835- 

Zyzomys rackhami s^. nov. from the Pliocene Rackham's Roosi Site, Riversleigh, nonhwesi- 
em Queensland is the first fossil member of this genus and only the second Tertiary murid 
described from Australia. It is known from many hundreds of dental fragments recovered as 
a part of an ancient megadermatid roosting cave. It appears to be the most plesiomorphic 
member of the genus and is part of adiverse suite of extinct murids from this site. M^f^rf^'nrra, 
Muridae, Zyzonjys, Pliocene, Riversleigh. 

H. Godthelp. School of Biological Science, University' of New South Wales, New South Wales 
2052, Australia; 4 November 1996. 



The murid genus Zyzomys conlaixis 5 living spe- 
cies which are resiricied lo tropical Au.stralia. 
Zyzonm has been placed in the tribe Conilurini 
(Baverstock, 1984) and is an 'Old Endemic' 
scnsu Ride (1970) or an 'Older Immigrant* of 
TdXc ( 1 95 1 ). Along with Mesembriomys and Con- 
Hums, Zyzomys comprises a group of laxa unified 
by a number of cranio-dentai characters, phallic 
mtirphology (Lidickcr, 1987) ajid chromosomes 
(Baverstock etal.. 1981). 

Dental nomenclature follows Musser (1981), 
Measurements arc in millimetres. Unless other- 
wise stated material is housed in the Queensland 
Museum (QMF). 

SYSTEMATICS 



M^'2; QMFIfl82l, partial right maxillary with M*'^: 
QMF23325, partial right maxillary M^--\- QMFiasig. 
left M^ QMF24001. panial left maxillary with M^-^; 
QMF24004, partial left maxillary with M'-^; 
QMF24002, right M*; QMF24003, nghi M^^ 

All from Rackham's Roost Site (19°02'O9" S, 
138*'4r60' E) at Riversleigh, NW Queensland. The 
site represents the remnants of an ancicm cave which 
has largely eroded away leaving the indurated lloor 
sediments exposed. The sediments of the floor contain 
myriad bt:)nes and teeth, mostly fragmented, which are 
interpreted to be megadermatid (Macroderma and 
Megaderma spp.) prey remains (Godthelp, 1988; 
Hand, 1994). Its age is Pliocene on the basis ot a 
macTOpodid similar to Proiemnodon snewini 
Bartholamai (I97S) which species occurs in the early 
to middle Pliocene Bluff Downs Local Fauna f Archer 
&Wade, 1976: Bartholamai, 1978). 



Order RODENTIABowdich, 1821 

Suborder MYOMORFHIA Brandt, 1 855 

Infraordcr MYODONTA Schaub. 1958 

Superfamily MUROIDEA Miller & Gidley, 

1918 

Family MURIDAE Grav, 1821 

Subfamily MURINAE Gray. 1821 

Zyzomys Thomas, 1909 

Zyzomvs rackhami sp. nov. 
(Figs 1-2. Table 1) 

MATERIAL. Holoiype QMF108 18, partial left maxil- 
lary with M^'^. 

ETYMOLOGY. For Alan Rackhani. the discoverer of 
the Rackham's Roost Site. Paratypc QMF 1 08 19. !cti 
maxillary fragment with M* and zygomatic plate. 
Other material QMF23365, partial kfl maxillary with 



DIAGNOSIS Zyzomys rackhami differs from 
other species of the genus by the following com- 
bination of characters: Relatively well-developed 
series of cusps (T3,6,9) particularly T3, relatively 
small proportions of the lingual series csi cusps 
(1,3,7), reduced molar gradient, frequent Tlbis; 
tooth row short and narrow. 

DESCRIPTION. Small lo mediunvsi/ed denUil 
arcade arcuate and concave lingually with the 
internal edge of M^ as the most lingua! point ot* 
the tooth row. M* longer than M-- M-^ small stnd 
relatively reduced. All cusps and cusp complexes 
with marked posteriorly inclined slant except in 
M^ with cusps nearly vertical. Molar overlap 
minimal. 

M'. Relaiively long and naiTow often with u 
prominent anteriorcingulum which formsascmi- 



330 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 1. Zvzomys rackhami sp. nov, 
QMF10818, SEM (stereo pair). 



Rackham's 



circle around the T2,3 complex in the holotype. 
Anterior cingulum with a series of small but 
apparently occlusally functional accessory cusps 
randomly positioned. Tl elliptical with its long 
axis obliquely inclined to the axis of the T2,3 
complex, directed to the rear of the tooth and 
positioned posterior to the base line of the T2,3 
complex. Tl joined to the T2,3 complex only in 
extreme stages of wear and in some specimens 
via a variably sized and shaped Tlbis. Tlbis 



structure in the holotype 
small, becoming more 
prominent with increased 
wear. T2 large and arcuate 
anteriorly with a straight 
posterior edge, more than 
half the width of the ante- 
rior portion of the tooth. T2 
joined to T3 at its buccal 
edge. T3 small and circular, 
with its posterior third be- 
hind the posterior edge of 
T2, almost entirely incor- 
porated into the T2 com- 
plex with wear. T4 
elliptical with its long axis 
inclined obliquely to the 
axis of the T5,6 complex 
and directed posteriorly. T4 
occlusal surface approxi- 
mating Tl in size and shape 
if unworn, larger in worn 
specimens, joined to T5 in 
early stages of wear, nearly 
wholly incorporated in 
older individuals. T5 large, 
with a subangular arcuate 
anterior edge, with posteri- 
orly concave posterior 
edge, with a smaller occlu- 
sal surface area than T2. T6 
strongly attached to T5 
even in early stages of 
wear, with junction of these 
two cusps marked by a cleft 
on the occlusal surface that 
continues anteriorly and 
ventrally as a furrow in the 
enamel indicating that the 
distinction between T5 and 
T6 is never lost. T6 circular, 
smaller than T3, approxi- 
Roost Site, holotype, mately half of T6 behind 
the posterior edge of T5. T7 
larger than T4, elliptical, 
with its long axis inclined obliquely to the axis of 
the T8,9 complex but in reverse to the angles of 
Tl and T4 and as such is directed forwards. T7 
and T4 in very close proximity, forming a single 
complex after moderate to extreme wear. T8 
large, with a nearly circular occlusal surface. T9 
barely discernible, incorporated into the T8,9 
complex at the onset of wear. Slight furrow in the 
anterior surface of the T8,9 complex marking the 
anterolingual edge of T9 (would remain even 



ZYZOMYSRACKHAMI SP. NOV. FROM RtVERSLOGH 



33) 




FIG. 2. Zyzomyx rackliatni sp. nov.. paratype* QMF10821, 
SEM showing zygomatic plate. 



with mixlerate wear). Posterior cingulum (z) ab- 
sent. 

M-. Tl large, lear-shaped, with a long axis 
obliquely inclined lo the anterior edge ot the tooth 
and directed posteriorly. T2 and T3 absent. T4 of 
moderate size, elhptical, posterior to the domi- 
nant T5. T4 joined to T5 in very early stages of 
wear. T5 subiriangular in occlusal outline, wiih 
its anterior most edge of the enamel boundary 
contacting the posterior edge of M' in extreme 
wear. T6 small, circular, joined to T5 with mod- 
crate wear. As in the T6 structure of M', the 
connection associated with a well-defined fur- 
re>w. this cusp always retaining its identity. T7 an 
elliptical cusp oi moderate size, wiih its axis 
running almost parallel to the main axis of the 
tooth, not observed to merge with the T8.9 com- 
plex in M- even with e\treme wear. T8 large, 
nearly circular, with distal edge extending well 
beyond the distal edge of T7, giving the posterior 
region of [he tooth an arcuate shape. T9 lost, with 
a remnant of the furrow that marked the position 
of the lingual side of the cusp. 

M-\ Tl smirlK almost circular. T2 and T3 ab- 
iicnl. T4 small, lear-shaped, joined to T5 after 



little wear, with an axis directed across the 
width of the tooth. T5 small, subtriangular. 
displaced toward the buccal edge of the 
tooth. T6 absent, with only a poorly de lined 
remnant of the lingual furrt)W fonning a 
weak buccal cingulum at the anterior edge. 
T7 moderate in size, shaped as a bisected 
semi-circle. T8 a mirror image of T7. with 
the 2 cusps jomed in early wear. T9 absent. 
M' with 4 roots, 3 well-deveU:»ped, fourth 
small and probably reduced; anterior root 
large, directed forwards, exposed occlus- 
ally; medio-linguai root long, narrow, pv^si- 
tioned under T I and the anterior edge of T4; 
medio-buccal root a remnant of a more sub- 
stantial root, with an alveolus for this root on 
all specimens examined even though there 
is not always a root. Posterior root large, 
nearly as wide as the tooth at its origin, 
directed towards the buccal edge of the max- 
illary. M- with 3 roots; anierolinqua) and 
anterobuccal roots of equal si/e. together 
below the anterior margin of the tooth; pos- 
terior root wide, running obliquely with re- 
spect to the anterior roots, with lingual 
extremity its most posterior point. M* with 
3 roots of equal si/c, forming a triangle with 
a root at each apex. As in M^ there arc 2 
anterior roots (buccal and linqual). 

Occlusal surface of the tooth row concave, 
with the highest points being the anterior 
third of M* and the M^. Anterior palatal vacuitv 
extending distal I y as far the anterior edge of M*^. 
Atiachmeni node for the origin of the superficial 
masselcr large and well defined. Anterior edge of 
the node alligned with the anterior edge of the 
zygomatic plate and just behind the maxil- 
lary/premaxillary suture. Zygomatic with ante- 
rior edge rising vertically and straight, of 
moderate width, with one nutrient foramen dis- 
tally near its base. 

COMPARISON. Although the nearest species in 
size Z. arguruSy Z rackhami is more similar to Z 
pedimculatus because of the relatively cuspidate 
nature of the teeth, the development of the buccal 
series of cusps and the relatively unreduced ap- 
pearance of M^. These similarities are probably 
simplesiomorphics. A nutrient foramen is found 
anterior to the tooth row in all species with the 
exception of Z rackhami and Z maini, in which 
the foramen is on the zygomatic plate. Unlike Z. 
maini however Z rackhami has a longitudinal 
palatal crest as in all other species oiZyzomys. 
Zyzornys rackhami differs from Pseudomys, 



331 



MEXtOIRS OF THE QUEENSLAND MUSEUM 



TABLE I Measurements (mm) of Zv:flw>t mrkfumi 

sp, Tiiiv. L=!eriglh. W-\vitlih. 



SPEC. NO. 






M' 


L \\ 


QMF108i8 


3 3 


4,3 


2 6 


i 3 


QMF10821 


5 6 


4.4 


2.7 


iJb 


QMF23325 


5 3 


4.3 


1.1 


1.7 


QMF108I9 






1-1 


1.7 


QMF23365 


. 


4,2 


_Z6 

2,6 ' 


1.5 


OMF24001 




4,3 


].6 


QMF24a-t2 


1 


2.-; ! i.5 


QMF24i:Ci3 


- 


2.7 1 ],S 


iiMr2-ioa4 


- 


4.2 1 2,6 


i.5 
1.5 


gMF30<J65 




4.2 


2.6 


gMP3cy)63 


- 


4.2 ! 2.6 


1..S 


QMF30062 


- 


- 


2.6 


1..S 


0NtF2430}i 




4.2 


2.6 


15 


QMFI0810 




- 


2.7 


1,6 


QMF30268 


- 


4.3 


26 


1 ,5 


QMF30265 




4,3 2 6 ' 1,5 1 


' 



Mastacomys^ Leporillus, Notontys, Leggadina, 
Rattus^ Meiomys and Uronm in having a well- 
dcvclopcd T7. Z rflrJUitwjiis dislingulshcd Irom 
Pogonofnys by the lack al' u pocitcrior cingulum 
(z) and by smaller and les» pn>nounced buccal 
cusps (T3,6,9). Z. rackhumi differs from 
Hyttrorny^ and Xenmyt in having 3 upper molars. 
Z. rflcWiflm/" differs from Corulurus in having a 
well-developed T3 and Tl isolated from anddis- 
lal 10 the T2,3 complex. Z rackhawi \s removed 
from Mesemhriomys by reiaining a T9 on M', 
close proximity of T7 to the T8.9 complex, re- 
duced molar overlap and less cuspidate nalurc ol 
the molars. 

DISCUSSION. Zyzmiys rackkami is the most 
abundant (marnly isolated molars) rodent in the 
Rackham's Roost deptisu but only a few maxil- 
lary fragments are known. Skulls apparently 
break up more readily than other muiids from the 
deposit. 

Lower jaws of rodents are uncomnion in ihe 
deposit and not weii preserved. No elements of 
the lower dentition of Z rackhami have been 
identified. There is apparently some taphonomic 
pajcess limiting the number of dentanes pre- 
served. This might be caused by some aspect of 
Ihe feeding behaviour ot Macroderma gign.s, the 
presumed predator, or by some physical process 
of sorting within the original cave system. In the 
laiicf case it is possible that we might find a 



concentraiitm of these elements elsewhere in H« 
deposit. Lower jaws and teeth of other mammal 
groups( i e. , imrsupials. bats ) found in the deposit 
appear to be as abundant as their upper counter- 
ports. 

Z rackhami is the only species of Zyzomys 
recovered from the Rackham's Roost Local 
Fauna and is represented by hundreds of speci- 
mens. This is in contrast to modern and 
Pleistocene faunas, in which there arc usually at 
least 2 species present and sometimes, as in tl>e- 
Nouraiangie Rock area, NT., 3 (Kitchener 
0989) reported only 2 but the author has sighted 
2 specimens of Z woodwatdi, CM72(K) and 
CM720O. from the area.) Areas on the east coast 
where only Z. ar^urus is now found appear to 
liavo last a second species, Z woodwardi, only 
recently, as is the case in the Chillagoe area. Z 
pedunculatm occurs in surficial cave deposits at 
Cape Range, Western Australia along with Z 
argums which is still extant locally. In Ihe 
Riversleigh areaZ argurus has been trapped and 
Z w'oodwardi has been recovered from Recent 
owl pellet deposits along the Gregory Ri vei Both 
Z argurus and Z woodwardi have been recov- 
ered from Macroderma gigas prey remains m 
Carrington's Cave on Riversleigh Station The 
deposits in Carrington's C^ive arc as yet undated 
but appear to range from Recent to Pleistocene. 
The presence ot both species m these deposits 
indicates that A/, gigas is capable of taking the 
relatively larger Z woodwardi as prey. PIitx:cnc 
M. gigas from Rackham's Roost were not dilTcr- 
enl in size (Hand, 1994) from the modern popu- 
lation and prey si/e would seem to he an utilikely 
explanation for the absence of a second Zyzamys 
m the Rackham's Roost deposit. 

ACKNOWLEDGEMENTS 

Work at Riversleigh has been supported by the 
Australian Research Council, the Department of 
the EnvironmenL Sport and Tourism, National 
Estate Programme Grants (Queensland), the Aus- 
tralian Geographic Society. ICL the Queensland 
Museum and the University of NSW. Access to 
comparative material was kindly provided by S. 
Van Dyck, J. Calaby, T. Flannery. L. Gibson' D. 
Kitchener and P. Jenkins. SEM photographs were 
taken by .1. Muirhead at the University of NSW. 
I ;im graielul to A. Gillespie and S. Williams who 
have done much of the preparation of the mate- 
rial. Particular thanks are due to the Rackham 
family for their unwavering enthusiasm and as- 
sistance throughout the past years. 1 must thank 



ZYZOMYSRACKHAMl SP. NOV. FROM RIVERSLEIGH 



333 



my colleagues M. Archer and S. Hand for their 
support and advice, and B. Turnbull and A. 
Baynes for constructive criticism of earlier drafts 
of this manuscript. 

LITERATURE CITED 

ARCHER, M. & WADE. M. 1976. Results of the Ray 
Lemley Expedilions, part !. The AIHngham For- 
mation and a new Pliocene vcncbrale fauna from 
northern Queensland. Memoirs of the Queensland 
Museum 17:379-397. 

ARCHER, M., GODTHELP, H., HAND. S.J. & 
MEGIRIAN, D. 1989. Pcriiminar\' overview of 
mammahan diversity, biostraligraphy, corrclaiiun 
and environmental change evidenced by the fossil 
deposits o\' Rivcrsleigh. northwestern Queens- 
land. The Australian Zoologist 25: 29-65. 

ARCHER, M., HAND, S.J. & GODTHELP. H. 1991. 
Riverslcigh. (Reed: Sydney). 

BARTHOLOMAI, A. 1978, The Macropodidae 
(Marsupialia) from ihe AUingham Formation, 
northern Queensland; results of the Ray E. Lemley 
expedition. Part 2. Memoirs of the Queensland 
Museum 18: 127-143. 

BAVERSTOCK, P.R. 1984. Australia's living rodents: 
a restrained explosion. Pp 905-91 3. in Archer, M, 
& Clayton. G. (eds). Vertebrate zoogeography and 
evolution in Australasia. (Hesperian Press:Perth) 

BAVERSTOCK, P.R.. WATTS. C.H.S., ADAMS, M. 
& COLE, S.R. 1981. Gcnetical relationships 



among Australian rodents (Muridae). Australian 
Journal of Zoolog>' 29: 289-303. 
GODTHEI-P. H. 1988. Riversleighscene4: Rackham's 
Roosl — the beginnings of the modem world. Pp. 
81-83. In Hand, S.J. & Archer, M. (eds). The 
antipodean ark. (Angus & Robertson: Sydney). 

1993. Zyzomys rackhami, a new species of murid 
from the Pliocene Rackhams Roost deposit, 
northwestern Queensland. Abstracts, CAVEPS, 
Adelaide. 

1994. The three R's of Riverslcigh: the Rackhams 
Roosl rodents (Placentalia: Rodentia). Abstracts 
Riverslcigh Symposium, UNSW, Sydney. 14. 

HAND. S.J. 1 995. First record of the genus Megadernia 
Geoffroy (Microchiroptera; Megadermalidae) 
from Australia, Palaeovertebraia 24: 48-66. 

KITCHENER, D.J. 1989. Taxonomic apraisal of 
ZKo//o'-^(Rt^eniia, Muridae) with descriptions of 
two new species from the Northern Territory , 
Australia. Records o\ the West Australian Mu- 
seum 14:331-373. 

ML'SSER, G. 1981. The giant rat of Flores iind its 
relatives east of Borneo and Bali. Bulletin of the 
American Museum of Natural History 169: 71- 
175. 

RIDE, W.D.L. 1970. A guide to the native mammals of 
Australia. (Oxford University Press: Melbourne). 

TATE, G.H.H. 1951 .The rodents of Australia and New 
Guinea. Results of the Archbold Expedilions. No. 
65. Bulletin of the American Museum of Natural 
History 97: 189-424. 



NEW MIOCENE LEAF NOSED BATS (MICROCHIROPTERA: HIPPOSIDERlDAE . 
FROM RTVERSLEIGH, NORTITvVTSTERN QUEENSLAND 

SUZANNE HAND 

Hand, S.J. 1997 06 30: New Miocene leaf-nosed bats (Microchiroplera: Hipposideridae) 
from Riversleigh, norihwesiem Queensland. Memoirs of the Qmensiand Museum 41(2): 
335-349. Brisbane, ISSN 0079-8835. 

Two new Australian Teaiar> hipposiderids are described on the basis of skull and dental 
material recovered from Bitesantennary Site, a Miocene cave-fill on the Site D Plateau. 
Riversleigh, nonhwesiem Queensland. The new species are closely related lo Hipposideros 
iBtachtpposiderosjnooraleebus Sigd. Hand & Archer from Rivcrsleigh's Micrositc, and the 
living northern Auslralian Rh'mnmcteris auraniitts (Gray). One species is refened to 
Rhinmictens. the other tentatively referred to Brachipposideros. The subgenus- 
Brachipposideros Sige is raised to generic rank to better reflect relationships of its species. 
D Miocene. Riversleigh. hipposiderids, leaf-nosed hats. 

Suzanne J. Hand, School of Biological Sciencff, Uni\trsLty of New South Wakx, New South 
Wales 2052, Australia: received 4 December 1996. 



Ritcsatitcnnar}' Site, in early Miocene (Archer 
clal., l989;Creaser, 1997) freshwater limestone 
Dn the NE edge of the Site D Plateau at 
Riversleigh (Hand el al., 1989; Archer el a!., 
1989,1994) covers approximately 150m- and 
contains thousands of bal skulls, limb bones and 
snails. Almost all are complete, suggesting 
fossilisaiion at or very near the point of accumu- 
lation. This deposit is interpreied as a cave-fill 
(Hand et al.. 1989) and contains at least 11 
niicrochiropteran species - 10 hipposidcrids and 
a mcgadermatid. At least 4 of the Bitesanlennary 
hipposidcrids are known from many hundreds of 
paitial and complete skulls. Two of the hip- 
posidcrids. which are morphologically similar to 
MIcrosite's Brachipposideros nooraleebiis Sige 
el al., 1982. are described and their phylogenetic 
reluiionships and palacoecology are discussed. 

Skull terminology follows Hand (1993, 1995); 
dental teirninology follows Sige et al. (1982). 
Slratigraphic nomenclature for the Riversleigh 
region follows Archer et al. (1989, 1994; Crcaser 
this volume). The prefix QMF refers to speci- 
mens held in the fossil collections of the Queens- 
land Museum, Brisbane. 



SYSTEMATICS 



Suborder MICROCHIROPTERA Dobson, 1875 
Supcrfamilv RHINOLOPHOIDEA Weber. 

1928 
Family HTPPOSIDERIDAE Miller, 1907 



RhinonicterisGray, 1847 

Rhinonicteri.s ledfordi sp. nov. 
(Figs 1-2, Table 1) 

M-^TERI^^L HolotypeQMF229IO, partial skull with 
RM", L,M". Paratypes QMF229! 1, partial skull with 
RP -Nr and LM-^: QMF22912. maxillary fragmcqt 
with RC -M , QM F22840, rostrum with LC-M-. 
types from early Miocene (System B) Bitesanlennary 
Site. Other material; Bitesanlennary Site: OMF22S3 1\ 
QMF2284]. QMF22842. QMF22845. QMF22854. 
QMF22859, QMF22865, QMF22S71. QMF22S9(), 
QMF22891, OMF22893. QMF22909. White Hunter 
Site (System A): QMF22921, QMF22922. RV Site 
(System B); QMF22930. QMF2293I, QMF22932, 
QMF229i3. Upper Site (System B): QMF22914, 
While I turner, RV and Upper Sites are at>ou( 2km SSW 
of the type locality. 



ETYMOLOGY. For Richard Tcdford. American Mu- 
seum of Natural History who described the first Trrti- 
ary mammals from Ri\erslctgh in 1967. 

ASSOCIATED FALTNA AND TAPHONOMY. 

The cave-fill (Handctal., 1989) at the type locd- 
ily contains thousands of well preserved, almost 
complete bat skulls and limb bones, suggesting 
fossilisaiion at or near the point of accumulation. 
Contact between the fill and older cave wall have 
been idenlificd. The deposit's many freshwater 
snails suggest that the deposili^mal area was open 
to light and under water for some period during 
its history. A travertine floor, including a large 
stalagmite, has been found at the base of die 
deposit. 



336 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 1. A-C, Rhinonictehs tedfordi sp. nov., QMF22910, holoiype, from Bitesanlennary Site, Rivcrsleigh, 
northwestern Queensland. A, dorsal view; B, lateral view; C, ventral view. D-F, Rhinonicteris aurantius, AR 
15400, Klondyke Queens Mine, Marble Bar, Western Australia. D. dorsal view; E. lateral view; F, ventral view 
Scale indicates 5 mm. 



Bitesanlennary Site contains R. tedfordi, ?B. 
watsoni and at least 8 other hipposiderids and a 
megadermatid with rarer frogs, lizards, a bold, a 
stork, swift, peramelids, a dasyurid and a 
bulungamayine macropodid (Archeret al., 1 994). 

In the complex lacustrine White Hunter, Upper 
and RV deposits the vertebrate faunas are much 
more diverse, with the Upper Local Fauna 
(Archer et al. 1994) one of Riversleigh's richest. 



DESCRIPTION. In comparison to Miocene B. 



nooraleebm Sige el al., 1982 and Recent 
Rhmonictehs aurantius (Gray, 1845). 

Skull 10-20% smaller than R. aurantius and 
approximately same size as B. nooraleebus 
(braincase may be slightly longer in R. tedfordi). 
Proportions similar to B. nooraleebus: rostrum 
wide and long with respect to braincase, approx- 
imately 2/3 braincase length, 2/3 maximum (mas- 
toid) width and twice interorbital width. 
Zygomatic width greater than mastoid width. 
Maximum height of the skull dorsal to the glenoid 
process as in R. aurantius. In dorsal view, poste- 



NEW MIOCENE LEAF-NOSED BATS, RIVERSLEIGH 



337 




FIG. 2. Rhinonicteris tedfordi sp. nov., QMF2291 2, paratype, maxillary fragment with C'-M^, from Bilesantenn- 
ary Site, Riversleigh, northwestern Queensland. A, oblique-occlusal view; B-B', occlusal view, stereopairs. 
Scale indicates 1 mm. 



rior margin of the skull quadrate rather than 
rounded as in R. aurantius and B. nooraleebus. 

Rostrum distinctly lower than the braincase, 
more so than in B, nooraleebus but less than in R. 
aurantius. Rostral inflations much more promi- 
nent than in B. nooraleebus and R. aurantius , 
mainly because of the very marked groove lead- 
ing to a deep frontal depression delimited sharply 
by well-developed supraorbital ridges. R. au- 
rantius with inflations better developed, with 
very little development of supraobital ridges, 
with frontal depression and groove between ros- 
tral inflations more limited in depth and extent. 
Infraorbital foramen wholly above M^ as in B. 
nooraleebus, but unlike R. aurantius (above M-' 
^), larger and more rounded than in B. nooralee- 
bus, smaller and slightly more elongated than in 
R. aurantius. Bar of bone forming its dorsal mar- 
gin (anteorbital bar; e.g. Hill 1963) straighter and 
wider anterodorsally than in R. aurantius (being 
roughly the same thickness throughout), (In R. 
aurantius this bone curved, about 3 times as wide 



postero ven trail y as anterodorsally.), more curved 
than in B. nooraleebus, in which it is roughly the 
same thickness throughout and very straight. Zy- 
goma (as in B. nooraleebus and R. aurantius) 
with an enlarged jugal projection occupying 
much of its length, as tall as the level of the lower 
insertion of the anteorbital bar, with slightly con- 
vex posterior margin, with its anterior edge 
posterodorsally directed (rather than vertically). 
Sagittal crest well-developed (but see 
QMF22871), much better developed than in B. 
nooraleebus and different to R. aurantius, with 
maximal height anterior to the middle of the 
braincase level with the posterior zygomatic 
roots, not terminating as abruptly nor in a for- 
wardly curving projection as in R. aurantius, 
extending further anteriorly onto the moderately 
constricted interorbital region, not joining the 
supraorbital ridges as distinctly as in B. nooralee- 
bus, extending posteriorly to the lambdoidal 
crest, rather than attenuating in the interparietal 
region as in R. aurantius. 



338 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE L Skull and dental measurements (mm) of type material. H=holotype; P=paratype; two measurements 
in parentheses in a column indicate (left) and (right), respectively. 





Rhinonicteris tedfordi 


?Brachipposideros watsoni 




QMF22910 
(H) 


QMF22911 
(P) 


QMF229I2 

(P) 


QMF229i5 

(H) 


QMF22828 

(P) 


QMF22916 
(P) 


Greatest skull length (dorsal) 


15.0 


15.4 




14.4 






Rostral length 


5.0 


5.5 




4.6 


?4.2 




Braincase length 


10.0 


9.9 




9.6 


10.1 




Rostral width (at lacrimal) 


5.3 


5.2 




4.5 






Min. interorbital width 


2.5 


2.4 




1.9 






Zygomatic width 


8.5 


8.8 




7.5 






Mastoid width 


(8.9) 






7.5 


8.3 




Rostral height 


4.3 


4.4 




4.0 


4.6 




Braincase height (max.) 


7.9 


7.2 




6.6 


7.3 




Palate length 


1.6 


1.6 




1.7 


1.5 




Palatal width (base of M') 


3.0 


3.2 




2.7 


3.0 




Interperiotic distance 


1.4 


L5 










CI-M3 






5.6 








P4-M3 






5.0 


(4.1) (4.1) 


4.3 




M1-M3 






3.5 


(3.3) (3.2) 


3.5 




c'l 






1.4 








c' W 






1.1 








P^L 




0.9 


1.1 


(0.9) (1.0) 


0.9 


1.1 


P'^W 




l.I 


1.3 


(1.0) (0.9) 


1.3 


1.2 


m'l 




1.4 


1.5 


(1-3)(1.3) 


1.3 


1.4 


m'w 




1.4 


1.4 


(l.Dd.l) 


1.5 


1.3 


M-L 


(l.3)(1.3) 


1.3 


1.3 


(1.2)(1.3) 


1.3 


1.3 


m'w 


(1.4) (1.4) 


1.5 


1.4 


(1.2) (1.2) 


1.5 


1.4 


M^L 




1.0 


0.9 


(0.8) (0.9) 


0.9 




M^W 




1.3 


1.4 


(!.l){l.2) 


1.5 





Premaxillae not known but make a V-shaped 
junction (often stepped) with the maxillae rather 
than a rounded V-shape as in R. aurantius and B. 
nooraleebus. Palate shorter, with posterior mar- 
gin extending to the level of the metacone of M- 
(rather than the anterior face of M^), marked by a 
short postpalatal spine, as in R. aurantius. 
Mesopterygoid fossa narrow anteriorly, necking 
in before broadening posteriorly, more similar to 
R. aurantius than B, nooraleebus in which it is 
broad and rounded anteriorly and of uniform 
width throughout its length. 

Lacrimal foramen much larger than in R. au- 
rantius and larger than in B. nooraleebus. 
Lateroventral fossa broader than in R. aurantius 
and similar in width to B, nooraleebus. Postpala- 
tal and sphenopalatine foramina much larger than 
in R. aurantius or B. nooraleebus (QM F 1 9039 
but not 19040), closely paired, more distant in R. 
aurantius and well separated in B. nooraleebus. 



Anteriordiploic, ethmoidal and cranio-orbital fo- 
ramina fused, larger than in B. nooraleebus, not 
fused and large in R. aurantius, separated from the 
optic foramen by a thick bar (rather than broader 
plate) of bone. Like R. aurantius, palate pierced 
by many foramina, none especially distinctive. 

Sphenorbital bridge relatively broad, not 
greatly constricted posteriorly, with sphenorbital 
fissure well-exposed. Hammular process very 
similar to R. aurantius, with a conspicuous wing 
projecting backwards making up at least half its 
length, with a laterally directed flange of variable 
length (long in QMF22859) posterior to the ham- 
mularprocess, as in R. aurantius and B. nooralee- 
bus. Sphenorbital fissure shorter and broader than 
in R. aurantius; optic foramen more lateral than 
in R. aurantius, with the orbitosphenoid splint 
separating them directed medially rather than 
posteromedially as in R. aurantius and B. 
nooraleebus. 



NEW MIOCENE LEAF-NOSED Bats. RIVERSLEIGH 



33? 



Basisphcnoid shallow, Basioccipilal widih be- 
tween Ihc pcriolics as in R auraniins (perhaps 
slightly narrower), narrower than in B. tUH>nitee- 
tuis. Puslglenoidi'ossa ('/temporal emissary lora- 
nicn) larger than in R. auratuins and fi. 
ni^jraleebus', puslglcnoid process also slightly 
bener developed than in R. aurnniius, and much 
belter than in B fworoleebus, Foramen ovnie 
very laige; a bar of bone separating the I'oranien 
ovale from u ?posicriorly opening fenestra in B. 
noomleebus is absent in R, reJJorJi ;ind R. au- 
ramiusii^ is the fenestra. The lamhdnidal crest is 
bolter developed thun in B. nooraleefni^, and in 
Ibis way more sitnilar to /?. tmrafitius (although 
in the latter this varies intraspccifically e.g. AR 
15400 and M84i6). Unlike R. aurantius, it is 
CYiotinuous across the occipilals m H. tedforcl't 
rather than attenuating at the ?nuchal point. Fora- 
men magnum more dorsally oriented than in B. 
ntKiroleebiis and R. aurantius. with indentation ol 
lis dorsal margin in R annmtiHs lucking in R. 
tt^iifordi and B. nooraievbus. 

Periotic, its orientation aitd its attachnictii to 
surrouttding basicranial elements similar to that 
in R. attranfius and B nooraleebiti 

Upper teeth approximately the sanK si/c in ihc 
3 species, those oi R. aunmtius morchypsidonl. 
Upper incisors unknown. C similar to that in U. 
nocfrateebus in width, length and posterior sec- 
ondary cusp, but with shallower anterolingual 
citigulum. removing its squared appearance (but 
sceQMF22S45). C wider and longer in the tooth 
row than m R. annmUus. P- extruded such that C' 
and P* are in close contact, almost Touching (eg 
QM F22845 ), being closer than in A', nnorab^fbNs 
{although this varies) and at least as close as m R. 
tmramiits. P^ nan'owei than in R. auraniius (es- 
pecially anteriorly), the lingual citigulutn deeper 
than in R. aurantius and similar to R. noomiee- 
bus, and the anterolingual cingular cusp better 
developed than in R. aurantius. M' with 4 roots. 
with heel similar to /?. aurantius and broader than 
in B. noaralevbiis, with a very strong dihedral 
crcsi and thickened poslerolingual cingulum. 
Lingual notch incipient, well-developed in 
QMF22840. M- with 4 roots, evenly spaced, as 
in R. aurantius; B. noomleebus with 3. Its heel 
much weaker than in R, aurantius, similar to B. 
nooraleebus but with the poslprotocrisiu reaching 
Itie base of the meiacone atid with a slight ridge 
(rather than cresl) issumg from its end point (oi 
more anteriorly in worn specimens) and entend- 
ing to the slightly thickened |-«)Sierc^liitgual cin- 
gulum. M^ similar in the 3 taxa. 



COMPARISON. This species differs from the 
Recent R. auraniius in its smaller size, relatively 
shorter braincase (especially in the poslglcnoicJ 
region), flattened rostral inflations, deep gr(X»ve 
between mliations. strong supraorbital ridges,C' 
with less pronounced posjerior ;icee$sory cusp. P* 
relatively naiTow with greater anterobuccal ex- 
tension and M* heol much less expanded. 

From Rivcrslcigh's Brachippositlfros 
ni>orait't'hus Sigc et al., 1982 it differs most con- 
spicuously in its relatively shorter palate with 
posterior medial spine, its. long* slim 
mesopterygoid fossa, well-developed sagittal and 
lambdoid crests, more inflated nasals. C^ without 
deep anterolingual cingulum. W with hiviadcr 
heel and M- with lour roots. 

It differs from fi. omani Sige, J 995, B. sp. cf. B, 
hronssatcmis or "¥<yrvf\ X* froni St Victor La 
Coste (Sig6 ci al.. 1982). B. sp. cf. B. branssaten- 
sis from La Colombiere (Sige et al., 19R2) and B. 
agniUvi Legeivdre. 1982 in M- having 4 routs It 
differs from B, coHon^i^nsis fDcperct, iS92) and 
B. dcchaseauxiSig6. 1968 in the heel oI'M'' not 
being poslcrohiKcally extended and M- invari- 
ably having 4 wots^ It diffe(*s froiU fi. JfrunwaWn^ 
sis (Hugueney, 1965} in its postcrolinguul 
development of the heel of M' - atid generally 
less conspicuous lingual notch separating pru- 
loconc from hoci it) hV'-. 

BrachipposiderosSig^, 19(^8 

?BrachipposJdert)is watsoni sp. nov. 
(Fsgs 3-4. TflbJe I) 

ETYMOLOGY. For Neil Watson in recognition of his 
long associaiian with the UniversUy of NSW. 

MATERIAL- Holoiypc QMF229I5. skull with LP- 
M-^ and RP^-M-^. Paratypcs QMr22828. skull with 
LCLm3 and Rp2-M-\ QMF229 i 6, maxillary fragment 
with LCL\P. Other material OMF22S24, QMF22«26. 
QMF22S33. QMF22S46, OMF22S.S7. QMF22K60. 
QMF2286I. QMF22862. QMF22K70. QMF22>i*M. 
QMF22S^6, OMF228*^8. QMF229I30, QMF22904, 
QMF229n7. .Ml material from the early Mioccoc 
BiteiaTiteniiary Site (discussed above). 

DESCRIPTION. IBrachipposideros mw/jw/ is 
described in comparison with the Miocene R. 
tedfordi sp. nov., fi, nooraleebus Sige et al., 1982 
and Recent R. aurantius (Gray, 1K45). 

Skull appnuimalely 10% shorter and narrower 
than R. tedfordi, 20-30% shorter than R m- 
rantitis, with braincase length similar to B, 
nooraleebus, w\h ^hniliif uver^dl protHKlions to 
fi. neoraleehits and fP. tedfordi, ralhcr Ihan /f. 



340 



MEMOIRS OF THE QUEENSLAND ML'SEUM 



nooraleebits, with rounded posterior 
margin. Sagittal crest lower anteriorly 
than in R. tedfordi and R. auranthis but 
probably slightly taller than in B. 
nooraleehus. As in H. riooraleehus\ 
maximal height ofbraincase more pos- 
terior than in R. tedfordi and R. ou- 
randus, being posterior to the glenoid; 
sagittal crest remaining tall anteriorly 
onto the postorbital region (unlike R. 
aurantius). joining the supraorbital 
ridges fairly distinctly (in QMF22S28 
supraorbital ridges almost develop 
wings or flattened plates like an incipi- 
ent frontal shield), of variable posterior 
extent (in QIVIF229 1 5 attenuating in the 
parietal region, as in B. nooraleehus 
and R. auranthis, but in QMF22828 and 
QMF22843 extending to the 
lambdoidal crest as in R. tedfordi. Zy- 
gomatic width greater than mastoid 
width as in R. tedfordi and unlike R. 
aurantius. 

Rostrum lower than braincase (not as 
low as in R. aurantius and R. tedfordi). 
Rostral inflations similar in proportion 
to R. tedfordi and R. aurantius, more 
distinct than in B, nooraleehus, less dis- 
tinct than in R. tedfordi and R. au- 
rantius. Trough between the inflations 
less pronounced than in R. tedfordi but 
more than in R. aurantius and slightly 
more than in B. nooraleehus. Frontal 
depression shallower than in R. tedfordi 
but deeper than in R. aurantius and B, 
nooraleehus, with an unpaired medial 
frontal foramen. Nasal opening dorso- 
ventrally compressed in anterior view 
compared to that in R. tedfordi and R. 
aurantius; bony nasal septum much 
longer than in R tedfordi and similar to 
R. aurantius; opening of the vomer 
sinus round as in R. aurantius rather 
than slit-like as in R. tedfordi. 

Infraorbital foramen dorsal toM-'^ as 

in R. aurantius, rather than M- as in /?. 

tedfordi and 6. nooraleehus, more 

FIG. 3. ?Brachipposideros watsom sp. nov.. QMF22915, holotype, elongate than in R. tedfordi and R. au- 

from Bitesantennary Site. Riversleigh, northwestern Quccnslimd. fantius Anteorbital bar slim and cencr- 

A, dorsal view; B, lateral view; C-C\ ventral view, stcrcopairs. ally the same width throughout, as in «. 

nooraleehus, sometimes with a flange 
or wing, often slightly curved as in R. 
tedfordi and R. aurantius. Zygomatic arch with 
very enlarged jugai projection (QMF22857) ex- 
tending upwards to at least the level of the upper 




Scale indicates 5 mm. 

aurantius (latter much longer in the poslglenoid 
region), witJi lambdoidal crest generally weaker 
than in R. tedfordi, more like R. aurantius and B. 



NEW MIOCENE LEAF-NOSED BATS, Rl VERSLEIGH 



341 




FIG. 4. ?Brachipposideros watsoni sp. nov., QMF22916, paratype, maxillary fragment with C*-M^, from 
Bitesantennary Site, Riversleigh, northwestern Queensland. A, oblique-occlusal view; B-B\ occlusal view, 
stereopairs. Scale indicates 1 mm. 



insertion of the anteorbital bar, directed slightly 
posterodorsally, with a rounded but narrow apex, 
and slightly convex posterior margin. 

Premaxillae unknown, with a V-shaped junc- 
tion to the maxillae as in R. aurontius, B. 
nooraleebiis and R. tedfordi. Palate extending 
posteriorly to the level of the anterior face of M^ 
as in fi. nooraleebus and R. aurantius, rather than 
the M^ metacone as in R. tedfordi. Bony medial 
palate spine absent, unlike R. aurantius and R. 
tedfordi (variable in B. nooraleebus). 
Mesopterygoid fossa more like that in B. 
nooraleebus than in R. tedfordi or R. aurantius, 
being broad and rounded anteriorly and uniform 
in width throughout its length. 

Laterovenlral fossa narrower than in R. ted- 
fordi, broader than in R. aurantius and similar in 
width to that in B, nooraleebus. Lacrimal and 
postpalatal foramina similar to those in B. 
nooraleebus and smaller than in R. tedfordi. Lac- 
rimal larger than in R. aurantius; postpalatal fo- 
ramen and sphenopalatine similar in size to R. 
aurantius (but proportionately larger), approxi- 
mately equidistant from each other and the three 



interorbital foramina (cranio-orbital, ethmoidal 
and frontal diploic), closely paired with the inter- 
orbital foramina more distant in R. tedfordi, with 
intermediate condition in R. aurantius, with the 
sphenopalatine not 'confluent' (i.e. 2 small fo- 
ramina (QMF19038, QMF19039), and the 3 ap- 
proximately equidistant) in B. nooraleebus. 
Orbitosphenoid splint separating the optic fora- 
men from the sphenorbital fissure, directed 
posteromedially like in B. nooraleebus and R. 
aurantius, rather than medially as in R. tedfordi. 
Sphenorbital bridge slightly more constricted 
posteriorly (posterior to pterygoid processes) 
than in R. tedfordi and R. aurantius. Pterygoid 
wings directed dorsally rather than posteriorly, 
resulting in shorter wings than in R. tedfordi and 
slightly shorter than in R. aurantius (proportion- 
ately). Postglenoid fossa slightly smaller than in 
R. tedfordi, but slightly bigger than in R. au- 
rantius. Postglenoid process similar to R. au- 
rantius and R. tedfordi and better developed than 
in B. nooraleebus. Foramen ovale similar to the 
other taxa; a posteriorly directed fossa relatively 
smaller than in R. tedfordi without bar. Inter- 



341 



MEMOIRS OFTIIE QUEENSLAND MLtSEUM 



(Xrriotic dii^tiince simibr \o that in R. tedfotiii, 
Periotic morphology un<i oiieniuiion itnd ;i«ach- 
mcnt to the basicrrniium similar tn olhcr 3 laxa. 
Foramen magnum similar to that in /?. tedfordi, 
directed more venlrally, us in/?, auraixtms ajid B. 
noomieehus, 

Teeth smaller than in R. aurantius and R. 
tedfordi, jpproximatcly jsume size as in ti 
rtooroleebu^. Upper incisors unknown. C pro- 
portionately ^ihoricr (in Ihc tooth row) than in R. 
tcdfordi <\nd probably B. nooroleebus, inure sim- 
ilar to R. aitraftttus. C cittgulum noT developed 
as in R. redfordi and B. noondeebus^ more like 
in R. iVtranfius: anlerolingual cinguluin follow- 
ing the tooth outline rather than Ihiekcning in the 
anterolingual corner. Posterior secondary cusp 
similar to that in R ledfordi hut perhaps lailer ( in 
buccal view, 1/3 to 1/2 C length rather thiui 1/3 
in B. tworaieebns and R, tedfordi). P' small and 
huccally extruded: C and P* generally not in 
cont;iCt'(but see QM F22907). generally closer, 
but not in conUKl in R. wdjordi and R ountntius 
P* narrower than in R, tutnwtiits and B. noorahe- 
bus, most similar to R. ledfordi. M' has 4 roots. 
With heel longer than in B. rworaleehus\ more 
sitnilar ioR, tedfordi and R. aurantius. M^ with 3 
roots, like B. uounilvobus and unlike R. tedfordi 
and R, aurantius, with heel more expanded ihan 
in R, tedfordi, similar to B. aooraU't^bus, much 
less expanded Ihan in R. autwidus. (Buccal and 
lingual lengths similar in ?R. watsoni and /?. 
tedfordi: buccal length greater than lingual length 
in B. mHfralei'bus,)M^"- crest and cinguhir dcvel 
opment and M^ similm* in the 4 taxa. 

COMPARISON. It differs from R. tcdfordt in its 
.slightly smaller size, shorter mcsopterygoid 
fossa, less anteriorly inOuied braincase. more 
elongate infraorbital foramen, lackot postpalatal 
spme and M^ with three roots. 

It differs from the Keceiil A!, aurantius \\\ its 
smaller size, much less anteriorly inflated brain 
case and pronounced sagittal crest, relatively 
shorter braincase (especially in postglenoid re- 
gion), (laticncd rostral inflations, deeper groove 
between inflations, strong supraorbital hdges. 
less pronitunced acecssi»ry cusp on C'. P-^ larger 
and less extruded from the toothrow, P^ relatively 
nanow vviih greater anterobuccal extension and 
M' heel much less expanded and having three 
roots. 

U differs from B. noofaleebus in itK C lingual 
cingulum being uniformly shallow, its narixjwer 
and shorter P*. more expanded M- heel, sharp rise 
in braincase height above glenoid, position of 



lOfraA^rbital foramen, deep frontal depression and 
more pronounced supraorbital crests. 

it differs from Brachipposiderosb/'anssatensis„ 
B. coKon^ensix and B. dechaseaaxi in M* invari- 
ably having three roots. It differs fiom B. omont 
in its larger si/c and better developed heel in M'. 
It differs from Form X' in its moreexpandcd he'd 
in M'. It differs from B. sp. cf. B. brattssatensis 
in ils posterolingual dcvclopmem of the heels of 
M'- and pronounced crests on the posterior Hank 
ol ihe proiocone. It differs Imm B ag(dtari)n M' 
having four roots. 

COMPARISONS OF THE NEW 
HIPFOSIDERIDS WITH RELATED TAXA 

These new species arc similar in sktill and 
dental morphology to norihcm Australia's livinp 
Rhinonicieris ourantius and Mscrosite's 
Brachipposiderosnooraleebus in proportions of 
the skull, broad rostrum, subparaliel tooth rows. 
palate and zygomatic <irch* crested premaxillae, 
basicranial. periotic and otic morphology, pro- 
nounced accessory cusp on C and little reduced 
upper and lower M3s. 

Sigc et al. (1982) recognised R, minvuius a.s u 
probable descendant of Ihe Austi'allan 
Btavhipposideros lineage. Brachipposideros is 
known trorn the Tertiiiry of Eim>pe, Atabia and 
Ausiral(a{SiEteJ%8;Si'e^etal., l9K2;Legcmlrc, 
1982: Zicgicr, 1993: Sigcet al.. 1995). The new 
Riversleigh species can be compared with Euro- 
pean and North African taxa only on their upper 
dentition because: 1) skull material has ntU been 
described for non-Australian taxa and 2) demur- 
ics cannot be posiiivcly referred to the 
Riversleigh ticva. 

A comhiualion ofdental characters is shared by 
Brachippojsidiros ixui} the new Australian laxa: 
small size, P- between C and F^^ near or on buccal 
margin of toolh row, C' with .secondary cusp, P^ 
slender with respect to other teeth, M' with lour 
roots (loss in some), M- with three roots (ad- 
vanced forms have four), heel of M' sepaiatcil 
from proiocone by a notch (loss secondary) and 
forming a posteriorly directed lobe. M'' heel rel- 
aiivcly weakly developed, primilively, 
postprtHocrista has promincm anterior portion 
and only incipiently developed posierior pari. 
Bnuhipposiderosuooralvebus shares with Euro- 
pean /^mc/i//j/>oi/cATc>5a small lower canine, low 
coronoid process and similar shape of ascending 
ramus (Sigdctal. 1982). 

The 3 Australian Mi^nrenc species differ from 
the early Oligoccne B. omatu (Sige ci al.. 1995) 



NtW MlOCeNELKAF-NOSED BATS, RIVERSLEIGH 



343 



in iheir largei* si/e, more recurved C with beuer 
developed secondary posterior cusp, and ^f' pro- 
lucohc wirh weaker dihedral crest. AdUilionally, 
/?. tedfcmh' diifcTs from B. omiVii in its4-rooicd 
M-. 

Compared with B. sp. cf. £f. branssatensis or 
'Form X' (Sige ci al. 1982) of the French Ime 
Oligoccne(Chaltian). Ihc Australian species have 
P^ smaller and further extruded from the 
Icothiow, P* wiih belter developed anlerulingual 
cjni»ularcusp, P^ wider with respect to M' • (clos- 
est to ?B. waisoni), M'- heels more posieroliiigu- 
aJly developed and posteriorly directed, M-^ heel 
more expanded with dihedral crest and 
poslerolingua! cingulum stronger in Australian 
l^a. M^ size is similar. 

The early Miocene (Lower Aquilanian) French 
species R hnvissatefisis (Hugucncy. I%5) has 
quae different M'- heel development from Aus- 
Ualian species, with heel expansion occurring a( 
ll)c posierolingital corner but directed buccally. 
and having a pronounced lingual notch, n variable 
Chiiracteristic rn Australian taxa. The M- heel is 
hcnei developed ihan in Ausiraliun forms but the 
dihedral crest is more pronounced in Australian 
U\a as is (pnibably ) Ihe po^ilerolingual cmgulum 
C' IS similar to that in 78. walsoni and R. tedford'u 
in which the lingual cingulum is uniform and 
follows Ihc curvature of the tooth, and hence 
unlike /?. noondeehus. P- posiii(»n and si/.e arc 
similur hut ni Ausiralian lorms P'^ is generally 
smaller and mure extruded. The infraorbital fora- 
men occurs dorsal to M- as. in H. redfordi and 5. 
^oondiH'bus. 

M'- heel expansion in the Australian taxa is 
more similar (o Ihat found m the French early 
Miocene (Lower Aquitanian) li. sp. v-f, H, 
branssatvnsis from La Colomhiere. in direction 
of expansion anil strong crest on ihe piKteritx 
flank of the nrotoeone. C' is smaller in si/c and 
the lingual cingulum uniform and even in depth, 
bui wilh similar thickening m its anierolingual 
comer as in R. fcdfordi. The posterior margin of 
I*^ is very curved, ihc anterior margin narrower 
and Uie anierobuccal extension greater than in 7B. 
wofSfttii :ind similar lo /<, noorak'fbtis M', and 
M' '/variably, has 4 roots. 

The r'rench early li» early middle Miocene 
(Upper Aquiianian) B, decliaseauKi Sigt5, I %S is 
larger than the Australian species. The posterior 
Hank on the M'"^ proiocone is simply rounded 
with the dihedral crest poorly dcvcUiped, Ihe heel 
is directed posteriorly to posterobuccally like B. 
hnvissatensi.\\ M'- width is very similar lo that 
nf M^, ^vaWably developed lrngu;il noieh sejmral- 



ing proiocone and heel. P* is narrow wilh respect 

\<y M'"^. possibly smaller than in ?B. yratscmi, its 
anterobuccalcxtensiun much grcaterihan in Aus- 
tralian tana, P" ifs outside Ihe toolhrow, but is 
probably .similar to Australian laxa in si/e ;ind 
position. C has a uniform lingual cingulum as In 
?B. watsoni and R. ledfotdi. 

M'-- heel development in the French e;irly \i> 
early middle Miocene (Buidigalian) Fi, of^tdUm 
Lcgendre, 19S2 is shaipcr than the AusU-alian 
species but the direction of expansion and cicM 
on the proiocone are similar. The posterolingual 
heel cingulum is noi well-developed. The ctcs\ is 
continuous wilh the posterior lingual cingulum in 
H agfiiiari tn M' - the ecloloph (s different the 
buccal edge ts angular rather than rounded a.s. in 
Ihe Australian laxa. P^ appears lo be relalivx'ly 
large and C gracile wilh a uniformly deep lingual 
cingulum like :^H watsani. 

The type species, B. collon^qetisis (Depcrcl. 
1892). from the French early middle Miocene 
(Upper Burdigalian) is similar in si/e to H, 
walsoni and B. noondeehus but P- is less ex- 
truded from the loot h row. M'^ heeb 
posicrobiiccally developed like B. hraN.s.sitft'twis 
and B. dechaseauxi and M- heel better developed 
hui with weakei dihedral crest especially in M^ 
whose proiocone flank is rounded. P^ is relatively 
wide with respect to M''^ vls in the Australian 

SjVCtCN, 

PHYLOCENETIC RFXATIONSHIPS 

On denial characK^s. iIk new Australian spc 
cics arc more similar lo each other and to B, 
noondetfbns Ihan to non-Ausiralian taxa. Sigi5 et 
aJ. (1982, figs K-9) found that that compared to 
liuropean species, the dental structure of B. 
noorakebiis was more advanced Ihan Aquitanian 
forms and as advanced as Burdigalian species. 
The Chatiiiin 'Form X' was considered close Ui 
the ba.se wS ihe European radiation, with B. 
branssatemis close to the group lhat gave rise lo 
the B. coUongcnsis and B. dechaseanxi lineages 
and H, sp cf . B. hranssaffnsis closer t\ ) B. a^nlluri 
and B nooraleehns, Apomoiphics shai'ed by H. 
sp. cl. B. branssatcnsis. B.aenHarf ;ind B. 
noondi'ehn.s included heel of M ' " separated from 
Ihe proiocone by a slight lingual notch and l*ccls 
developed posierolingually and directed poswl- 
orly. Bra<hi{)fu}.\id('rosugtdldri and B. noomke- 
bus share I urther reduction oj P- so Ihal C' and f-*^ 
are close and someiimes in contuci. P^ relatively 
larger and M^^ pratoconc with pronounced dihe- 
dral CfVM 



344 



MEMOIRS OF THE QUEENSLAND MUSEUM 



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FIG. 5. Phylogenetic hypotheses of hipposiderid relationships presented by Hand & Kirsch (in press) resulting 
from analyses conducted on 40 taxa and 59 dental, cranial and skeletal characters: A, Strict consensus of 4 
PAUP trees, all unordered characters (CI=0.25; 87.5% resolution); B, Strict consensus of 8 PAUP trees, some 
ordered characters (Cl=0.23; 82.5% resolution). C, Hennig86 Nelson consensus, unordered characters. See 
Hand & Kirsch (in press) for characters and character states. 



NEW MIOCENE LEAF-NOSED BATS, RIVERSLEIGH 



345 



TABLE 2. Distribution of character-stales used in a 
phylogenelic analysis of relationships among 
Brachipposideros and Rhinonicieris species, and re- 
lated taxa. based on dental characters only. 0=inter- 
preted plesiomorphic condition, N3=apomorphic 
states, ?=missing data or character does not apply. 



Tax on 


Character states 


B. omani 


7177 7777 oi 71 1 


B. brossolensis 


1007 001101011 


B. sp. cf B. branssatensis 


1000 001101 oil 


B. collongensis 


lOUOlOl 11011 


B. aguilari 


1012112201 Oil 


B. dechaseauxi 


lOlOOOIl UOII 


B. nooraleebus 


1002 0022 01011 


?B. watsoni 


10010022 01011 


R.. fedfordi 


10120022 11 Oil 


R. attrantius 


I0120122120I1 


Hipposideros aier 


0201 7022 01 111 


Amhopsomaius 


0100 7021 '>0 100 


Ancestor 


0000 000000 000 



Characters: 

1 : Height of ascending ramus of dentary: 0=tall, 1 =low 
2: C^ accessory cusp: 0=present, l=poorly developed, 

I =absent 
3: P" extrusion: 0= extruded but still separating C' and 

P^, l=C' and P** in contact or nearly so 
4: P^ width wrl other cheekteeth; O=narrow, l=mc- 

dium, 2=wide 
5:M' no. of roots: 0=4, 1=3 

6: M' heel development/length: O=moderate, l=slrong 
7: M' heel direction: O=none, l=posterobuccal, 

2=posterolingual 
8: M' lingual notch: 0=absent, l=inconspicuous, 

2=conspicuous 
9: M^ dihedral crest: 0=absent, l=weak/medium, 

2=strong 

IO:M" number of roots: 0=3, 1=4 
ILM-^ heel length/development: O=none, l=slight, 

2=great 
12:M^ heel direction: O=none. l=posterobuccal, 

2=posterolingual 



13:M- dihedral crest: 0=absent, 
2=strong 



:weak/medium. 



The new Bitesantennary species also share 
these apparent aponioiphies and are assigned to 
that clade. Although tedfordi shares with the B. 
branssatensis, B. coll on gens is and B. 
dechaseauxi lineages a fourth root on M-^ it does 
not share the distinctive posterobuccally ex- 
panded heels on M'-^. In this case a four-rooted 
M- is interpreted to be homoplastic; it occurs also 
in R. aurantius. 

A phylogenelic analysis of the interrelation- 
ships of 12 hipposiderid species including 10 



species of Brachipposideros and an hypothetical 
ancestor, based only only dental features (13 
characters) (Table 2) and using the clustering 
program PAUP 3.1.1 (Swofford 1993), was un- 
able to resolve relationships within the group 
(percent resolution of trees 18.2%). Tree resolu- 
tion did not improve when the most poorly known 
species, B. omani , was removed, nor if character 
states were ordered. However, majority rule trees 
(50%) did show the European B. branssatensis, 
B. dechaseauxi and B. sp. cf. B. branssatensis 
(Form X) clustering in 67% of trees, as did B. 
collongenis, B. aguilari, R. tedfordi and R. au- 
rantius. Hand & Kisch (in press) found in their 
phylogenetic analyses of 37 hipposiderids that 
dental features (20 characters) were not sufficient 
to interpret relationships among genera and spe- 
cies groups of the Hipposideridae. They found 
that resolution of trees was less than 33% when 
dental features only were used, compared with 
87.5% resolution with a combined data set of 
cranial, dental and skeletal characters. 

Brachipposideros Sige, 1968 was erected as a 
subgenus of ///ppos/J^To^ Gray, 1831. However, 
probable patristic relationships between 
Brachipposideros and Rhinonicteris, indicate 
that the evolutionary relationships of these taxa 
are not adequately reflected by current taxonomy. 
Hand & Kirsch (in press) found Hipposideros to 
be almost certainly paraphyletic, as did 
Bogdanowicz & Owen (in press). Huguency 
(1965), Sige (1968) and Legendre (1982) all sug- 
gested that Hipposideros was paraphyletic. 

Hand & Kirsch (in press) also found that B. 
nooraleebus was more closely related to 
Rhinonicteris (aurantius and fedfordi), and pos- 
sibly other Australian Miocene hipposiderids 
than to Hipposideros (Fig. 5A-C). Because that 
analysis was based on cranial as well as dental 
characters, European Brachipposideros taxa 
could not be included, and precise relationships 
between non-Australian and Australian 
Brachipposoideros species remain unclear. 

Relationships between Brachipposideros, 
Rhinonicteris and other Australian Miocene 
hipposiderids were not completely resolved in the 
analyses by Hand & Kirsch (in press). However, 
in all trees nooraleebus occurred as the 
plesiomorphic sister-species to a clade consisting 
of, or containing, aurantius and tedfordi. In .some 
trees, aurantius and tedfordi formed part of a 
broader group of Australian Miocene 
hipposiderids including Xenorhinos and 
Riversleigh (Fig. 5B-C). 

When watsoni was included in PAUP analyses 



346 



MEMOIRS OF THE QUEENSLAND MUSEUM 



A 



c 



fE 



Anthops ornatus 
Xenorhinos halli 
Riversleigha wiltiamsi 
Triaenops persicus 
Rhinonicteris aurantius 
Rhinonictehs tedfordi 
Brachipposideros nooraleebus 
?Brachipposideros watsoni 
Cloeotis percivali 
Coelops frithi 
Aselliscus thcuspidatus 
Rhinolophus megaphyllus 
Rhinolophus affinis 
Rhinolophus simulator 
Ancestor 



A 



rf^ 



-c 



rC 



Anthops ornatus 
Xenorhinos halli 
Riversleigha williamsi 
Triaenops persicus 
Rhinonictehs aurantius 
Rhinonicteris tedfordi 
Brachipposideros nooraleebus 
?Brachipposideros watsoni 
Cloeotis percivali 
Coelops frithi 
Aselliscus thcuspidatus 
Rhinolophus megaphyllus 
Rhinolophus affinis 
Rhinolophus simulator 
Ancestor 




Rhinonicteris aurantius 
Rhinonictehs tedfordi 
Brachipposideros nooraleebus 
?Brachipposideros watsoni 



100 



67 



C 



Rhinonicteris aurantius *- 

Rhinonicteris tedfordi 
Brachipposideros nooraleebus 
?Brachipposideros watsoni 



FIG. 6. Phylogenetic hypotheses of relationships of 40 hipposiderids plus ? Brachipposideros watsoni resulting 
from PAUP analyses conducted on 59 characters (Hand & Kirsch, in press). A, Strict consensus of 44 trees 
(Cl=0.24; 55% percent resolution), all unordered; B, 50% majority rule tree of 6A; C, D, % support (majority 
rule) for clustering of Rhinonicteris and Brachipposideros taxa, trees based on unordered and ordered characters 
respectively. 



of the same taxa and characters used by Hand & 
Kirsch (in press), resolution of relationships be- 
tween hipposiderid taxa fell (from over 82% to 
less than 60% in all analyses). Relationships 
among crown groups (i.e. Hippos ideros, Asellia, 
Palaeophyllophora and Pseudorhinolophus) re- 
mained unchanged from those shown in Fig. 5 
(indicated by broken line in Fig. 6A-B), but res- 
olution at the base of the trees (e.g., among 
Brachipposideros, Rhinonicteris, Coelops and 
Cloeotis) decreased markedly (cf. Figs 5A and 
6A). Majority rule trees (50%) clustered species 
of Rhinonicteris and Brachipposideros (e.g., Fig. 
6B), but with little consensus on relationships 
between watsoni , nooraleebus and an aurantius- 
tedfordi dside (Fig. 6C-D). 

On the basis of all analyses (Hand & Kirsch in 
press and herein), watsoni and tedfordi are as- 
signed to a clade also containing B. nooraleebus 
and R. aurantius. However, the interrelationships 



between these taxa is not as clear. Skull morphol- 
ogy of nooraleebus (e.g., its poorly developed 
sagittal crest, shallow frontal depression and 
poorly inflated nasals) is less derived than thai of 
watsoni, but its dentition (e.g., large P^) would 
exclude it from being a structural ancestor to 
watsoni. Here, watsoni has been tentatively as- 
signed to Brachipposideros, and tedfordi to 
Rhinonicteris. Brachipposideros nooraleebus is 
known from fragmentary material with key fea- 
tures of the sphenorbital bridge area lacking 
(Hand, 1993, fig.l). However, both nooraleebus 
and watson lack a number of apparent apomorph- 
ies shared by tedfordi and aurantius, including an 
anteriorly vaulted braincase and low but conspic- 
uously inflated rostrum, a round infraorbital fora- 
men bordered by a curved anteorbital bar, a 
postpalatal spine, a narrow, scalloped 
mesopterygoid fossa, a poorly (posteriorly) con- 
stricted sphenorbital bridge with long (an- 



NEW MIOCENE LEAF-NOSED BATS, RIVERSLEIGH 



347 




difficult to determine on dental morphol- 
ogy alone. Until further information be- 
comes available, all non-Australian and the 
least derived Australian laxa (i.e., those 
lacking obvious synapomorphies for 
Rhinonicteris) are referred to 
Brachipposideros as perhaps the simplest, 
if not entirely accurate, reflection of the 
group's evolutionary relationships. 

'1 Brachipposideros sp. (Fig. 7), a maxil- 
lary fragment of a Miocene hipposiderid 
from Riversleigh's Upper Site, preserves 
M' and M- which are strikingly similar to 
those of the B. collongensis and B. 
branssatensisVmesiges, particularly in their 
poslcrobuccaily-directed heel develop- 
ment which is quite unlike any other known 
Riversleigh hipposiderid. 
The Bitesantennary Site is a Miocene 
FIG. 7. ?firac/n/7/?oi7Weraysp.,QMF22917, maxillary fragment cave-fill in which '?B. watsoni and R. 
wiihM'^fromUpperSite, Riversleigh, northwestern Queens- ft'^yt^rrf/ occur with at least 8 other 
laiKl.A-A\stereopairs,oblique-occlusal view. Scale indicates hippositierids, 5 of which are yet to be 

described. Five of the 10 Bitesantennary 
hipposiderids, including ?B. warsoni and R. 
tedfordi, are well represented, each by tens or 
hundreds of complete skulls; the other 5 
hipposiderids, and a megadermatid (cf. 
Macroderma godthelpi), arc represented by 
fewer, more fragmentary specimens. The gener- 
ally very fine preservation of the remains (often 
with periotics in situ) suggests that fossiiisation 
occurred quickly with little transport, probably in 
still water rather than guano (in which biodegra- 
dation would be expected). Few juvenile bats are 
among among the thousands represented, sug- 
gesting that this cave (or part thereof) was not 
used as a maternity roost. 

By analogy with modern bat communities, the 
high diversity of hipposiderids in the 
Bitesantennary deposit suggests wami, humid 
conditions in the cave, and probably outside it. In 
Europe, appearance of Brachipposideros in the 
fossil record coincides with a period of steadily 
increasing temperature and their disappearance 
probably correlates with the climatic deteriora- 
tion across Europe in the later Pliocene (Aguilar 
et al., in press). In Australia, 6 hipposiderids are 
restricted to northern tropical areas. Rhinonicteris 
aurantius roosts in very warm, humid caves in 
colonies of 20 to several thousand individuals 
from NW Queensland to NW WA. It emerges at 
dusk to feed, mostly on moths but also on beetles, 
shield-bugs, parasitic wasps, ants, chafers and 
weevils (Jolly & Hand, 1995). Although R. au- 
rantius and the Miocene Rhinonicteris tedfordi 



mm. 



leroposteriorly) pterygoid wings, and M- with 
four (rather than three) roots. Fewer apomorphies 
appear to be shared between watsoni and 
nooraleehus, but potentially include the posterior 
extension of the supraobital crest and elongated 
infraorbital foramen. 

Recent R. aurantius can be distinguished from 
the Miocene R. tedfordi by its larger size, rela- 
tively longer braincase (especially in postglenoid 
region), little or no groove between inflations, 
weaker supraorbital ridges, more expanded heel 
on M-, more pronounced accessory cusp on C', 
P* relatively wide with little anterobuccal exten- 
sion, and P- small and further extruded. In 
Riversleigh' s Pliocene Rackham ' s Roost deposit, 
an early population of R. aurantius occurs syn- 
topically with other as yet undescribed 
Rhinonicteris and/or Brachipposideros species, 
and today R. aurantius is still found in the general 
area. 

DISCUSSION 

I raise Brachipposideros Sige, 1 968 from sub- 
generic to generic level. Tentatively, it would 
include non-Australian species fi. branssatensis, 
B. collongensis, B. dechaseauxi , B. omani. B. 
aguilari and B. sp. cf, B. branssatensis) as well 
the Australian Miocene species, nooraleehus and 
watsoni. Although this may be a paraphyletic 
group, evidence is conflicting and relationships 
between Australian and non-Australian taxa arc 



348 



MEMOIRS OF THE QUEENSLAND MUSEUM 



are closely related and similar in many skull 
features, the extinct species lacks the forward- 
projecting development of the sagittal crest that 
characterises R. aurantius, and it is unclear 
whether or not they could be described as ecolog- 
ical vicars. 

Hipposiderid bats promise to be useful 
biostratigraphic indicators in the limestones at 
Riversleigh. They are the most common bats in 
Riversleigh's Miocene deposits, the best pre- 
served, and, with megadermatids, currently the 
best understood in terms of their phylogenetic 
relationships as well as their morphological vari- 
ability (Sige et a!., 1982; Hand, 1993, 1995, 
1997). Brachipposideros nooraleebus is known 
only from Microsite and ?B. watsoni only from 
Bitesantennary Site. Rhinonicteris tedfordi, how- 
ever, is known from Bitesantennary Site in the 
Verdon Creek Sequence, RV and Upper Sites on 
Godthelp's Hill, and White Hunter Site on Hal's 
Hill. None of the species described herein has 
been recorded from System C sites (Archer et al., 
1989, 1994; Creaser, this volume), but close rel- 
atives (?descendanls) occur at sites such as 
Gotham City and Dome North Sites suggesting 
that lineages may be identified within the 
Riversleigh limestone sequence. 

ACKNOWLEDGEMENTS 

Work at Riversleigh has been supported by the 
Australian Research Council, the Department of 
the Environment, Sport and Territories, National 
Estate Programme Grants (Queensland), Queens- 
land National Parks and Wildlife Service, the 
Australian Geographic Society, the Linnean So- 
ciety of New South Wales, ICI, the Queensland 
Museum and the University of NSW. I thank 
Michael Archer, Henk Godthelp and Bernard 
Sige for their help and encouragement and two 
referees for constructive criticism of this paper. 
Skull photographs are by Ross Arnett and Robyn 
Murphy, University of NSW. 

LITERATURE CITED 

AGUILAR, J.-J., LEGENDRE, S., MICHAUX, J., & 
MONTUIRE, S. (in press). Pliocene mammals 
and climatic reconstruction in the western Medi- 
terranean area. American Association of Strati- 
graphic Palynologisis, Contribution Series. 

ARCHER, M., GODTHELP, H., HAND, S.J. & 
MEGIRIAN, D. 1989. Fossil mammals of 
Riversleigh, northwestern Queensland: prelimi- 
nary overview of biostratigraphy, correlation and 



environmental change. The Australian Zoologist 
25: 35-69. 

ARCHER, M., HAND, S.I & GODTHELP, H. 1994. 
Riversleigh. The story of the animals of 
Australia's ancient inland rainforests. 2nd Ed. 
(Reed: Sydney). 

BOGDANOWICZ, W., & OWEN, R.D. (in press). In 
the Minotaur's labyrinth: a phylogeny for the 
Hipposideridae. In Kunz, T. &Racey, P (eds). 
Proceedings of the 10th international Bat Re- 
search Conference, Boston. (Smithsonian Institu- 
tion: Washington). 

CREASER, P 1997. Oligocene-Miocene sediments of 
Riversleigh: the potential significance of topogra- 
phy. Memoirs of the Queensland Museum 41: 
303-314. 

HAND, S.J. 1993. First skull of a species of 
Hipposideros (Brachipposideros) ( Mi c ro- 
chiroptera: Hipposideridae), from Australian 
Miocene sediments. Memoirs of ihe Queensland 
Museum 31: 179-192. 
1995. Hipposideros bernardsigei, a new 
hipposiderid (Microchiroptera) from the 
Miocene of Australia and a reconsideration of the 
monophyly of related species groups. Munchner 
Geowissenschaftliche Abhandlungen. 
1997. Miophyllorhina riversleighensis gen. et sp. 
nov., a new Miocene leaf-nosed bat 
(Microchiroptera: Hipposideridae) from Queens- 
land. Memoirs of the Queensland Museum 
41:351-354. 

HAND, S.J., ARCHER, M. & GODTHELP H. 1989. 
A fossil bat-rich, Oligo-Miocene cave-fill from 
Riversleigh Station, northwestern Queensland. 
Conference on Australasian Vertebrate Evolution, 
Palaeontology and Systematics, Sydney, March 
1989, Abstracts: 7. 

HAND, S.J., & KIRSCH, J.A.W. (m press). A southern 
orgin for the Hipposideridae (Microchiroptera)? 
Evidence from the Australian fossil record. In 
Kunz, T. & Racey, P. (eds) 'Proceedings of the 
10th International Bat Research Conference, 
Boston'. (Smithsonian Institution: Washington). 

HUGUENEY, M. 1965. Les chiropt^res du Stampien 
superieur de Coderei-Branssat. Documents du 
Laboratoire geologique de la Faculte des Sciences 
de Lyon 9: 97-127. 

JOLLY, S. & HAND, S.J. 1995. Orange Leafnosed-bat 
Rhinonicteris aurantius. Pp. 464-465. In Strahan, 
R. (ed.), The complete book of Australian mam- 
mals. 2nd Ed. (Reed: Sydney) 

LEGENDRE, S. 1982. Hipposideridae (Mammalia: 

Chiroptera) from the Mediterranean Middle and 

Late Neogene and evolution of the genera 

Hipposideros and Asellia. Journal of Vertebrate 

^ Paleontology 2: 386-399. 

SIGE, B. 1968. Les chiropteres du Miocene inferieurde 
Bouzigues. I. Etude systematique. Pal- 
aeovertebrala 1: 65-133. 

SIGE, B., HAND, S.J. & ARCHER, M. 1982 An Aus- 
tralian Miocene Brachipposideros (Mammalia, 



NEW MIOCENE LEAF-NOSED BATS, RIVERSLEIGH 



349 



Chiroptera) related to Miocene representatives 
from France. Palaeovertebrata 12: 149-71. 

SIGE, B., THOMAS, H., SEN, S., GHEERBRANDT, 
E., ROGER, J. & AL- SULAIMANI, Z. 1994. Les 
chiropteres de Taqah (Oligocene inferieur, Sul- 
tanat d'Oman). Premier inventaire systematique. 
Munchner Geowissenschaftliche Abhandlungen 
A, 26: 35-48. 

SWOFFORD, D.L. 1993. PAUP: Phylogenetic analy- 



sis using parsimony. Version 3. 1 . (Computer pro- 
gram distributed by the Illinois Natural History 
Survey, Champaign, Illinois). 
ZIEGLER, R. 1993. Die Chiroptera (Mammalia) aus 
dem Untermiozan von Wintershof-West bei 
Eichstatt (Bayem). Mitteilungen der Baycrischen 
Staatssammlung fur Palaontologie und 
Hislorische Geologie 33: 1 19-154. 



MIOPHVLLORHINA RIVERSLEIGHENSIS GEN. ET SP. NOV., A MIOCENE LEAF- 
NOSED BAT (MICROCHIROPTERA: HIPPOSIDERIDAE) FROM RIVERSLEIGH, 

QUEENSLAND 

SUZANNE HAND 

Hand, S.J. 1 997 06 30: Miophyllorbina rnersleighensi.s gen. ei sp. tiov.. a Miocene leaf-iioscd 
bal (Microchiroplera: Hipposideridai.^) from River.sleigh. Queensland. Memoirs of the 
Queemland Muaeum 41(2): 351-354. Brisbane. ISSN 0079-8835. 

A new Australian Ternary hipposiderid is described on the basi.s of a maxillary fragmcm 
froni RV Site, on Godthelp flill, Riversleigh. northwestern Queensland. Miophyllorhma 
menlfi^hensis £^en. ei .sp. nov. isdislinguishcd from all other hipposiderids in its loss of P", 
releniion of a large M^. and P* longer than wide wiili well-developed anierolingual cingular 
cusp. Its phylogenelie relationships to other hipposiderids are obscure. □ Miocene, 
Riversleigh Australia, hipposiderid, leaf-nosed but 

Suzanne J. Hand. School of Biological Science, University of New South Wales, New South 
Wales 2052. Australia: received 4 December 1996, 



Tertiary deposits in the Riversleigh World Heri- 
tage Fossil Silc, Lawn Hill National Park, NW 
Queensland are rich in Old World bats of the 
Hipposideridae (Sigc ci al., 1982; Hand, 1993, 
1995, 1997). They comprise the majority of bats 
in all Riversleigh Oligocene-Miocene sites, with 
as many as 8 species occurring synlopically in 
dept>siis such as Upper Site (Archer ct al., 1994). 
Many species are known fron) complete or nearly 
complete skulls as well as disarticulated but com- 
plete postcranial material. Several hipposiderid 
genera or subgenera are represented and others 
await description. 

A fragment of a maxilla from RV Site (Archer 
et al., 1989. 1994: Crcaser, 1997) represents a 
new hipposiderid genus distinguished by a 
unique combination of features, h has ntit been 
identified from adjacent, possibly contemporane- 
ous deposits, such as the better-sampled RSO Site 
(Greaser. 1997). Other vertebrates from RV Site 
are generally fragmentary and of small to me- 
dium-sized animals. They include a skink, 
madtsoiid. small crocodile, chelid, pcramelid» 
pscudocheirid and the phascolarctid Nimiakoalo 
grcystanesi (Black & Archer 1997). Other bats 
irom the site include Rhinonicien's tcdfordi 
(Hand. 1997) which also occurs at adjacent sites, 
a vespcrlilionid possibly Leuconoe and a 
moiossid. 

Dental lerminology follows Sige et al. (1982). 
Specimens held in the fossil collections of the 
Queensland Museum (QMF), Brisbane. 

SYSTEMATICS 

Suborder MIGROGHIROPTERA Dobson, 1875 



Superfamily RHINOLOPHOIDEA Weber. 

1928 
Family HIPPOSIDERIDAE Miller. 1907 

Miophyllorhina gen. nov. 

TYPE SPECIES. Miophyllorhina riversleighensiH isp. 

nov. 

El YMOLOGY. Greek phyllo. leaf and rhino, nose; 
Mio' refers to the interpreted Miocene age. 

DIAGNOSIS. P^ lost; P^ longer than wide, with 
deep lingual cingulum and weli-devcloped an- 
terolinguaIcinguIarcusp;largeM-^.as wideasM- 
and with premciaerista 3/4 paracrista length. 

Miophvllorhina riversleighensis sp. nov. 

(Fig. 1) 

MATERIAL. Holot^pe QMF30566, a left maxilla 
fragment \s ilh P^. M^"-^ from Early Miocene tufa al RV 
Site in System B on Godihelp's Hill. Riversleigh 
(Archer etal.. 1989. 1994; Megirian. 3992: Greaser. 
1997k RV Site is perhaps slightly younger than the 
South Australian Kutjamarpu Local Fauna 
(WoodbumeeiaL, 1985). ' 

ETYMOLOGY. For the Riversleigh World Heritage 

Fossil Site. 

DIAGNOSIS. As for genus. 

DESCRIPTION. Teeth worn but not broken. Al- 
veolus for C' indicating this tt)oth wider and 
longer than P^. P- lost, with no sign of an alveolus 
for this tooth either within or extruded (lingually 
or buccally) from the toothrow. P^ longer than 



352 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. !. Miophyllorhina riversleighens'is gen. el sp. nov. from RV Site, Godlhelp's Hill, QMF30566, holotype. 
A, antero-occlusal view showing lack of alveolus for P~. B-B',stereopair, oblique occlusal view. Scale = 1mm. 



wide, narrower particularly anteriorly than M-^ 
and as long. Lingual cingulum deep, well-devel- 
oped; anterolingual cingular cusp well-devel- 
oped. M^ and M- with 4 evenly-spaced roots. M^ 
with a posteriorly-directed, small but conspicu- 
ous heel. Protofossa probably open posteriorly 
but with wear the postprotocrista reaching the 
base of the metacone. Postprotocrista with a 
slight ridge or crest issuing from what was prob- 
ably its end point (more anteriorly in this worn 
specimen) and extending to the thickened 
posterolingual cingulum. M^ not greatly reduced, 
as wide as M-, with premetacrista 3/4 paracrista 
length. 

MEASUREMENTS(mm). Holotype QMF 
30566:- p4-M-^L=3.77; C' (alveolus)-MM.71; 
M--M-^=1.77;P^L=0.88; P^l=0.81; M- L=L18; 
U- 1= 1 .42; M^ L=0.82; M^ 1=1 .40. 

COMPARISONS. In lacking a P-. this species 
differs from all other Riversleigh hipposiderids, 
namely, Brachipposideros, Rhinonicteris, 
Xenorhinos, Riversleigha and Hipposideros 
bernardsigei from the Oligocene-Miocene 
Neville's Garden Site (Hand, 1995) and H. sp. 
from the Pliocene Rackham's Roost Site (//. bi- 
color group). The lack of P- characterises living 
and extinct Asellia, Cloeotls and H. (Syn- 



desmotis), and P- is very reduced or lacking in 
some members of the H. cyclops group (including 
the only fossil taxon H. bernardsigei). It is rarely 
absent in other extant species of Hipposideros 
(i.e., possibly H. sabanus). However, in these 
cases: 1) M^ is also very reduced (\.Q.,Asellia and 
Hipposideros)', or 2) P^ is large and anteriorly 
very wide, with the anterolingual cingular cusp 
reduced, absent or located near the buccal margin 
of the tooth (i.e., Cloeotis percivali and some of 
the H. cyclops group, including H. semoni and H. 
stenotis)\ or 3) M-^ is reduced and P"^ is wider than 
long (i.e., Syndesmotis megalotis). P- is retained 
in all other hipposiderids, i.e., Tertiary Pal- 
aeophyllophora, Pseudorhinolophus and Vayiat- 
sia, and extant Coelops, Paracoelops, Triaenops, 
Anthops and Aselliscus. It is also retained in the 
Rhinoiophidae, the immediate sister-group of the 
Hipposideridae. 

DISCUSSION. On available material it is not 
possible to determine the relationships of this 
species to other members of this family. Hand & 
Kirsch (in press) found that dental features alone 
are not sufficient to interpret relationships within 
the Hipposideridae. 

In its dentition M. riversleighensis exhibits a 
mixture of what appear to be plesiomorphic and 
apomorphic features. For example, the loss of P- 



NEVV UEAF-NOSED BATFROM RIVERSLEIGH 



353 



IS pnibably a derived fealure for hipposiderids. 
iiulependently acquired in a number of separate 
lineages. The anteriorly-narrow P* wilh promi- 
nent anlerolingual cingular cusp is more dilTiculi 
m inierpret. bui is possibly plcsiomorphic among 
hipposiderids (Hand, 1995. Hand &. Kirsch. in 
press). The large M^ on the other hand i^prohahly 
plcsiomorphic among hipposidcrids ic.g., 
Hipposideros, Cloeoiix pvrci vali ix%\A I h e 
lirachipposideroS'Rhinonictefis group) but may 
he secondarily derived in other groups (e.g., ihe 
H. Cyclops group; Hand. 1995). 

All hipposidcrids known in Rlvcrslcigh's 
Oligo-Mioccne and Plicxrenc sediments retain a 
P^ as do most living Au>iraliiiin hipposiderids (N. 
iiunmiats, H. afer.H. ceninus, H. semoni and//. 
(hodema). Only living H. srenotis of NW Aus- 
tralia, a highly specialised member of the H. 
tydop.s group (Hand. 3995). lacks d P-. Af. 
riversleighensis may he related to the Bntc- 
hipposidcros-Hliinwucterh group, sharing a sim- 
ilar P* and large M\ 

M nverslcij^hensis, m lacking a P^, may repre- 
sent an aberrant Bravhipposklcros. However, no 
other species of the Brach ipposideros ■ 
Hhinonicteris group shows this abnonnality de- 
spnc hundreds of specimens bemg available. The 
other vcrj' distinctive hat laxa in RV Site (a 
vcspcrtilionid and a molossid) lend wcighi to Ihe 
argument that Miophyllorhmah also a di^vtinctivc 
but poorly represented laxon. 

Hand & Kirsch (in press) suggested it close 
relationship between Brachipposkienn and 
Cheoiis, early autapomorphically speciaJived 
branches of the hipposidend radiation. Hill 
(1982) grouped Rhinonicieris, Cloeous and 
Trwenops". Koopman (1994) referred them t(» a 
separate subtrihe, the Rhinonycicritia. Perhaps 
XSiophyltorhina is part ol" this larger group of 
relatively plcsiomorphic hipiK)siderids, CUftfotis 
shai-es wiih Miophyilorlwm a very large M-^ and 
liK'k of P^ but its P^ is autapomorphically quite 
distinct and its M- heel very poorly developed. 

Alternatively, M- nversleighensis could be a 
disxanl relative of//, hernardsigei of the //. cy- 
Hops group, interpreted by Hill ( 1 963), Flannery 
& Colgan (1993) and Hand & Kirsch (in press) 
as derived hipposiderids. However, although it 
shares with Miophyllorhina a similar M\ it rc* 
tains a reduced P- and its P^ is derived. 

ACKNOWLEDGEMENTS 

WoTk at Rner^lcigh has been supported by the 
Australian Rcseiirch Council, the Depaiimcnt of 



the Environment, Sport and Teirilortes. National 
Estate Grants Programme (QldJ. Qrieensland Na- 
tional Parks and Wildlife Serv ice, the AusinaUan 
Geographic Socieiy, the Linnean Sixiety of 
NSW. ICI, Ihe Queensland Museum and the Uni- 
versity ol NSW. This study would not have |->ccn 
possible without the support and encouragement 
of Michael Archer and Henk Godthelp. The fol- 
lowing people kindly provided access to compar- 
ative specimens in Iheir insiiruiions: B. Enge.ss^* 
H. Fcltcn, T. Flannery, W. Fuchs, L. Gibson, J. E. 
Hill, M. Huguency, P. Jenkins. D. Kitchener, K. 
Kcjopinan. P. Mem. R. Rachl. B. S»g<S. N. B. 
Simmons. G. Siorch, ajid S. Van Dyck. i am 
grateful to Coral Gilkcsun for access to an SEM 
at MucqtJiiric Universily- 

f. ITER ATU RE CITED 

ARCHER. M.. GODTHELP. il.. HAND, Si. & 
MEGIRIAN, D, I9.S9. Fos.sil mammaK of 
Rtvcrsleigh. nonhwcsiem Uueensland. prelimi- 
nary overx'iew of biostraligraphy, coirvlation and 
environmental change. The Auslralian ZcxjIi^U 
25: 35-69. 

ARCHLR, M . HAND. SJ & GODTHELP H. 19*^ 
Rivcisleigh, The siary of the animals of 
Auslnilia's ancient inland rainforests. 2nd BJ. 
(Reed: Sydney) 

BLACK, K. & ARCHER. M 1097. Nmuokoida gcau 
nov. (Marsupialia: Phascolarciidac) from 
Riversleigh, nonhwe.^lem Queensland, wilh are* 
N'lsion ol" Litokoida. Memoirs of the Queenslaafid 
Museum 41 :20)-228. 

CREASER, P. 1997 Oligocenc-Miocenesedimei5t.st)f 
River&lclgh: ihc potential significance of topogra- 
phy. Memoirs of Ihc Queensland Museum 4T 
303-314. 

FLANNERY, IX k COLGAN. D.J 1993. A new 
spcciey and two new subspecies ofHipfJoaidtfas 
(Chiroplera) from western Papua New Guinea. 
Records of the Ausualian Museum 45: 43-57. 

HAND. .SJ 1993. Firsi skull of a species of 
Hipposidrro.s {BrtHhtpposttUroy} 

(Microchirnptf^ra* HippositlcridJC), Ironi AuMfa- 
lian Miocene sediments. Memoirs of (lie Queens- 
land Museum 3 1 : 179-192 
1 995. Hippoaideros bernardufiei, a new 
hipposiderid (Mitnichiroplcra) truni Ihc 
Miocene of Australia and a rcconsideraiion of Uk 
montiphyly of related species groups. Miinchitci 
Geowissenschafiliche Abhandlungen. 
1997 New Miocene Icaf-nosed hais 
(Microchiropicra: Hipposideridac) from 
Riverslcigh, northwestern Queensland. Memoirs 
of the Queensland Museum 41 ■ 335-349, 

HAND. S.J. & KIRSCH. J.A.W. in press. A soutncm 
ongin for Ihc Hippvisidcridae iMicrochiropicm)'^ 
Evidence from the Au.sirulian fossil record. In 



354 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Kunz, T. & Racey, P. (eds) Proceedings of the 1 0th 

International Bat Research Conference, Boston. 

(Smithsonian Institution: Washington). 
HILL, J.E. 1963. A revision of the genus Hipposideros. 

Bulletin of the British Museum (Natural History), 

Zoology 11: 1-129. 
1 982. A review of the leaf-nosed bats Rhinonyctehs, 

Cloeotis and Triaenops (Chiroptera: 

Hipposideridae). Bonner zoologishe Beilrage 

33:165-86. 
KOOPMAN, K.F. 1994. Chiroptera: syslematics. 

Handbook of Zoology, VIII, 60. Mammalia: 1- 

217. 
MEGIRIAN, D. 1992. Interpretation of the Miocene 



Carl Creek Limestone, northwestern Queensland. 
The Beagle, Records of the Northern Territory 
Museum of Arts and Sciences 9: 219-248. 

SIGE, B., HAND, S.J. & ARCHER, M. 1982. An 
Australian Miocene Brachipposideros 
(Mammalia, Chiroptera) related to Miocene rep- 
resentatives from France. Palaeovertebrata 12: 
149-171. 

WOODBURNE, M.O., TEDFORD, R.H., ARCHER, 
M., TURNBULL, W.D., PLANE, M.D. & 
LUNDELIUS, E.L. 1985. Biochronology of the 
continental mammal record of Australia and New 
Guinea. Special Publications, South Australian 
Department of Mines and Energy 5: 347-363. 



THE FIRST FOSSIL PYGOPOD (SQUAMATA, GEKKOTA). AND A REVIEW OF 
MANDIBULAR VARIATION IN LIVING SPECIES 

MARK N. HUTCHINSON 

Hutchinson, M.N. 1997 06 30: The first fossil pygopod (SquamaLa, Gekkota). and a review 
of iTiandibular variaiion in livinc species. Kfemoin of the Queensland Museum 41(2): 
355-366. Brisbane. ISSN 0079-8835. 

The STiake-like Auslratiun pygopod li/ards (Pygopodtdae in iradilional taxonomies) show 
considerable variation in mandibular and dental structure, correlated w ith dietary Sfxjcialisa- 
lion in several genera. Following a review ol this variaiion, a fully toothed Miocene deniary 
from Riverslcigh, northwestern QueensJand. is identified as a pygopod, Ihe first in the fossil 
record, Pygoptis hortukmia sp, nov. i& specifically distinguishable tmm Ws'mgPygopits by 
looih morphology and proportions of the syn>physiaj region of the deniary- U Fygopodv, 
Itzards. osJeology, fossils. Miocene. 

Mark N^ Hutchinson, Sauih Australiun Museum, North Terrace, Adelaide, ^oulh AusiruUu 
5000, Australia; 4 December 1996. 



One of the most distinctively Australian squa- 
itiiile radiations is a group of snake-like, virtually 
lifiiblcss lizards, variously knov^n as lliip-fooiod 
li/ards or snake-lizards (Bustard, 1970) These 
have no external trace of ford i nibs while hind 
limbs are reduced to fin*like flaps on each side of 
the vent (Kluge. 1974; Shea. 1993). There arc 35 
species tn 8 genera (Greer. 1989; Shea. 1991; 
Cogger, 1992). All ore restricted to Australia and 
New Guinea. 

Flap-footed li/ards have long been regarded as 
forming a distinct family, the Pygopodidae, 
closely related to the Gekkonidae (Underwood, 
1954. 1957), It has been suggested that the sister 
gnmp of Pygopodidae is noi all Gekkonidae, but 
only the Australian Diplodaciylinae, or some part 
of that subfamily (Kluge", 1987; King & 
Mcngdcn, 1990k Acceptance of this phylogc- 
dettc hypothesis (dissenting views exist; see 
Hsics et al., 1988) would mean changes in taxon- 
omy, with the diplodacty lines becoming a sub- 
family of the Pygopodidae, as proposed by Kluge 
(1987) or the pygopodids becoming a subfamily 
of the Gekkonidae (Bauer, 1990). Pending acon- 
ccnsus view 1 use Underwood's (1957) contrac- 
tion of their traditional family name, 'pygopods\ 
as an informal collective term. 

Pygopods show considerable ecological and 
morphological diversity. /^/jravuv, PleflwlcLS and 
Ophidiocephalns, exhibit fos.sorial adaptations 
and behaviour (Kluge, 1974; Ehmann. 1981; 
Shea & Peterson, 1993). However. PleihoUvc 
j^racilis and at least some species ofAprasia are 
regularly active on the surface by day (Shea & 
Peterson, 1993; pers. obs.). Species in the largest 
genus, Delma. and that regarded by Kluge ( 1 974; 



1976) as most generally primitive. Pygopus, as 
well as Lialis, Aclys and Parodelma, ore surface- 
dwellcrs( Wilson & Knowles. 1988; Greer I9S9). 
While most pygopodids appear to be active for- 
agers feeding on invertebrates, the two species of 
Lialis are ambush predators of scincid lizards 
(Palchell& Shine, 1986a, h; Murray ei al., 1991). 

The mandibular and dental anatomy of 
pygopods t.s varied, and at least panly correlated 
■with the ecological diversity jii.si mentioned. The 
one overview of pygopod osteology attempted 
(Stephenson. 1962) gave .scant attention to the 
mandible. Kluge (1974, 1976) employed some 
mandibular characters in his analysis of the cladc, 
and Rieppel (1984) noted some correlations be- 
tween anatomy and miniaturisation in Aprusia 
and /^/^r/;o/rt.v when compared to Pygopus. Parker 
(1956) noted sexual dimorf)hism in Apnisia, in 
which only males have pretnaxillary teeth. 

Austndia's fossil lizard fauna is poorly known 
tEstcs. 1983a. b; Covacevich et ai., 1990; Hutch- 
inson, 1992); no fossil pygopods have been iden- 
tified Fossils are essential for establishing a 
minimum estimate of the lime a taxon has inhab- 
ited an area and of the time since the taxon first 
evolved. If pygopods are the sister group of 
diplodactylines. especially the Dipiodactylini 
(Kluge. 1987). then it is likely that their differen- 
tiation occurred in Australia and should be re- 
corded in the Australian fossil record. This paper 
is the first report oi a fossilised pygopod. 

MATERIALS AND METHODS 

Dried skeletons (2 1 species in 8 known genera- 
Appendix I were examined to assess inlcrspcciilc 



356 



MEMOIRS OF THE QUEENSLAND MUSEUM 




cui ±L nDAf) nr\rsr\^fV)rf] /r) 





ami 



FIG. 1. Labial (A) and lingual (B) views of the righi mandibular ramus of a pygopodid lizard. Delma J'raseH 
(SAMAR2291 K Ctxomalbidgup, W.A.) showing major features. Abbreviations; adfo=adductor (=MeckeJian) 
fossa, amf=anieriur mylohyoid foramen; asf=anierior surangular foramen; t=coronoid; cp=dorsal process of 
coronoid; cpd=compound bone - fused prearticular, surangular and articular; c(f=foramen ft>r chorda ty mpani; 
d=tkniar);iaf=inferiora]veolarforamen:prp='prearticularprocess' (medial articular process of the surangular); 
psf=postcrior surangular foramina; rap=Tetroiirticular process; s=spleniaL Scalc=5mm. 



variation in the sutural relalionships of the man- 
dibular bones, positions of foramina, tooth num- 
ber and tooth morphology. With the exception of 
Pygopus, this sample did not permit study of 
mtraspecific variation. Unless otherwise speci- 
fied, the descriptions refer to the anatomy as seen 
in intact mandibles. Outgroups used to infer de- 
rived character states are diplodacty lines as the 
sister group of pygopods. gekkonines the sister of 
ihe.se two and other scleroglossans as the most 
distant outgroup (Estes ci a!., 1988). 

PYGOPOD MANDIBLE (Fig. 1). 

The dentiiry is the largest bone of the pygopod 
mandible. It consists of the tooth-beanng body of 
the bone and a relatively kmg, pusieriorly di- 
rected angular prexress that covers much of the 
labial and ventral surface of the mandible. 
Pygopods share with other gckkonoids and mem- 
bers of several other families the complete oblit- 
eration by dentary overgrowth of the groove for 
Meckel's cartilage, but differ, ai least from all 
diplodaciylines examined, in that the angular pro- 
cess extends on the labial surface of ihe mandible 
to well behind the coronoid (the dentary also 
extends posteriorly to a marked degree in the 
gekkonine Paroedura, Klugc, pers. comm.). The 
dentary of dipiodaclyline and gekkonine geckoes 



is generally more slender and incurved than that 
of pygopods and the angular process terminates 
at about the level of the coronoid. 3-8 menial 
foramina open along the labial surface of the 
dentary. the series generally extenditig posteri- 
orly to 1/2-3/4 the length of the tooth row. There 
is no posterior extension of the bony internal 
septum separating the Meckelian cartilage from 
the inferior alveolar nerve; bony separation is 
limited to the immediate vicinity of the mental 
foramina, Estes ct al. (1988) described this char- 
acter (their number 56) in terms which 
emphasised that a vertically-oriented, posteriorly 
extended intramandibular septum is well-devel- 
oped in anguimorphs. but failed io note that it 
occurs 10 almost the same extent in lygosomine 
skinks (Shea & Hutchinson, 1992, fig. 3). 

Adult geckoes typically have large numbers oi 
small dentary teeth (Bauer & Rusself 1990: 
Klugc & Nussbaum. 1995); in a sample of 9 
dipiodaclyline, 1 sphaerodaclyline and 14 
gekkonine genera, denlarycounls ranged from 25 
iPbelsumo mackigascahensis) to 62 {Saltuarins 
safebrosus), with a mean of 4(_).2 (Edmund. 1 969; 
pers. obs.). In adult Delrna and Adys tootJi num- 
ber is very much like that of dipkxlaclyline and 
gekkonine geckoes, typically in the range 25-35. 
Reductions to 24 or fewer, or increa.ses to 50 or 
more are ihcrcfore likely to be apomorphic 



PYGOPOD MANDIBLE 



^57 



(KIu^, 1976). Diplodaclylincs amJ gekkonincs 
usual Jy have slender, upright teech wiih acuie 
crowns he^iring a pair of noimed apical cusps 
scparatcdby agrw)VtMSum»da& Murphy. 1987). 
Teeth wiih ihis morphology occur in some 
pygopods {Delma and Aciys), and iire probubly 
plesi amorphic for the group. Within pygopcds 
ihcrc is marked iniergencric varialion. including 
robust, upright or siigluly recurved leelln less 
robust bui more strongly recurved tccih, or vcr>' 
small leoih with compressed, sharp-edged 
crowns. All retain an apical groove, bul the bicus- 
pid structure is latgely lost, the labial cusp enlarg- 
ing (o become the tooth apex, while the lingual 
Cusp ail but disappears. In Ualis tooth crowns are 
so compressed that (he apical groove is faini arid 
only discernible in unworn teeth. 

The coronoid consists old lateral ly compressed 
dorsal process, an amcriorly-dirccled dcnlary 
process and a ix)Sterior process. The deniary pro- 
cess is bifurcated and clasps the denlury hone 
behind Ihe end ot the tooth row both labially and 
lingually. On the lingual face of the mandible, the 
anterior exlremiiy of the dcntary process termi- 
nates posterior to, cocxiensively with, or anterior 
to the front ol the spicnial, the latter 2 character 
scales being apomorphic with respect to other 
gckkonoids and other liziu"ds. The posterior pro- 
cess of the coronoid extends to the anterior ex- 
tremity of the Meckelian fossa. The form of the 
dorsal process vanes from lall and fin-like in 
several genera to very low in Ualh (Klugc, 
1976). A well-developed dorsal process \^ the 
rule in geckoes and other li/ards and is likely to 
be plesiomorphic for pygopods. 

"The spleniai is reduced in all pygopods com- 
pared to the development seen in diplodactylines 
and other geckoes, usually failing to extend ante- 
riorly beyond the level of the distal two or three 
teeth. In most pygopods the spleniai is further 
reduced in length or depth. The spleniai is com- 
pletely absent in Apnisia (pcrs. obs.; Parker, 
1956; 'very slight'. Stephenson, 1962). The 
spleniai, when present, comptclely surroufids (as 
in diplodactylines) the inferior alveolar foramen 
ar>d bears a notch for the anterior mylohyoid 
foramen on its ventral suture with the dcntary. 

Like the majoriiy of geckoes (Kluge. l9fi7X 
pygopods lack a distinct angular. The spicnial in 
Dehna and Pygopus has a posteriorly extending 
process that separates the deniary and prcarticu- 
lar, as would an angular, suggesting thai the an- 
gular has been iosi via fusion wiih Ibe spleniai. 

The surangular is fused labially and p<>stcriori> 
wiih the fused prcariicular-aniculur In aduh 



pygopods, but is completely disiinct in juveniles 
{Delma inotleh, Uaih burtonis and Pygoput 
icpidopcuJui), In aduhs of mo^i genem a suture 
persists on the lingual fac« of Ihe mandible run 
ning antenorly (rum the Meckctian lossa; in in- 
tact mandibles this suture may be concealed by 
the coronoid. 3 foiamma are usually pitisent on 
the labial surfaceof the surangular. The antcrinrly 
directed opening of Ihe anterior surangular fora- 
men lies on the labia* suifijcc on or jusi posterior 
to the point of inicrscctinn of the sutures bep^ocn 
the coronoid, dcnun- and Mirangular. 2 ochcr 
foramina usually lie towards the pusterolnbial 
region of the surangular, but there is iniergenetic 
variation. In some pygopods there may be only a 
single foramen, as in diplodactylines. but in nto&l 
there are 2. The 2 openings may be close togeiljer, 
or moderately separated, and ihc more 
posterodorsal of the 2 may itself be subdivided. 

Mast lizards have only'a .single posterior sur- 
anguUir foramien, and sc the additional (mive 
anterior) foramen iT»U-si be identified. Klu^,c 
(1967] and Grismer ( 19H8) de^ignaied \hc m«.w 
ventral of the posterior foramina as the posterior 
mylohyoid and Ihc more dorsiU as the posterior 
surangular foramen. My survey of gckkonoid 
mandibular variation sug;gcsis that this interpre- 
tation IS inLorrcct. 

Tlie posterior mylohyoid nerve innervates lite 
throat musculaliii'e (Camp. 1923; Poglaycn-Ncu- 
wall 1954) aiMl typically exits lbn>ugh a medially 
or vcntrx>mcdiallydi reeled ibramen in the angular 
bone. While the angular i& absent in most 
gek'konoids I.KIugc, 1987) sonw of those exam- 
in«;d (the gekkonines GMo^ Fhehumo and f*hy- 
Uodaaylm \=Christimis\) retain a small forank?n 
on the dentary-splenial suture in the expected 
topographic position of Ihc posterior mylohyoid 
foramen. Other gekkonines and other gckkonoids 
examined, including all pygopods. liick a fora- 
men in this position. 

The foramen identified by Klugc (1967) and 
Grismer ( 1 9S8uis the posterior mylohyoid opens 
on the labial surface iif the mandible and run;* 
through to open lingually mio the Meckelian 
fos.sa. Thi.s foramen thas follows the course of Ihc 
posterior surangular foramen of other li/ards, and 
it too iransmiis a branch of the mandibular nerve 
to the adductor (picrygoidcusl musculature (pers. 
obs.). The foramen and nerve arc remote from the 
throat musculature which, hy definition, the pos- 
terior mylohyoid supplies. It therefore seems to 
me more probable that the additional foramen on 
(he labial surface of the surangul<ir in pygopods 
sind other gekkonoids reprvs^ents « duplic}tti4jn of 



358 



MEMOIRS OF THE QUEENSLAND MUSEUM 











FIG. 2. Labial views of right mandibular ramus of the 8 pygopodid genera. A, Delma mornaia (SAMAR22408. 
no data); B. Aclys concinna tSAMAR38060, Badgingarra National Park, W.A.); C, Pygopiis nighceps 
(SAMAR2I029, no data); D. Pamdelma onenlalis (QMJ30250, Cracow, Qld); E, Ophidiocephahis taeniatus 
(SAMAR28365, Abminga, S. A.); F, Pletholax gracilis (SAMAR3806 1 , Jandakot, W.A.); G, Aprasia stholata 
(SAMAR35569, Mylor, S.A.); H, Lialis hurtonis (NMVDl 5399, Warby Ranges, Vic). Scales=5mm. 



PYGOPOD MANDIBLE 



359 










FIG. 3. Lingual views of same specimens in Fig. 2, drawn to same scales. 



360 



MEMOIRS OF THE QUEENSLAND MUSEUM 



the posterior surangular foramen, not a displaced 
posterior mylohyoid. 

The fused prcanicular-articulur logcihcr with 
ihc sunmguliirconfili lutes the posterior 1/3 of the 
pygnpod mandible. As in di|)IodacTylincs and 
Other gelvkonoids the ariiculiiiing facet is oriented 
to face postcrodofsally rather than dorsaliy, re- 
sulting in a marked *stcp' down from the level of 
the upper edge of the adductor fossa lo the level 
of the reiroarticular princess. The latter structure 
is variable intergcncrically in its shape (spoon- 
shaped to rod-like) and the degree of media! or 
ventral inflection (Klugc, 1976). A foramen for 
the n. chorda lympani opens on the dorsolingual 
aspect of the bdse of the retroarticuiar process. 
The posterior region of the Mcckelian fossa, 
which contains the internal opening for the pos- 
terior surangular tbramen. nu^y be demarcated 
from the anterior region. 

MANDIBULAR VARIATION IN LIVING 
PYGOPOUS 

DEtMA. This genus has the most generally 
'gccko-likc' iTiandibles. However, there is signif- 
icant variation within the genus in proportions, 
lOiMh cn^wn shape, si/e of splenial and other 
fealuncs. Deitnafraseri (Fig. I) and D. inornata 
(Fig* 2A. 3A) show some of this variation, with 
the latter species tending to retain more 
ple.siornorphie features than the former. The 
pygnpod synapomorphy of extensive posterior 
extent of the dcntary on the labial surface is 
present in all Dclnui , but the teeth are nunierous, 
with unmodifiCii (e.g. O impar, D. inomaia, D. 
nasuta) or slightly expanded crowns (e.g. D. 
fraseri, D. mitellaV Whether thiii is inter- or in- 
traspcci fie variation will retjuiixsa iwreexfcnsivc 
survey. Tooth crowns retain the bicuspid mor- 
phology of diploitactylinc geckoes. The deniary 
is moderately slender and the splcniol little re- 
duced, although it usually fails ti> exicfvJ anteri- 
orly as far as the poslcri(jrmosl loolh {O. inornata 
specimens showed inu-aspecitlc variation, the 
splenial failing to reach the tooth row in 
NMVDI5448. reaching the second-last tooth in 
SAMAR35570 and extending as far as the sixth- 
last m SAMAR22408, Fig, 2A). The posterior 
surangular foramina are moderately to narrowly 
separated (variable both inter- and intraspecifi- 
cally) The Relatively unspccialiscd dentition is 
associated with a generalised arthropod diet 
(Shine & Putchcll, 19S<ia; Coulson. 1990). 

AC/,r5 (Figs 2B,3B J. In general the mandible of 



this genus is an elongate version of Detma's, 
Elongation of the jaw occurred by lengthening of 
the region between the tip of the coronoid process 
of the dentary and the posterior ax^d of the tooth 
row, with hypertrophy of the lingual ramus of the 
anterior process of ihc coronoid reducing the 
exposure o\' the splenial and widely separating it 
from the tooth row. Height ol the dorsal process 
of die coronoid is^ reduced relative to most Delma 
and the medial articular process (Fig. I) of the 
surangular is elevated, both trends foreshadow- 
ing the exicnsivc coronoid flattening and sur- 
angulitr elevation oi Lialis, The derived features 
of the jaw of Aclyt are all seen, although not lo 
the same degree, in some Delma, especial ly those 
u iih more elongate skulls such as D butler t and 
D. nasuta, 

/0*CO/'C*S(Figs2C.3C,4B.4D).The mandibles 
of the two species placed in this genus arc very 
similar to one another, and probably mdislin- 
guishahle. The form of the mandible is 
ttpomorphic with respect \ok\\\xXo\ Lkinm in being 
shorter and deeper. The dentition is also 
apomorphic, the teeth being fewer (<2.5), much 
more robust, and with .sharp, tapering, recurved 
crowns. The tooth crowns retain a pronounced 
apical groove, but the typical gckkonoid bicuspid 
structure is reduced, with the labial cusp being the 
principle tooth apex while the lower, lingual cusp 
is little more than the acute-angled inner margin 
of the apical groove, Tl^e mandible is more 
plcsiomorphic than most Delma in that ilic splen- 
ial extends forward lo under ly the posieiiormost 
leeth-The posterior surangular foraminu are nnxl- 
eraicly to widely separated (intraspecifically 
variable). The genus is characterised by a pro- 
noutwed mesio^tistal decrease in inoth size, the 
mesial teeth (second to fifth) being 30-40% taller 
than the mid-dcniiiry teeih (tenth lo twelfth). The 
enlarged teeth at the front ot the jaw arc some- 
what procumbent and arc supported by a deep 
syinpbysial region. Pianka (1986) described P. 
rtii^ncepsas a scorpion specialist, and Patchcll & 
Shine 1 1986a) found thai the major prey oi f\ 
Icpidopodus were mygalomorph and lycosid spi- 
dci*s. Possibly the relatively powerful front teeth 
arc adaptations for rapidly disabling such poten- 
tially dangerous prey. 

f'AKADE/J^A (Fig.s 21^, :^f^). This nionodypie 
genus, like Pygopus. has a reduced loolh number 
(21 ) compared with Delma but the teeth are dtS- 
tinciive. beittg more slender than in Pygopusand 
having recurved crowns. The tooth apices arc like 



PYGOPOD MANDIBLE 



m 



Ihose of Fygopus in ihal the apical groove is 
prcscni bui the lingual cusp is barely developed. 
Compared with RyKoptn, the jaw is less tobusi 
and more bowed. The diet is unknown. 

OPHJDIOCEPHALUS (FigslE. 3E). The jaw of 
this small fossorial form is relatively robust, short 
and deep, similar in proportions lo that of 
J*y^opNS, but is apomarphic in several characters. 
The splenial is greatly shortened and shifted pos- 
teriorly compiircd with Pygopus. The sin^jle jaw 
examined is distinctive m thai the lingual ramus 
of the anterior process of the coronoid is reduced, 
«X|K»sing the bone beneath. Based on the position 
of the prcarticular-surangular suture exposed 
below the dorsal process of the coronoid. this 
anteriorly exposed bone is the surangular. The 
posterior surangular lt)ramina arc narrowly sepa- 
rated. The teeth are similar in to those ofPygopns, 
but are fewer in number (13-15 in adults versus 
17-24 in hatchling to aduU Pyiiopus) and have 
moderately recurved crowns. Recorded prey in- 
diciiics a relatively generalised arthropod diet 
fF.hmann, 1981). 

PLETHOIAX (FigS 2F. 3F). The jaw is long and 
slender, but with a wcll-dcvelopcd dorsal process 
of the coronoid. Tooth counts arc below 20 
fKluge, 1976), with the mesial teeth relatively 
robust, pointed and slightly recurved while the 
n»ore distal teeth are markedly reduced in si/e 
The splenial is reduced to a narrow splint but still 
encloses the inferior alveolar foramen and forms 
ti\i' dorsal margin of the anterior myli»hyoid fora- 
men. There is a single posterior surangular lora- 
men. Shea A Peterson (1993) found that most 
guts of this species contained little chitinous ma- 
icrial but often included short cut lengths of grass, 
consistent with digestion from the bodies of in- 
sect prey. They suggested the most likely diet was 
poorly-sclerolised, readily digested prey such as 
leriniles, Ehmann (1993) suggested that 
PlviholcLK is a neciar feeder. The w cak jaws and 
small, widely spaced teeth suggest it would be 
unlikely to deal effectively with the tough exo- 
skelctons of typical invertebrate prey. 

APRASJA (Figs2G, 3G). The mandible has only 
3 elements; dcntary, coronoid and a compound 
bone representing the remaitidcr; the splenial is 
absent, whether through loss or fusion is unclear. 
The inferior alveolar foramen lies on the suture 
between the anterior extremity of the compound 
bone and the dcntary. The posterior surangular 
foramen is smglc. The dorsal process of the 



coronoid is reduced and the retroarticular process 
is abbreviated. The teeth are very greatly reduced 
in number, restricted to a patch of 3-4 rclativxrly 
robust, pointed, recurved teeth situated close \o 
(but not on) the symphysis. The diet is restricted 
10 the eggs, larvae and pupae oi ants (Webb & 
Shine. 1994). The mandibular and dental anat- 
omy oiAprcuiia is thus convergent to some extent 
on that of ant-eating scolecophidian snakes, 
which also have weak, almost edentulous jaws 
save for a few teeth m the upper (Typhlopidue) or 
lower (Lepwyphlopidae) ntandiblc. 

UAUS (Figs 2H. 3H). This genus has a highly 
derived mandibular morphoktgy. The denlary Is 
greaUy ailcnuated. with many (to over 60) small. 
recurved, sharply pointed teeth having ligamen^ 
lous basal attachntents (/> bitrtonis\ Patchell & 
Shine. 1986b; L jicari. C. Shea. pers. comm.)» 
Other peculiarities include the greatly reduced 
height of the coronoid and a peculiar fan-like 
medial articular process of the surangular ascend- 
ing well above the level of the articular region, 
and higher than the coronoid. Functionally this 
braces the jaw against the quadrate ramus of the 
pterygoid, preventing lateral displacement of the 
jaw when the mouth is open Along with these 
apomorphies. Lialu retains a picsiomorphic, rel- 
atively large, antenocly extending splenial. The 
postetiof surangular foramen is single or double. 
The wide gape and spcciali.^d dentition are re- 
lated to the ohiigalc lizard-eating (especially 
skink-ealing) habits of this genu;s (PatcheJI & 
Shine, 1986b). 

SYSTEA1AT1CS 

Infraivder GEKKOTA Cuvicr. 1KI7 
Family PYGOI*ODIDAE 

Pygopus Merrem, 1870 

TYPE SPECIES. Bipcs lepidopodus Laccpcde, 1804 

Fygopus hortulanus sp. nov. (Figs ) 

ETYMOLOGY. Latin horiulanus. ol or belonging lo 
a garden; alluding to the Neville's Garden Site. 

MATERIAL. Holoiype QMF16875 (Fig. 4), a right 
dentary preserving ihe complete looih row and sym- 
physial region but minus the angular process from early 
Miocene. System B (Archer et al.. 1994), Neville's 
Garden Site on D Site Plateau at Riversleigh, NW 
Queensland; the site is interpreted as a'prcsentin^t^n 
accumulation m a pool close lo a cave entrance . 



3^ 



MEMOIRS OF THE QUEENSLAND MUSEUM 






FIG. 4 Comparison of the dcntaries ofPygopus hortulanus SP. NOV. (A and C, QMFl 6785) and P. lepidopodtts 
(B and D; SAMAR38928). A-B, labial views; C-D. lingual views. Note larger mesial (anleriormost) teeih and 
deeper symphysis in P. lepidopodus. Scale=5mm. 



DESCRIPTION. Right dentary, complete and 
undamaged except for the absence of the angular 
process posterior to the level of the end of the 
tooth row. Dental arcade in occlusal view straight 
from distal tooth anteriorly to about seventh tooth 
(counting mesiodislally), then curving gently me- 
sially. Lingual rim of dental sulcus distinct, 
rounded in cross-section. Groove for Meckel's 
cartilage completely obliterated by overgrowth of 
dentary. Inlramandibular septum not extended 
posteriorly. In labial view, acute-angled splenial 
notch extending to level of penultimate (twenti- 
eth) tooth, and superficial facet for anterior exten- 
sion of splenial extending as far as seventeenth 
tooth. Wedge-shaped facet for anterovenlral ex- 
tremity of coronoid incised into posterior margin 
between dorsal edge of splenial notch and last 



tooth. Labial surface with 5 mental foramina, last 
level with fourteenth tooth. Shallow fossa on 
labial surface tapering anteriorly from 
poslerodorsal margin of dentary. Total length 
6.0mm; depth at level of last tooth 1. 4mm (tooth 
excluded). 

Tooth loci 21. Counting from mesial to distal, 
all hut first (broken), ninth and fifteenth (empty) 
with intact teeth. Twentieth tooth lost subsequent 
to preparation of Fig. 4. Length of tooth row 
5.2mm. 

Teeth robust, 2.5-3 times as high as wide, 
crowns tapering rapidly to sharp points. Well-de- 
fined groove traversing each tooth crown, setting 
off weak second apical point, lower and lingual 
to main tooth apex. Tooth crowns slightly in- 
curved. Teeth reducing gradually in size from 



PYGOPOD\tANDIBL£ 



363 



mesial to distal, but ameriormosi teeth onI> 
slightly larger than mid-dcntary icclh, 

REMARKS. Idcnliflcation of the fossil as a 
gckkonoid is bused un aponioqihic features: ]) 
obliteration of the groove for MeckcPs cartilage 
by overga>wih of the. denlary, 2) apex o( Uie 
splcnial notch level with ihc posterior end ol* the 
loolh row. with a narrow tongue of the splenial 
extending forwiird over the lingual face ul Ihc 
dcntary below the looih row, and 3) n>csial (i.e. 
anieriormost) leeth largest (rather than mid-den- 
lary teeth). 

Many Australian skinks (Scincidac) also have 
ifxr Meckelian groove obliterated by ihc dentiiry, 
hut in these the splenial notch exieiidii well for- 
ward under the tooth row (the tiny Norosrhirus, 
loolh row <:3mm, is an exception), and scincid 
denlancs wilh a closed Meckel's groove all have 
n vertically oriented, posteriorly extended in- 
iramandibular septum. Skinks, unlike 
^ekkonoids (Bauer* 1990; Bauer & Russell, 
1990), have the mid- dentary teeth larger than the 
nocsial leclh. Scincid dcniarics also have a pro- 
nounced coronoid process on the labial surface 
(Esles, 1983); although this part of the specimen 
is incomplete, existing outlines indicate that very 
lilllc of this region is lost and that there was 
scarcely any development of this process. The 
only other lizard families in which dcntary oblit- 
Cfalion of ihe Meckelian groove occurs ai'e the 
Xantusiidac (all), many Gymnophdialmidae and 
manviguanians(Prcsch. 1980; listescla!., 1988; 
EJhcridgc & dc Queiroz, 1988; MacLcan, 1974). 
These can be excluded on the basis of loolh crown 
shape (McOovvell & Bogen. 1954. Sumida & 
Murphy. 1987; MacLcan. 1974; Ethcridgc & de 
Queiroz* 1988), and having mid-deniary leeih 
larger than mesial teeth. 

Among gckkonoids, only pygopods have the 
stout, straight dentary shape, low dentary tooth 
counis and leeili of ihc robust form seen in P. 
hortulanus. 

Fygopus hortulanus shows two seemingly de 
rived characier states within pygopods, the rcla 
lively shore, deep dentary and the fewer robust 
Icclh. These relate il not only lo I'ygopus but also 
OfMdiocephalus. The fossil is plcsiomorphic 
with respect to Ophidiocephalus in straighter 
looth crowns, more teeth and greater anterior 
extent of the splenial. it is plesiomorphic with 
re->j>ect to Fygopus perhaps in the less enlarged 
mesial leeih and (?coTTelated) shallower sym- 
physial region. AltcmaLivcly the more even tooth 
row of F. hortiilamts could be inierpreied as 



autapouioriJhic, because enlarged anterior ie<;ih 
are cointnon in pygopods. The placement of the 
species in Py^opus rather than Ophtdiocephalns 
reflects ihc fewer specialisatiomi shared with Op- 
hidiocephalus than with Pygopus. 

The cladistic analysis of intergeneric nelalion- 
ships reported by Klugc (1974, 1976) not only 
concluded that Pygopus is the most primitive 
extant genus but also placed Pygopttx as a gra<k 
group at the base of the pygopod radiation and 
suggested that Paradeima orienudis is more 
closely related to Pygopus uigrkeps than the 
latter is to Py. tepidopodus. The placement ot the 
Ibssil in Pygopus could thcrelbre be taken to 
ntean only that the fossil is a plcsiomorphic pygo- 
pod. However, the short, deep dentary and robust 
teeth of Pygopus are probably apomorphic in 
pygopods. and l therefore maiiMain the concept 
of this genus employed by Cogger ( 1985), Grccr 
(1989) and Shea (1993). 

This early Miocene pygopod i.s consistent wilh 
Kluge's (1987) suggestion that pygopoOs 
evolved on the Australian continent subsequent 
to a Late Cretaceous vicariant event isolating tl>c 
ancestral diplodaciylinepygopod siock. This 
find, with its apomorphic teeth, is not u 
generalised ancestral pygopod, implying the ori- 
gins of the group must be older than the Miocene. 

Archer et al (1989; 1995) suggested tliaf the 
Mioccnccnvironmcnt of Riversleigh was primar- 
ily tropica! closed forest, an assessment sup- 
ported by many genera in Systems B andC whose 
living representatives arc restricted to closed for- 
est environments. Mcgirian ( 1 992) suggested that 
scdimcntoiogical evidence argued foramoreiirid 
clitnaie with any rainforest Hmhcd to water 
courses. Creaser (this volume) reports that sedi- 
ment patterns regarded by Megirian as being 
confined to arid depositional environments occur 
today in mid-montanc New Guinea. 

No modem pygupud inhabits rainforest al 
though Lialis iuid some DHmu (Shea, 1987) in- 
habit vine scrubs and cucalypt forest on the 
margins of Tainforcsl. The presence of /^ 
hortulanus in System B could be inierpreied to 
indicate either that ibctc were drier, open patches 
nearby, or that P. hortulanus wiis a rainfonr-st- 
dwelJer wilh no living analogue, 

ACKNOWLEDGEMENTS 

I thank Mike Archer for enabling me lo work 
on the Rivcrslcigh lizard fossiU. G.M. Shea pre- 
pared (he dentary of Ihlnm vxMUi, A.G. Klugc 
and G. M. Shea commented on ihc fossil. Rjr 



364 



N4EM0ERS OF THE QUEENSLAND MUSEUM 



Kfrni-sston to prepare skclcral malerial from cot- 
rlions in iheir care and for ihe loan or exchange 
of .specimens, I ihank J. Cnvacevich and P. 
Coupcr (Queensland Museum), K. Aplin and L. 
A. Smilh (Wesiern Australian Museum) J. Cov- 
entry (Museum of Victoria) and R. Sadlicr (Aus- 
Indian Museum). Fieldwurk at Riversleigh is 
supported by the Australian Research Grants 
Scheme, Australian Dcpanment of ihc Arts, 
Sport, the Environment, Tourism and Territories, 
National Esiaie Pro^^^amrne Grants Scheme, Uni- 
versity of NSW; Wang Computers, AusUalian 
Geographic Socieiy. Mount Isa Mines, Queens- 
land Museum, Australian Museum. Royal Zoo- 
logical Socieiy of NSW, Linnean S<xiety of 
NSW. AnsetVWridgways, Mt Isa Shire Council, 
Rivcrslcigh Society and the Friends of 
Riverslei^h. 

LITERATURE CITED 

ARCHER. M., GODTHELP. H.. HAND, S. J & 
MEGIRIAN, D. ]W9. Fossil mammals of 
Rivcrslcigh, nonhwestcm Queensland: a prelimi- 
nary mvrv'iew of biostratigrnphy, correliition and 
ciivtaKiment^tl change. Australian Zoologist 25: 
29-65- 

ARCIIER, M, HAND, S. J. & CJODTHELP. H. 1994. 
Rivcrslcigh 2nd cd. (Rccd: Sydney) 
1995. Tertiary environments and biolif cliangc in 
Ausualia. Pp. 77-90. In Vrba.E.S.. Denton. G.H.. 
PiirtrKlgc.T.C.& Buakle,L.H Paleoclimale and 
evolution, with emphasis on human origins. 
(Yale University Press: New Haven). 

BAUER. A- M 1990. Phylogcneiic sysicmatics and 
biogeography of the Carphodaciylini (Rcptilia: 
Gekkonidae) Bimner ZovdogiNches 
Monographicn 30. I '217. 

BAUER, AM. & RUSSELL. A P. 1990. Dentitional 
diversity in Khacodactyhix (Rcptilia: 
Gcklconidae). Memoirs of the Ouccnslaod Mu- 
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36S 



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APPENDIX 

Pygopodid skeletal specimens exanuTicd:/lc7>jr 
cotuitifta SAM A R38060; Aprasia inaunia 
SAMA R 14275: A. pseudopitlchella SAMA 
R406A; A. striolata SAMA R35569. R41S25: 
Aprasin .sp, SAMA unregistered alizarin speci- 
men, Mma aitsfralis SAMA R 1595S; D. Intfleri 
SAMA RI4913. R16843A; D. fraseri SAMA 
R22911; a impar NMV D 1 5446; D. inomma 
SAMA R22408, R35570, NMV D 15448: D. 
niitellQ AMS R65264 (panial deniary only); /). 
ntolhn SAMA R2254CI. R35572: O, na-UfUi 
SAMA R22517: D. plei>eia QM J5K9 1 : D. timla 
SAMA RJ51S9A; UaUs burwnts SAMA 
K15882.R4U031,QMJ47481.NMVDI53yy;iL. 
jicari SAMA R I 1 44 I ; Ophidiocephotus 



366 MEMOIRS OF THE QUEENSLAND MUSEUM 



taeniatus SAMA R28365 (mandible only); 
Paradelma orientalis QM J30250 (mandible 
only); Pletholax gracilis SAMA R38061; 
Pygopus lepidopodus SAMA R 19604 (mandible 
only), R35571, R38924-28; P. nigriceps SAMA 
R1250,R21029. 



TWO >^W EARLY MIOCENE THYLACINES FROM RIVERSLEIGH, 
NORTHWESTERN QUEENSLAND 

JE ANETTE i4UIRHEAD 

Muirhead, J. 1997 06 30: Two new early Miocene ihylacines from Riversleigh. northwestern 
Queensland. Memoirs of the Queensland Museum 41(2): 367-377. Brisbane. ISSN 0079- 
8835. 



Thylacines, Wabulacinus ridei gen. el sp. nov.and Ngamalacinuslimmutvaneyi gen. ei sp. 
nov., are described from ihe early Miocene of Riversleigh, northwestern Queensland. Both 
show carnivorous adaptation iniermediaie between thai of the plesiomorphic Nimbac'inus 
dicksoni and derived Thylacinus. The family concept is revised lo include these new taxa. 
All known Ihylacinid genera occur in late Oligocene to middle Miocene Riversleigh faunas 
and sonic may have overlapped in lime followed by a decline in family diversity since the 
Miocene O Thylacine, marsupial, carnivore. Miocene, Riversleigh, Queensloftd. 

J. M airhead. School of Biological Sciences, UrdversUy of New South Wales NSW 2052 
Australia: received 25 June J 995. 



The Thylacinidae consists of three .species of 
Thylacinus (T. cynocephalus Hams, 1808, T. 
fX}iens Woodbume, 1967 and T. macknessi 
Muirhead, 1992) and the monotyxAcNimbQCinu^ 
dicksoni Muirhead & Archer, 1990 from the late 
Oligocene to middle Mjocene of Queensland and 
the Northern Territory (Muirhead & .^Vrchcr. 
1990). It is the oldest and most p^rimilive ihy- 
lacinid, more closely resembling dasyimds in 
many plesiomorphic features. Thylacinus potens 
from the laie Miocene Alcoota Local Fauna 
(Woodbume, 1967) is considered (Archer, 1982) 
live sister species of modem T. cynocephalus and 
is almost as specialised. Thylacinus macknessi. 
From early to middle Miocene Riversleigh faunas. 
Is also a highly specialised thylacine. Because it 
retains some plesiomorphic features, it is consid- 
ered to be die sister species to die T. potens -7. 
cynocephalus clade (Muirhead, 1992). T^^x) new 
early Miocene thylacinids from Riversleigh are 
described here. In many features they provide a 
continuum in morphological change from the 
plesiomorphic dentition ofN, dicksoni to thai of 
specialised Thylacinus. Dental nomenclature fol- 
lows Flower ( 1 869) and Luckett ( 1 993) where the 
adult dentition includes PI -3 and Ml-4. Taxo- 
namic nomenclature follows Muirhead & Archer 
(1990). Material is housed in the Queensland 
Museum (QMF) or Northern Territory Museum. 



SYSTEMATICS 



Order DAS\XTROMORPHIA (Gill. 1872) 
Superfamily DASYUROIDEA{Goldfuss, 1820) 
Family TH\XACIMDAE rBonaparte, 1838) 



Wabulacinus gen. nov. 

TYPE SPECIES. WabuIaciAus ridei gen. ei sp. nov. 

ETYMOLOGY. Wanyii Wabula, long ago; Greek 
kynos^ dog. Masculine. 

DIAGNOSIS. Iniraorbital foramen surrounded wholly 
by themaxillary and positioned low and anterior to M ' ; 
cenuocrista and prcparacrisia parallel, fomiing contin- 
uous straight line on M*; entoconid absent (on M3); 
hypoconulid enlarged (on M3). 

COMPARISON, Wabulacinus differ fn^m A^. 
dicksoni by larger size, lack of stylar cusps B and 
D on M', lack of stylar cusp B on M^ and the 
minute sjze of St D on this tooth, the straight or 
almost straight centrocTista on M^ and M^, ante- 
rior cingulum of M^ has no notch for placemcni 
of preceding premolar, iheanteriorrooi of M' lies 
directly under the cingulum , U>c anterior width of 
the upper molar crowiis are less than that of ibc 
buccal Icngdis, wider angle of crests at ihc 
paracone ar»d roetacone, exueme reducuon of the 
talonid basin and protocone, particularly on M* 
with concurrent loss of meiaconules on this iooth» 
extreme reduction in size of the metaconid, ab- 
sence of entoconid, reduced lalonid basin by the 
more lingual position of the hypoconid and luck 
of diasiemata between Pi and P2. 

Species of Wabulacinus differ from all species 
of Thylacinus in the extreme reduction of the 
iak)n and protocone on M', the more parallel 
alignment of the preparacrista with the cen- 
trocnsta on M^ , a small melaconid (at least on the 
M3). less elongate snout by lack of diasiemata 
between the prenvolars as well as between Pi and 
the canine. Wabuiacinus ridei is similar in molar 



368 



MEMOIRS OF THE QUEENSLAND MUSEUM 



size to T. macknessi, but lacks 
an anterior cingulum on M'. 

Wabulacinus ridei sp. nov. 
(Fig. 1) 

ETYMOLOGY. For David Ride 
for his long-term commitment to 
Australian vertebrate palaeonto- 
logy. 

MATERIAL. Holotype. 

QMF16851, right maxillary frag- 
ment containing M * "^ (Fig. 1 A-C). 
Paratype. QMF 16852 left dentary 
fragment with broken M3 (Fig. 
ID- F) from early Miocene (Sys- 
tem B) Camel Sputum Site, 
Godtheip Hill, Riversleigh. 

DIAGNOSIS. As for genus. 

DESCRIPTION. Maxilla 
partly preserved. Infraorbital 
foramen enclosed within the 
body of maxilla, above the pos- 
terior alveolus of P^. 

Buccal crown of M^ length 
exceeds anterior width. 
Metacone largest cusp fol- 
lowed by paracone, protocone 
and St E. No other cusps. 
Postmetacrista longest crest, 
curving buccaily at the poste- 
rior end. Preparacrista orien- 
tated almost parallel to the 
tooth row, terminating at the 
anterior tip of the crown. Pre- 
metacrista and postparacrista 
connecting as a straight cen- 
trocrista which parallels the 
preparacrista. Lacking pre- 
protocrista, postprotocrista, 
protoconule, metaconule, sty- 
iar shelf or stylar cusps anterior 
to St E. Buccal flank of crown 
forming continuous slope from 
paracone and metacone to low- 
est buccal edge of the crown. 
Protocone small. 

M- similar to M' except : St 
E minute. Stylar shelf region 
high, of many tiny indistinct cusps and crests, 
especially on the more posterior half of the 
crown. Postmetacrista longest crest on the crown, 
followed in declining length by preparacrista, 
postprotocrista, preprotocrista, postparacrista 




FIG. 1. Wabulacinus ridei A = QMF16851 (M' and M^) lingual view. B = 
QMF 16851 (M^ and M-) buccal view with infraorbital foramen arrowed. 
C and C' = QMF1685 1 stereo occlusal views. D = QMFl 6852 (P2 and M3) 
buccal view. E = QMFI6852 (M3) lingual view showing small metaconid 
(arrowed). FandF' = QMFl 6852 stereo occlusal view. 

and premetacrista. Postparacrista and pre- 
metacrista forming a wide angled centrocrista. 
Postmetacrista leaving metacone almost parallel 
to the premetacrista, curving buccaily. Pre- 
paracrista straight, connecting to the postpara- 



TWX)NEW EARLY MI(X:E^fETHYLAClNES FROM REVERSLEIGH 



^ 



crista al appmximately 9iT and oblique K> the 
looTh row No Si B present. Trigon basin wider 
than on M'. Lingual flank of irigon basin *V*- 
shaped with a distinct ridge running vertically 
down its centre. The prcprolocrisia and 
posiproiocrista prominent with a minute pro- 
loconule and melaconule. Ectoflexus on the buc- 
cal surface of this looth slightly developed. 
Anterior cinguJum tenninaiing anterior to base of 
the paracone. lacking a notch. 

Menial foramen under the anterior root of P2- 
AJvcoli lorFi-?, Mi-2and the anterior root of M4. 
Mionly molar present. Symphysis beginning ad- 
jacent to the anterior root of P?. No diasiemaia 
between alveoli. All alveoli pairs orientated par- 
allel to the loulh row except Pi oblique, indicating 
sonic crowding of P| against the canine. Alveoli 
si/e indicating relative lengths of P3> P2> Pi, 
M^=M:> Ml. 

M^ with cusps in decreasing height paxaconid. 
melaconid, hypoconulid, hypoconid. All cusps 
prominent except tiunute melaconid; enioconid 
and related crests absent. Protocristid longest 
cicst, followed (in decreasing length) by the 
poslhypocrisiid and crislid obliqua. Remains of 
the metacrisiid connect to the Mtmll melaconid. 
Small lalunid basin open on die lingual side, 
Hypoconid almost medial to the irigonid basin. 
Poslhypocrisiid and cristid obliqua orientated 
oblique to the dcniary, meeting al the hypoconid 
at rigbi angles. Anterior cmgulum conlinuing 
buccal ly past the anlernbuccal corner of tooth, 
with wide notch. Posterior cingulum poorly de- 
vcloped, a small bulge in the enamel. 

Ngamalacinus gen. nov. 

*rVPE SPECIES. Nf^anuxiacinus timmulvaneyi eC sp. 
nov 

BTYMOLOGY. Wanyii Nsianuila. died oul; Greek 
kyhos, dog. Masculine. 

DIAGNOSIS. Moderately specialised among 
thylacinids in the reduced conules, reduced stylar 
shelf, anteropo&teriorly elongated molars. Re- 
taining small St B and D, melaconid, enloconid 
and hypoconulid. 

COMPARISON, hl^amahdnus differ?^ from N. 
dickwfti in its larger si/ie, reduced meiaconules 
and protoconulcs, reduction of Si D particulariv 
on M-. 

Mgofmilacinus dili'crs (roni W. riilei and Thy- 
hwinus in its; smaller si/e; narrower angle of 
crests at the paraconcs metaeone and proiocone. 



niirrower angle of ccnlrocrista; less reduced siy lar 
shelf with retention of prominent Si B, St D and 
Slylar shelf crests on M' and M-; less reduced 
talon basin, particularly on M'; lcssanlcri)postef- 
ior elongaiion of the molars and as.MKiaicd crest 
lengths; larger talonids: and larger metaconid 
(larger than ihe paiacooid) and with a disliiwi 
melacrislid. 

NgamaUuinus further differs from W ndei\x\ 
the more posterior position of the infraorbital 
foramen, presence of an entoconid on the lower 
molars and smaller hypoconulid. 

Ngamalacinus timmulvaneyi $p. nov. 

(Figs 2.3) 

ETYMOLOGY. For Tim Mulvaiicy. a long^ilne sup- 
porter of research at Riversleigh. 

MATERIAL Holotype QMF 16853 right dentary with 
Ml-4 (Fig. 2| from early Miocene (System B) Inalicy- 
ance Site, Godlhelp Hill. Riversleigh. Paraivpe.* 
QMF30300 left maxillary wiih P^-M? (Fig. 3A-C). 
from early Miocene (System B) Camel Sputum Site. 
Godlhelp Hill. Referred specimen QMFi6855, right 
M- (Fig. 31)), from the type locality. 

DIAGNOSIS. As for genus. 

DESCRIPTION. All articulating surfaces of 
dentary broken. Coronoid process rising from the 
ramus al approximately 1 20^. All four molars and 
the poslcrioi alveolus of P-^ present. No dia&lc- 
maia between these teeth. Degree of eruption of 
M4 indicating ajuvenilc. 

Protoconid of Mi tallest cusp, followed (in de- 
creasing height) by melaconid, paraconid, 
hypoconid, hypoconulid andcntoconid. All cusps 
distinct, with crests. Paracnslid lon^'esl crest cm 
crown followed (in decreasing length) by 
posihypocristid, melacrislid. cristid obliqua, pre-, 
postcntocristid. Anterior cingulum with a very 
small noich. Pat:4trrislid almost straight with a 
very wide angle connecting the paraconid and 
protoconid Talonid basin entirely enclosed by 
crests, large and deeply concave lo CA^nual poinu 
Hypoconid more buccally positioned than pro- 
tocomd. Cristid obliqua conlinuing Up the poste- 
rior wall of tlie protoconid. Posterior cingulum 
distinct, uniform in thickness 10 the base of the 
crown, with a slight tJOich formed between it and 
the hypCKOnulid 

M> same as Mi except: Metaconid relatively 
large, distinct, taller than the paraconid. All 
cnsiids higher and dislirtci. Anterior cingulum 
broader and Ihe notch mure distinct. Angle al 
cfCslAon the proTOconid approximately lOO-LIty. 



370 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Paracristid and metacristid longer. Metacristid 
straight; paracristid changing orientation at the 
valley between the paraconid and protoconid. 

Ml same as M: except: the anterior half of the 
crown thicker than the posterior because o[ the 
more lingual position of the paraconid and 
metaconid. Paracristid and metacristid elongated. 
Hypoconulid and enlex-'onid slightly more to pos- 
terior, with entoconid slightly smaller than on 
M:. Posthypocrisiid bending poslcnoriy to con- 
nect to the posterioriy positioned hypoconulid. 
Paracristid proportionally longer than on M:. 

Ma same as M3 except: Talonid basin reduced, 
well defined and enclosed by crests- Entoconid 
minute; hypuconid small; hypoconulid highest 
cusp on talonid. Small posterior cingulum pres- 
ent. 

No obvious sutural boundaries on the maxilla 
except a posterior suture that may have connected 
to either the jugal or the lachrymal. Maxilla indi- 
cating that the canine was large, its root extending 
deeply into the maxilla. Infraorbital foramen 
above M:. The region immediately posterior to 
the intraorbital foramen damaged but a depres- 
sion in the maxilla here and sutural boundaries of 
the jugal indicate that the jugal is likely to have 
contacted the exiemal opening of the infraorbital 
canal. Maxilla with large extension projecting 
back towards and contributing to the zygomatic 
arch. No maxillar>' palatal vacuities in the region 
of the premolars. 

Small diastemata between the upper premolars. 
P- triangular in lateral view with tioth an anterior 
and posterior cusp, with a crest from the major 
central cusp to the posterior cusp and a less well 
defined crest anteriorly to the smaller anterior 
cusp, with posterior region wider than the ante- 
rior, with ridges extending along both sides (lin- 
gual and buccal) of the posterior cusp. P^ larger 
than P^ and similar except for: anterior and pos- 
terior cusps relatively larger, anterior cusp with 
ridges off the lingual and buccal sides, posterior 
crest from the major cusp more prominent but not 
straight, posterior half of tooth relatively wider 
with enlarged crests bordering the posterolingual 
and posterobuccal edges of the crown, with an 
additional posterobuccal cusp. 

M' damaged, with anterior cingulum, a large St 
D larger than the distinct St B, a slylar crest 
running posteriorly from St D 10 the metastylar 
comer, talon broad with a possible proioconule, 
postmetacrista long and straight, crests at the 
paracone at approximately 90", preparacrista 
connecting to St B, almost perpendicular to the 
tooth row. 




FIG. 2. Ngamalacimus ummuhaneyl lower dentition. 
A = QMF16853 (dentary with M]-5) lingual \iew. B 
= QMF16853 buccal view. C and C* = QMF16853 

M- sam.e as M' with: in occlusal view 
posterolingual dimension longest followed by 
buccal length and anterior width. Anterior cingu- 
lum not notched in QMF16855 but is in 
QMF303{>0, cingulum terminating at the anterior 
face of the base of the paracone without connect- 
ing to the lalon basin. No posterior cingulum. 
Metacone highest cusp on the crown, followed (in 



TWO NEW EARLY MIOCENE THYLACINES FROM RIVERSLEIGH 



371 



decreasing height) by: paracone, St B, 
metastylar cusp(s) and protocone. 
Postmetacrista longest crest on the 
crown, followed by the preparacrista, 
premetacrista, postprotocrista, pre- 
protocrista and postparacrista. All 
crests relatively straight. Enamel sur- 
face slightly raised about the pro- 
toconule. Metaconule not present as a 
distinct cusp. Slightly raised 
postprotocrista connecting the pro- 
tocone to the metacone base where a 
sharp crest runs up the lingual surface 
of the metacone. A less distinct ridge 
running down the lingual side of the 
paracone and protocone. Slight 
ectoflexus at the buccal side of the 
crown due to bulging of enamel 
around St B. St E a raised part of the 
stylar crest. Between St E and B are 
minute cusps on QMF16855 but St D 
is more distinct on QMF30300. One 
crest connecting the metastylar 
cusp(s) to the posterolingual corner of 
the crown. Talon basin large with a 
broad, flat base. Preprotocrista and 
postprotocrista relatively low. Cen- 
trocrista at approximately 100°. 

M^ same as M- except: Ectoflexus 
stronger and all stylar cusps reduced to 
cuspules. Stylar crest not continuous 
along the lingual edge of the crown. St 
B largest stylar cusp. Anterior cingu- 
lum with less distinct notch than in M- 
of QMF30300. Preparacrista and 
postmetacrista longer; paracone rela- 
tively larger but smaller than 
metacone. Paracone more lingually lo- 
cated. Centrocrista at approximately 
90°; postparacrista strongly curved. 
Talon narrower. Protoconule and 
metaconule with ridges connecting to 
the lingual face of the paracone and 
metacone respectively. 



THYLACINID PHYLOGENY 




. Ngamalacinus timmulvaneyi upper dentition. A = 
)300 maxillary fragment with P^-M^ and showing infraorbi- 



FIG. 3 
QMF30300 maxillary 
tal foramen (arrowed). B and B' 



QMF30300 stereo occlusal view 



of P^^ and molars. C = QMF30300 lingual view. D and D' 



Ngamalacinus timmulvaneyi and W. qmfi 6855 (M-) stereo occlusal views 
ridei do not share any apomorphies 
that are not also found in Thylacinus. 
These, therefore, cannot be considered to repre- 
sent members of the same genus. 

Wabulacinus ridei and Ng. timmulvaneyi have 
combinations of features that place them between 



plesiomorphic N. dicksoni and apomorphic Thy- 
lacinus but do not form an independent dichot- 
omy (Fig. 4). Neither species can be placed in a 
known genus because: neither shares any 



372 



MEMOIRS OP THB QUEENSLAND MUSEL^M 



TABLE 1. Character and eUics aoiong tfaylacicies. 

1. Infiaorbical foramen: 0. not bound by jugal. 1 bound 
by jvgiU. 

2. Cenlzocristii. I . angled. 2. szraighc (as indxated by 

3. Prcparacrtsu on M . 1 . angled almost perpendicular 
CO the u>oth row axis. 2. wider soigk than state 1 . 3. 
straight 

4. Angle of crests at paraoone and tnxitaconc. L wider 
than on plesiamarpfiic das>'urids. 2. TurUieT widened. 

5. Entocordd- L smaU. 2. minute, a. either absent or 
posteriorly positioned and combined with the 
hypoconulid. 

6. Hypoconulid size. 0. large. 1 . reduced. 2. minute. 

7. Stylar shelf size. 1. crests and cusps present but 
reduced compared to plesionorphic dasyurids. 2. re- 
duction in size of some cusps and crests. 3. further loss 
of cusps and crests (mostly absent on M ). 4. complete 
loss on crests, only a single small cusp present on the 
posterior of the ctowti. 

8. AntcriorctnguIum.O.completeonM .l.inoovnplc^c 
on M . 

9. Mciaconulc and proloconule. 0. present «id laigc 1 . 
present and reduced. 2. further reduced or absent 

10. Mctaconid size. 1. reduced compared lo 
plcsiomorphic dasyurids. 2. small. 3, absent but retcn- 
sion of crest anangement in poslenor molars. 4. com- 
plete absence of cusp and associated crests. 

1L Talonid basin size. 0. unreduced 1. reduced by 
ling^ial placement of h>*poconid. 2. further reduction. 

12. Talon size. 1 . rejiuciion of talon width compared to 
plcsiomorphic das>'urids with associated lengthening. 
2. loss of metaconid and further widening of the crests 

13. Diastemata and size of SU. 0. no diastemata in 
premolar region, M4 shorter than M3. 1 . diastemataand 
M4 equal in length U> M3. 2. diastemata and M4 is 
longer than M3. 



apomoiphy with N, dicksoni that is not also 
shared with Tkylacinus: lo include cither in Thy- 
lacinus would expand it beyond any other 
dasyuromorphian genus. Wabulacinus ndei 
shows character conflicl in the plcsiomorphic 
nature of the infraorbital foramen which is more 
plesiomorphic than in N, dicksoni and Ng, 
timmuhaneyL This character may have under- 
gone reversal in W. ridei. 

The jdngle most parsimonious Uiee of thylacmid 
relationships was found using an Exhaustive 
Search PALT 3.1 (SwotTord, 1993) with 13 or- 
dered characters (Tables 1 & 2) using 
pletsomorphic dasyurids as the outgroup. Each 
laxon represents the sister species of all ihyla- 
cines immediately to ils right. In general, the 



TABLE 2. Character state dLstribulian among Ihyla- 
cines. {a = either or aut^morphic combination of 
entoconid and hypoconulid. ? = unknown saaic). 



Dasyurids 


OOOOOOOXIOOOO 


Nimhacinus dicksoni 


inn iiooioio 


Ngamaiacintds limmulvaneyi 


1111112011010 


Wahulacinuf ridei 


0232a 03 112 12? 


Tkylacinwt nackmssi 


7222224023221 


Thylacinus poieris 


12222 24024222 


Thylacinm cyr^ct^^ius 


12222 24r:4 222 



more specialised carnivores are located on the 
right 

Wabidacinus ndei and Ng. timmuivaneyi arc 
more plesiomorphic than Thytocinus in the larger 
size of the metaconid (small on W, ndt!i and much 
larger on Ng. tvrunulvaneyi) and ilic lack of ex- 
pansion of (he premaxillary region. Both species 
(and particularly W ridei) are more specialised 
than N. dicksoniirt die reduction of ihe stvlar shelf 
and the roetaconule and prDtoconule, lalon basm 
and degree of ectoflexus on NP. 

WABULACfNUS RIDEI, Features dial are more 
apomorphic than in N. dicksoni and are syn- 
apomorphic with Thylacinus are: the straight cen- 
trocrisia; the widened angle of ihe preparacrtsta 
relative to the postparacrista, particularly on the 
M^ where this crest is parallel with the an- 
teroposterior dimension of the tooih; an increase 
in Ihe size of the angle formed by crests at ihe 
pa/aconc and meiacone. diereby increasing over- 
all tooth length; fuilher reduction in si/e of the 
stylar cusps than that seen in either Ng. 
lifyvmdvaneyi and N. dicksoni; reduction m size 
of liie entoconid; reduction of the metaconid 10 a 
minute cusp; reducuon in size of the talonid basin 
by the mwe Ungual position of the hypoconid; 
and reduction in size of the talon basin. 

Wabulacinus ridei exhibits some au- 
tapomorphies not seen in any other thy lac in id » 
some of which are considered specialisations be- 
yond thai of T. cynocephalus. The preparacrisia 
on M' of W. ridel is parallel with the toodi row 
and the ccntrocrista. The preparacrisiae on .V. 
dicksoni and Ng. timmulvaneyi show the 
plesiomorphic state similar to most dasyurids in 
wtiich itliesalmostpcrpendiculaiio the tooth row 
and i'oims almost a 90"" angle with respect to the 
postparacrisia. The morphocline otherwise 
shown in thylacjnes from .V, dicksoni through to 
jf. cynocepkalus is a widening of ihe angle at 
which these crests meet (Fig. 5). This elongates 
the uxHh in an anteroposterior direction and pro- 



TWO NEW EARLY MIOCENE THYLACINES FROM RIVERSLEIGH 



373 



§ 



ID 
fiD 



□ plesiomorph>c sUto 
m 1st intermediate state 
B 2«^t intermedidte state 

■ 3rd intermediate state 

■ mt»Mt apomopphic slate 
^ unttrlaint}', we text 



3 m 

ft p 
7 ^ 

8D 

nD 

13 a 



FIG. 4. Cladogram of ihylacincs showing character 
state changes. Cladogranri is the single most parsimo- 
nious iree or n steps (CI = 0.906. HI = 094, RI = 
0.9 1 7. RC == 0.83 1 ). Striped box = unknown state of 
eitlicrplcsiomorphic or highly derived. For characters 
and their distribution see Tables 1 &. 2. 

duces an enlarged longitudinal blade formed 
from the posimetacri&ta. cenlrocrisla and pre- 
paracrisia, Only on M* of W. ridei doej. the pre- 
paracrislii lie parallel to the tooth row, a condition 
more derived than that in any other thylacine. The 
lalon basin on the M' of W ndei is also more 
derived in its degree of reduction ihan that of 7: 



macknessi but is similar 10 the condition in T. 

cynocepbalns. 

The anterior cingulum on M' of W. ridei is 
reduced compared to that of N. dicksoni (it is 
unknown in Ng. timmidvaneyi). In W. ridei it is 
incomplete while in N. dicksoni it continues lin- 
gually to join the talon basin. This feature is more 
plesiomorphic than in T. cynocephalus where the 
cingulum is lost, but more specialised than in T, 
nmcknessi where a complete cingulum is re- 
tained. In addition, the anterior portion of M' of 
W. ridei is unique in that the anterior root lies 
much further forward under the crown than itt 
other ihylacines. 

Another trend in ihylacines is for the entoconid 
to become reduced. Only in W. ridei is this cusp 
completely lost. 

WabLdacinits ridei h autapomorphic within the 
family in having an enlarged hypoconulid. In 
oihcrlhylacines the hypoconulid shows reduction 
(e.g., in N. dicksoni, T. cynocephalus) and may 
also move posteriorly (e.g., in T macknessi). Th\s 
enlarged cusp in W. ridei may be compensate lor 
loss of the entoconid, or alternatively, it may 
represent a combination of the hypoconulid and 
a more posteriorly placed entoconid. 

A feature previously used to distinguish ihyla- 
cines from dasyurids is the posterior position of 
the infraorbital foramen posteriorly delimited by 
thejugal(Muirhead& Archer, 1990). It is known 
in T. cynocephalus. T. potens, Ng. timmulvaneyi 
(Fig. 3A) and N. dicksoni. Wahulacinus ridei has 
the infraorbital foramen anterior to M' and well 
distant from the jugal (Fig. IB). This position is 
similar to dasyurids in which it most frequently 
occurs above M'/M"^ (e.g., in Dasyurus and An- 
techinus). The anterior position of this foramen 
in these dasyurids indicates that posterior place- 
ment near the jugal in most thylacines is 
apomorphic (Archer, 1976). The anterior place- 
ment of the jugal in IV. ridei is therefore 
plesiomorphic relative to other ihylacinids. 

Wabutocinus ridei is plesiomorphic in many 
respects to Thylacinus excluding it from Thy- 
lacinus. W, ridei has a number of features unique 
among ihylacinids placing it outside the range of 
variation within Thylacinus. 

NGAMALACINUS TIMMULVANEY!. This spe- 
cies shares with H'. ndei and Thylacinus the 
apomorphic reduction in the slylar shelf com- 
pared to N. dicksoni (Fig. 4). This" includes reduc- 
tion in size of St D of M- On QMFI 685.S, Si D 
is further reduced and replaced by a number of 
minute cusps thai border the stylar shelf On other 



374 



MEMOIRS OF THE QUEENSLAND MUSEUM 



molars, size of the stylar shelf h comparable lo 

thai in N. dicksonL 

The protoconules and mctaconulcs al* W. ridet 
are upomorpliically reduced compared to those of 
N. dicksoni. The lalon basin is iilso slightly nwre 
reduced than that o^ A/, dicks^tii. This species 
I'urihcr dilFers from /V. dkksoni in |h^ less ex- 
treme cctoflexus, an apomorphic feature. These 
specialisaiions of N^. timmulvaneyi compared to 
N. dicksoni arc less marked than the degree of 
specialisation of these same features m W. rUlei 
and Thylaciniis. Ngamalacintis riwmulvanevi and 
N. dicksoni share several picsiomorphies and. in 
terms of overall similarity. Ng. ummuhwwyi h 
much closer to ."V dicksom (han to any other 
thy lacinid(Fig.4i.Thcsciwo species <lotvot^hare 
any apomofpl>y not also found in other thyla- 
cines. 

PALAEOECOLOG Y OF RIVERSLEIGH 

Tmi-ACINIDS 

Thylacinids described from the Riversleigh as- 
semblages arcM /lickMmi, % rhnmul\w\eyi, W. 
ridei ^TidT. fnackftessi. This diversity raises ques- 
tions about niche diversification. Allbitugh only 
one thyiacinc appears to have been present at atiy 
one lime in lale Miocene (Alcnola. T. poteny). 
Pliocene (Awe Si Chinchilla, T. vsttocephalus) 
and Quaternary (many assemblages, T vyn- 
ocephalus) local faunas of Australia and New 
Guinea (Archer. 1982: Dawson, 1982). prior to 
the late Miocene, available resources enanled the 
*thy!acinc niche" to he more finely divided. Part 
of the e\pbnaiion may be found in the a|>parent 
absence from the Riversleigh local fauniisof any 
large dasyurlds as specialised for cami vory as the 
late Cainozoic species of Glaucodon, Suno- 
p}iilns and Dasyurus. Presence ol Jar^e carnivo- 
rous dasyunds appears inversely coirelaied with 
thylaciniddiversity The subsequent rise of these 
dasyunnes may, therefore, have accompanied 
laie Miocene decline in ihylacinid diversity. 

Although there is a greater diversity of thyla- 
cines in the Oligo-Miocene Riversleigh deposits 
than later, a wider range of large carnivores was 
cdso present in these Riversleigh local faunas. For 
example in single local faunas, ihere were ofien 
3 crocodilians (P. Willis, pers. coinin.). at least 2 
large snakes (madtsoiids ^^nd pytbonids; J. 
Seanlon, pers. comm.), 2 lineages of thy- 
IrtColconids {.Wukaleo aivd a genus similar to 
PfiscitefX, Archer ci aJ. , 1 989 ). a possibly caraiv - 
omus kangaroo (Anther & Flannery. 1985; Wroe 
& Aa-hcr. 1995: Wruc. 1996) and an unknown 



number of raptorial birds (Bole^, pers. eomm.l 
Archer ei al, 1994). Hence it is probable that the 
relatively high diversity of Riversleigh thylacincs 
reflects an overall higher biotic diversity in the 
rainforests of the Riversleigh region. 

Camel Sputum is the only Riversleigh site to 
have produced more than one thylacine: A^^. 
limmulvaneyi and IV. ridei. These laxa arc very 
similar in si/e. Tiic maxilla of A^^. timmuh'aneyi 
recovered from Camel Sputum Site differs from 
the maxilla of W. ridei in the position of the 
infraorbital foramen (in A/^. liiimulvaneyi it typ- 
ically lies above M- and was probably i^oundcd 
by (he jugal while in VV. ridei it lies anterior lo 
M') and die more plesiomorphic structure of the 
molars in Ng, timmulvcmeyi , These differences 
cannot be accounted forby intraspecific vanaiion 
and the specimens clearly represent (wo diftereni 
species. 

it is not clear how many of Riverslelgh's thyla- 
cincs co-cxistecLM/Tiile 2 arc present in IheCamel 
Sputum assemblage, the niorc generalised N. 
divkaoni may have been present throughout the 
Oligo-Mioi.-ene iSysiems A to C; Muirhcad & 
Archer, 1990). Thylacinus macknessi in Systems 
B and C (Muirnead. 1992) suggests thai by the 
early to middle Miocene, all 4 genera co-existed 
al Riversleigh By late Miocene Alcooia lime, 
one lineage is known: T. patens. 

In late Camoz-oic deposits from other areas of 
AusU'alia (i.e., cave assemblages in eastern, 
southern and western Australia; Ride, 1964; 
Archer, 1974, 1982; Dawson. 19S2), thylacinid 
remains are common. Sites where thylacinid re- 
mams iire abundant (e.g., Thylacine Hole on the 
Western Australian Nullarbor; Lowry, 1972) 
may he mterpreted to represent lairs or traps 
where carnivores were preferentially attracted, 
perhaps by the presence of other animals. In the 
Riversleigh deposits, most of which appear to 
have accumulated in shallow pools within 
rainforest environments (Archer ct al., 1989; 
Archer et al 1994), thylacinid remains are rela- 
tively rare and therefore may more fairly repre- 
sent natural frequencies. 

THYLACINID DL'XGNOSIS AND 
MORPHOLOGICAL TRENDS 

Thylactnids differ fn;im dasyurids and other 
polyproitKiom mai^upials hy having, in combina- 
tion, the loltowing features The prcmeiacrista 
and postparacrisia join as a ccntrocrisia. The 
angle formed by ihcse crests is straight t)r almost 
straight in occlusal view in al least M- of the 



TWO NEW EARLY MIOCENE THYLACINES FROM RfVERSLEIGH 



375 



J 




Ai 



Bl 



(C^^C^^ 




-} 



ci 



Di 



<€<y}^/Z^^ 



K>7i 




FIG. 5. Upper and lower deniiiions of all known ihylacine genera showing cresi oriemalion compared lo a 
dasyund. A=Dasyunis. B=Ni/nbacintis dicksont (P8695-92) C=Ngamalacinus tlmmulvaneyi. D=Wahulacini4S 
r'ldclE^ =Th\hwinusmachiessiE=Thytaani4ScynocephaUts. Upper dentitions include P-* where known. Scales 

= 0.5mm- 



uppcr dentition. The cristid obliqua continues up 
the posterior Hank of the protoconid from the 
lalonid region rather than terminaiing at the base 
of the protoconid. This functions in elongating 
this crest and becomes more prominent as the 
meiaconid is reduced (e.g., in Thylacinus). The 
stylar cusps are reduced. This occurs most prom- 
inently on M-* but also occurs to varying degrees 
on more anterior molars. The si/.e of the 
meiaconid is reduced on all lower molars. This 
reduction is correlated with the more posterior 
placement of the meiaconid relative lo the pro- 
toconid, functioning in widening the angle of 
crests at the protoconid and enlarging the irigonid 
basin. Reduction of the meiaconid is also found 
lo progress in degree from the more reduced 
condition on anterior molars to posterior molars 
(Muirhead & Gillespie, 1995). The size of the 
lalonid basin is reduced because of the more 
lingual position of the hypoconid. This cusp oc- 



cupies much of the surface of the lalonid basin 
such thai no flat surfaces occur on the basin lloor. 

Structural morphoc lines of the family (appar- 
ent in more specialised forms) include Ihe follow- 
ing. There is an increase in the angles formed by 
crests of the paracone and metaconc, increasing 
the length of the poslmetacrisla. More anlero- 
poslerior orientation of the preparacrisia. The loss 
of extreme cctollexus particularly in M^ (related 
lo Ihc overall elongation of the teeth). A reduction 
in si/c of the protoconule and mctaconule as well 
as the entire talon basin and reduction in si/c of 
the stylar shelf All of these fealures of the upper 
dentition increase the anleroposlerior length of 
the molars with ihe entire looih row acting as a 
.system of antcroposieriorly orientated blades 
( Fig. 5 ). These are lypicai specialisations in mam- 
malian ciu-nivorcs. 

In the lower molars, the trends arc for comptcie 
lo.ss of the meiaconid and opening of the Irigonid 



376 



MEMOJRS OF THE QUEENSLAND MUSEUM 



biisin. Here, like Ihe upper molars, the lower 

molar crests become orienlaicd anteroposterior! y 
(Fig. 5). The paracristid becomes the anierior 
crest with the elongated cristid obliqua function- 
ing as the posterior crest (the posiprolocrista) 
(Muirhead and Gillespie, 1995). The lingual side 
of the lalonid is also reduced through reduction 
of ihecntoconid. 

Only in Thylacmus is the snout elongated by 
b»Hh Jiasiemata between the canine and premo- 
lars and elongation of M4 (such thai il is longer 
iliun preceding molars). Extreme posterior place- 
ment of the infraorbital foramen and partial en- 
closure by the jugal \s also a possible 
synapomorphy of Thylacinus related to snout 
elongation. 

All thylacinids plesiomorphically retain the 

paraconid on Mu remnants of posterior and ante- 
rior cingula on the lower molars and poslenor 
increase in size from Pi to P^. 

Variation among thylacinids that docs not ap- 
pear to follow these 'carnivorous trends" includes 
position of the infraorbital foramen which, 
plesiomorphically and unlike all other known 
thylacinids. is more unicriorly positioned in W. 
nde't above P^. 



ACKNOWLEDGEMENTS 



The study was underliikcn with support fmm 
the Queen Elizabeth 11 Silver Jubilee Trust For 
Young Australians and the Australian Common- 
wealth Department of Employment, Education 
and Training 

Material on which this study is based was due 
to support from: the Australian Research Grant 
Scheme; the University of New South Wales; the 
National listaie Grants Scheme, Queensland; the 
Department of Arts, Sport, the Environment, 
Tourism and Territories; Wang Australia; ICl 
Auslialia; the Queensland Museum; the Austra- 
lian Museum; the University of New South 
Wales; the Australian Geographic Society; MIM; 
Ansell Wridgways; and Surrey Beatty and Sons. 

I am grateful for the advice and suggestions of 
refetecs Ken Aplin and Mike Archer and for the 
assistance given by the Australian Museum (m 
paniculai l.inda Gibson), the Northern Tcrnit>ry 
Museum (in j^articular Dr Peter Murray), Robyn 
Murphy. Anna Gillespie, Henk Godthclp (Uni- 
versity of New South Wales). 



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FLOWER. W.H. 1869. Remarks on the homologies and 
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MUIRHEAD, J. & GILLESPIE, A. K. 1995. Additional 
parts of the type specimen of Thylacinus 
macknessi (Marsupialia: Thylacinidae) from 
Miocene deposits of Riversleigh, Northwestern 
Queensland. Australian Mammalogy 18; 55-60. 

RIDE. W.D.L. 1964. A review of Australian fossil 
marsupials. Proceedings of the Royal Society of 
Western Australia 47: 97-131 . 

SWOFFORD, D.L. 1 993. PAUP (phylogenedc analysis 
using parsimony). Documentadon for Version 
3.1 . (Illinois Natural History Survey: Campaign). 

WOODBURNE. M.O. 1967. The Alcoota Fauna, Cen- 
tral Australia: an integrated palaeontological and 
geological study. Bulletin. Bureau of Mineral Re- 
sources, Geology and Geophysics, Australia 87: 
M87. 

WROE, S. 1996. An investigation of phylogeny in the 
giant extinct rat kangaroo Ekalladela (Pro- 
pleopinae, Potoroidae, MarsupiaIia).Joumal of 
Paleontology 70: 681-690. 

WROE, S. & ARCHER, M. 1995. Extraordinary 
diphyodonty-related change in dental function for 
a tooth of the extinct marsupial Ekaltadeta ima 
(Propleopinae, Hypsiprymnodontida£). Archives 
of Oral Biology 40: 597-603. 



APPENDIX 
All measuremenis are actual distance between 
cusps except those with *(horiz)' for which mea- 
surements were made from a horizontal plane 
above the cusps (occlusal view). 





Wabulacinus ridel dentiuon (mm) | 


QMF16851 
OMF16852 


M' 


M^ 


M. 


Para-meia 


3.56 


3.59 


5 Of. 


Meia-proio 


4.20 


5.01 


- 


Proio-para 


2.64 


3.53 


- 


Anicrior/width 


5.20 


6.60 


5.20 


Buccal/1 engih 


9.00 


8.92 


9.40 


Poslerolingual /uppers 


7.37 


8.36 


- 


para -m eta (horiz) 


2.90 


3.40 


4.36 


meu-prou> (horiz) 


2.35 


3.55 


, 


proto -para (horiz) 


2.35 


2.85 


_ 


hypo-hypoconuUd 


. 


- 


- 


hyp-cnlo 


- 


- 


1.81' 




Ngamalacinus ttmmulvaneyi upper dentition (mm) 


QMF30300 

QMF16855 


p2 


P^ 


M* 


M- 


M^ 


M^ 


Para-meia 






3.64 


3.66 


3.3^1 


3.03 


Meta-proco 






2.86 


3.36 


3.65 


5.00 


Prolo-pani 






3.77 


5.14 


5.02 


. 5.52 


Anterior/width 


2.25 


4.26 


4.90 


7.68 


7.2f 


1 8.25 


Buccal/lengih 


5.56 


7.92 


s.n 


9.10 


8.7t 


> 8.17 


Posterolingu /uppers 






8-15 


10.29 


9.3f 


1 10.27 


para-meta (horii) 






3.30 


3.42 


3.31 


2.80 


meia-pioio (horiz) 






2.28 


3.05 


2.84 


3.04 


proio-para (horiz) 






3.34 


3.25 


3.35 


3.20 




Ngamalacinus timmulvaneyi lower dentition (mm) 


QMF 16853 


Ml 


M2 


M3 


M4 


Para-meu 


3.36 


3.97 


4.53 


4,37 


Meia-proio 


. 


3.09 


3.35 


3-47 


Proto-para 


. 


4.11 


4.92 


479 


Amerior/widih 


3.34 


4.21 


4.25 


4.78 


Buccal/length 


7.69 


8.37 


870 


8,24 


para-meia (hon/) 


3.62 


3.98 


4.50 


4_27 


mela-proto (hori?) 


- 


2.51 


2-64 


2.43 


proio-pars (horiz) 


- 


3.88 


3.89 


4,03 


h ypo - hypocon ulid 


2.05 


2.26 


2.10 


1.34 


hvp-cnto 


0.76 


108 


1.13 


0.90 



KUTERINTJA NGAMA (MARSUPIALA. ILARIIDAE): A REVISED SYSTEMATIC 

ANALYSIS BASED ON MATERIAL FROM THE LATE OLIGOCENE OF 

RIVERSLEIGH. NORTHWESTERN QUEENSLAND 

T..I. MYERS AND M. ARCHER 

Myers. TJ. & Archer, M. 1997:06:30. Kuierintja ngama (Marsupialia, llariidae); a revised 
systematic analysis based on material from the Ute Oligocene ot" Rivcrslcigh, northwestern 
Queensland. Memoirs of the Queensland Museum 41(2): 379-392. Brisbane. ISSN 0079- 
8JC35 

Tlie Riversleigh ilariids come from the late Oligocene White Humer Site and are Kuierintja 

n^'ima Pledge. 1987. Molar cusp morphologies are compared with those of other iiariids and 
vombatiforms and several morphoclines identified. The range of variation is similar ld that 
m P/7ii5c<?/urf 105 t'i//^rfM5. Cladislic analysis suggests several hypotheses about inirafaminal 
relalionships: I ) Ku. ngamo is an ilariid; 2) Koobor us not an ilariid; and 3) ilariids form a 
monophyletic clade with the wynyardiids, although ihe relationships of ihese laxa to other 
vombalomorphians are not resolved. □7/an(Waf, Oligocene, Vombatijormes, While Hunter 
Site, Riversleigh. 

TJ. Myers & M. Archer. School of Biohsica! Sciences. Universiry of New South Wales. 
Svdnew New South Wales 2052. Australia: received 10 November 1996. 



Ilariids are extinct marsupials discovered in late 

Oligocene deposits of central Australia. Ilaria 
includes /. illumidens and /. lawsoni (Tcdford & 
Wuodburne, 1987); Kurerintja contains Ku. 
ngama { Pledge, 1 987). There is also some contro- 
versy concerning the placement of ^^(7o/?orwj(.bin 
the Phascolarciidac because Pledge (1987) sug- 
gested (Hal Ku. ngama may have been ancestral 
to Ko. jimbarratii, making the iaticr a potential 
ilariid. Tcdford & Woodburnc ( 1 987) found sim- 
ilarities in upper dentition between 7. ilhmiidens 
and Koobor, namely: a paraconule on M', no 
paraconule or neometaconule on M-' or M-^, but 
considered them symplesiomorphic, concluding 
that Koobor shared more synapomorphies with 
phascolarclids than with ilariids. 

We review Kuterintja ngama based on material 
from the While Hunter Local Fauna at 
Riversleigh, NW Queensland. White Hunter Site 
on HaFs Hill, on the D-sile Plateau (Archer el al., 
1994; Creaser, 1997) was quesljonably assigned 
to eariy Miocene System B (Archer el al., 1989, 
1 994) but the fauna now suggests laie OHgocene 
System A. A lenuuivc corrclaiion is made of 
While Hunter Local Fauna with the Ngama Local 
Fauna of the Etadunna Formation al Lake Pal- 
ankarinna. South Ausiralia. 

Pledge (1987) observed that Kuterintja ngama 
differs from flaria in being smaller, having larger 
cusps, pre- and poslcrislae on ihe siylar cusps, 
postprotocnsla and premetaconulecrisla sepa- 
rated by a crevice, and an anterior cingulum di- 
vided by a stronger preprolocrista. Similarities to 
f. illumidens include a selenodonl structure and 



well-developed buccal stylar cusps. Pledge 
(1987) described the hololype (S.\M P24539) of 
Ku. ngama as a LM"*. However, material from 
Riversleigh suggests that the holotype is a LM\ 

SYSTEMATICS 

Material is deposited in the South Australian 
Mu.seum (SAMP), and the Queensland Museum 
(QMF). Homology of molars and the dP-^ follows 
Luckelt(1993). Homology of ihcoiher premolars 
follows Flower (1867). Cusp homology follows 
Archer (1984). Tedford & Woodburnc (1987) 
and Pledge (1987). 

Order DIPROTODONTIA Owen, 1 866 
Suborder VOMBATIFORMES Woudburrve. 

1984 
Infraorder VOMBATOMORPHIA Aplin & 

Archer. 1987 

Family ILARIIDAE Tedford & Woodburnc. 

1987 

Kuterintja Pledge, 1987 

TYPE SPECIES. Kuterintja ngamaPliidgc. 19S7 from 
laie Oligocene Etadunna Formation at Lake Pal- 
ankarinna. northern South Australia. 

DIAGNOSIS. Relative to Ilaria: Small, lack- 
ing iransvcrse li nking crests on Ihe check teelh. Ii 
with low, almost horizontal inclination, dorsally 
flattened, transversely compressed and with an- 
terior portion intlecled. 

P3 subrectangular. with I large anterior cuspid 
and two .smaller posterior cuspids only slightly 



38a 



MEMOIRS OP THE QUHENSLAND MUSEUM 



tracts^ ctic VBlley 



(ll-i) cU»H*, 



ptulocoDJd 




byitocon"t 



etitocDoid 



MWCUQI i-Hi^itiluin 



FIG, 1. Kmerimja n^ama. OMF20810. 2330Ci. left 
dentary (P3 -M4J. 

separaled. one in a posterolingual position, other 
in a posteromedial posiiion longitudinally 
aligned with the anterior cuspid. M] subovaic, 
with anterior cingulum medially intlected and 
less developed, with lingual faces of the buccal 
cuspids near vertical, with less developed prc- 
prolocristid and posthypocnsiid, with prc- 
protocristid and posthypocnsiid lenminating in 
line with the 'cenlrar cuspids, with small lingual 
basin on the hypolophid; 'cenirar cuspid on the 
protolophid in transverse ahgnment with the lin- 
gual and buccal cuspids on M1-M3; metaconid 
separated slightly from the 'central* cuspid of ihe 
paKolophid; M: with 'centrar cuspid on die pro- 



tolophid and hypolophid of similar widths and 
more closely linked, with 'central' cuspid on the 
hypolophid not linked posteriorly to the en- 
loconid: M: and M3 with preprotocrisiid and 
posthypocnsiid not extending as far linguuljy; M3 
with lingual basins less developed, with 'central' 
cuspid on the hypolophid greatly reduced, with 
posterior cingulum relatively small, with 
postprotocnstid and prehypocrisiid not blocking 
trans verse valley; M4 with compressed posterior, 
with 'central* cuspid not distinguishable on the 
hypolophid, with the postprotocrisiid and pre- 
hypocrisiid poorly developed (Fig. 1). 

P^ subovate, much wider both anteriorly and 
posteriorly compared to the P3, with naiTow an- 
terior portion, with large cusps, iri-cusped, lack- 
ing the posterobuccal cusp, with cusps subequal 
in height, with twinned central cusps separated by 
a larger tmugh, with a Uirger crevice separating 
posteromedial and posterolingual cusps, with an- 
terolingual cingulum, with well-developed rib 
running from the apex of posteromedial cusp to 
Ihc posterobuccal edge of the posterior cingulum. 

M'""* with nearly vertical buccal surfaces on 
cusps. M^ with stylar cusp C almost as large as 
the paracone. with stylar cusp D as large as the 
metacone. with buccal border slanting sharply 
posierobuccally, with posterior cingulum around 
convex structure, with all cusps subequal in 
height; M- with stylar cusp C relatively small, 
with the cristae forming the borders of the buccal 
basin on the paracone strongly developed, witti 
the poslparacrista separated from stylar cusp C, 
with stylar cusp E greatly reduced, with pre- 
prolocrista strongly developed and dividing the 
anterior cingulum; M- and .W with lingual cusps 
transversely aligned with the buccal cusps; M- '^ 
with the anterior portion of the \ooi\\ litrgcr than 
the posterior; M* with stylarcusps B und C equiv- 
alent in height to the paracone, with the buccal 
basin on the paracone enclosed at its buccal mar- 
gin, with stylar cusp D larger, M-^ and M^ without 
stylar cusp E; M"^ with the lingual half of the 
transverse valley inHecied less towards the 
posterolingual comer, with stylar cusp C vari- 
able, with metaconule variable in position, and 
thus the lingual basin variable in size; (Fig. 4). 

COMPARISON: Kuterintja ngama differs from 

phascolarctids in lacking a paraconule and 
neometaconule, having longer molars, simpler 
selenes, separation of buccal selenes. better de- 
veloped stylar cusps, a strongly developed trans- 
verse valley, poorly developed postprotocrista 
and prcmetaconulecrisla, a proiocone that is more 



KUTERINTJA NGAMA FROM RiVERSLElGH 



3SI 




FiG. 2. Kuierinrjcuigiwm. A, QMF3C«:i57, R.NV, occlu- 
sal view, stereo pair B. QMF3 1 299, RM"^'**. occlusal 
View, stereo pair. C, QMF3I301, RI1-M2. buccal 
view. 

compressed relative lo the meiaconule t)n M'- 
M^. significant separation of slylar cusps C and 
D, no protoslylid, a lingually convex metaconid. 
a proioconid that is larger than the melaconid, 
lingual cusps that are nol compressed towards 
each other, larger crown height, a well -developed 
posteroJingual cusp on P^, no posterobuccal cus- 
pid on P3, central cuspid, having a posierolingual 
cuspid on P^, a non-bladed P.^ or P-^. no longitu- 
dinal valley, and a bulbous P^^. 

KiL ngamci is distinguished from Koobor by its 
larger slylar cusps, higher crown, larger molars, 
and continuous crest between protocone and 
meiaconule (Pledge. 1987). Other differences in- 
clude, i) more conical stylar cusps; 2) lower 
selene angles on the buccal basins of the upper 
molars; 3) Koobor lacks a lingual basin on the 
iransverse valley; 4) Koobor has a poorly devel- 
oped anterior cingulum; 5) the absence of a pos- 



terior depression on the meiaconule, as exists on 
most ilariid molars; 6) A'ootorhas molars which 
are slightly compressed lingually; 7) a much 
wider and longer longitudinal valley exists in 
Koobor, 8) more poorly developed postproto- 
crista and premetaconulecrisla; 9) a protocone 
thai is compressed longitudinally relative to the 
mctaconule on M^-M'\ 1 0) no paraconulc on M'; 
1 1 ) Koobor lacks the posterolingual cusp on P-^ 
and 12) Koobor has an elongated, rather than 
bulbous, P-\ 



Kuterintja ngama Pledge, 
(Figs 1-5,7) 



987 



MATERIAL. Hololype SAMP24539. LM \ presumed 
to he a left M"^ by Pledge {19S7) from the saddle 
between Mammalon Hill and main escarpnieni, NW 
corner o\ Lake Palankarinna. 100km N of Marree, 
South Australia m the late Oligoccnc (Woodbunie el 
ak, 1993) Ngama Local Fauna within the Eiadunna 
Formation. Other malcnal.QIVlF3 1 302, a right demar> 
fragment containing P"^, Mi and M2; QMF23306, 
QMF208 10. u left denlar> with all cheek teeth and the 
alveoli for 1 1 ; QMF3 1 30 1 , anterior portion of ajuvenilc 
nght dentary. with 1 1 , dP^. and M 1 . P3 is removed from 
Its crypt, and M2 has only pari of the proioconid 
remaining; QMF17527, RM3 ^^'ith roots missing; 
QMF3I300 RM4 with the anienor portion of the 
irigonid missing; QMF3O057. RM *; QMF23203, LM^' 
with a broken anterior cingulum; QMF30058. RM-; 
QMF3 1 299, right maxillary fragment containing M-***; 
and QMF24604, right maxillary fragment with M-^ and 
M"*. and alveoli for M' and M-; QMF3()332, partial 
nght maxilla, with partial palate, anterior zygomatic 
arch, P-^, M- and the alveoli for M^ All except type 
from late Oligoccne While Hunter Site, Riversleigh. 
N W Qucenshtnd; previously regarded as possihiy Sys- 
tem A or early System B ( ^^che^ et al., 1989; Archer 
et al. 1994), late Oligoccne or early Miocene. This 
species suggests comparable age lo the South Austra- 
lian type locality. 

DIAGNOSIS. As for genus. 

DESCRIPTION. Dentary, Deepest below the 
posterior half of M3. In lateral aspect alveolus lor 
li inclined .slightly on its ventral side, hori/untal 
on dorsal side. Mental foramen at the posterior 
end of this alveolus in the dorsovcniral midline, 
and just anterior to P3, only foramen on the den- 
tary (break dorsovcntrally from the junctu:)n of 
M: and M3 may obscure olhers). 

Ii. Lower first incisor projecting horizontally 

from the dentary, curving lingually at its anterior 
(distal) extremity, suhcylindrical, transversely 
compressed, with dorsal surface transversely flat- 



382 



MEMOIRS OF THE QUEENSLAND MUSEUM 




HG. 3 . Kuterintja ngama. A, QMF208 1 , left dentary. 
occlusal view, stereo pair. B. QMF23306, buccal 
view. 

tened, with enamel from the buccal to ventral 
surfaces. 

dP3. Same as P3 except in size, tricuspid, sub- 
triangular. 

Anterior cuspid tallest, with widest base. 
Smaller cuspid posterolingually from anterior 
cuspid. Third cuspid posterobuccally from ante- 
rior cuspid and equivalent in size to the 
posterolingual cuspid. Ail cuspids closely linked, 
conical, with wide bases. 

P3 (Fig. 1). Transversely compressed, tricuspid. 
Large, subovate, conical cuspid anteriorly larger, 
taller and more broadly based than the twin cus- 
pids posteriorly. More buccal of these an- 
teroposteriorly aligned with the large, anterior 



cuspid. Posterolingual cuspid taller than its worn 
buccal counterpart. 

Thin, low cristid running from the an- 
terolingual comer of the apex of the anterior 
cuspid, anterolingually to the base of the cuspid, 
then turning posterolingually and running further 
down towards the root, then turning posterobucc- 
ally up the cuspid and terminating about half way 
up the height of the cuspid, in line with the 
posterior side of the apex of the large cuspid. 
Anterior cuspid located over the posterior portion 
of the anterior root; posterior cuspids located 
directly over the posterior root. 

A minor crevice on the anterolingual comer of 

TABLE 1. Measurements (mm) of dentition of 
Kuterintja ngama. Le=length; M w=maximum width; 
Ha=height of anterior cuspid; Aw=anlerior width; Pw 
= posterior width; Hp=height of paracone; 
Hpr=height of protocone; Hm=height of metacone; 
Hml=height of metaconule; Hprd= height of piro- 
toconid; Hmd=height of metaconid; He=height of 
entoconid; Hh=height of hypoconid. Italicised num- 
bers indicate dimension may have been lessened by 
wear. 



QM No/ID 


U 


Mw 


Ha 


Aw 


Pw 


Hp 


Hpr 


Hm 


Hml 


30332 RP^ 


8.9 


7.5 


42 


- 


- 




. 


- 


- 


30057 RM' 


9.7 


- 


- 


8.7 


7.7 


5.4 


4.6 


5.2 


4.7 


31299 RM^ 


10 


- 


- 


8.6 


8.0 


4.8 


5.4 


4.9 


5.1 


30058 RM^ 


9.4 


- 


. 


8.9 


8.8 


4.7 


5.4 


5.1 


5.1 


30332 RM^ 


10 


- 




9.5 


9.5 


7 


7 


33 


3.i 


31299RM^ 


8.4 


- 


- 


7.6 


6.5 


4.4 


5.3 


4.1 


5.1 


24604 RM^ 


8.5 


- 


- 


8.1 


7.0 


4.5 


5.6 


4.4 


4.8 


31299 RM^ 


7.2 


- 


- 


6.7 


4.9 


3.8 


4.4 


3.3 


3.3 


24604 RJVf* 


7.6 


- 


- 


6.8 


5.5 


4.0 


5.1 


3.4 


3.5 


23203 LM^ 


10 


- 


- 


8.3 


8.0 


42 


4.8 


4.2 


4.2 


31301 dP3 


3.9 


3.6 


2.8 


_ 


- 


- 


- 


- 


- 


31301 RP3 


6.7 


4.9 


5.7 


- 


- 


- 


- 


- 


- 


31302RP3 


6.6 


5.0 


4.8 


- 


- 


- 


- 


- 


- 


31301 RMi 


10 


- 


_ 


6.8 


7.4 


5.6 


5.0 


5.7 


6.4 


31302 RMi 


9.5 


- 


- 


6.5 


7.3 


3.7 


3.7 


42 


4.0 


31301 RM2 


9.8 


- 




7 


? 


6.5 


7 


7 


7 


31302RM2 


9.9 


- 


_ 


6.9 


7.2 


3.8 


3.6 


4.0 


3.7 


31301 RM3 


10 


- 


- 


6.3 


5.8 


6.0 


4.4 


4.3 


6.1 


17527 RM3 


9.7 


- 


- 


6.4 


6.0 


6.3 


4.4 


4.4 


6.2 


31300 RM4 


9.1 


- 


- 


7 


5.6 


3.8 


3.5 


3.2 


3.7 


23306 LP3 


6.8 


5.4 


5.9 




- 


- 


- 


- 


- 


23306 LMi 


9.9 


- 


- 


6.4 


6.9 


4.9 


4.5 


4.7 


5.4 


23306 LM2 


9.7 


- 


- 


7.4 


7.5 


4.9 


4.4 


4.5 


5.1 


23306 LM3 


9.4 


- 


- 


7.2 


? 


4.4 


4.9 


4.1 


5.0 


23306 LM4 


9.2 


- 


- 


6.8 


5.7 


4.1 


4.2 


3.7 


3.7 



KUTERINTJA NGAMA FROM RIVERSLEIGH 



383 



the tooth. A deeper crevice dividing the tooth into 
sub-equal halves, with the large anterior cuspid 
on the anterior side and the twinned posterior 
cuspids on the posterior side, blocked half way 
along by a crest linking the anterior cuspid to the 
posterobuccal cuspid. A shallower crevice be- 
tween the posterior cuspids blocked by a minor 
crest running from the apices of these cuspids. 

Small cristid running posteroventrally from the 
posterobuccal corner of the apex of the 
posterobuccal cuspid, turning posterolingually, 
joining a wider posterior cingulum. Posterior cin- 
gulum curving anterolingually before joining the 
base of the posterolingual cuspid. 




protocone 



Lower Molars. Subrectangular. Mi -3 subequal; 
M4 smaller. Crown heights decreasing from Mi- 
4. Tooth row curving posterolingually (Fig. 1 ). 

Ml: 'Central' cuspids on the protoiophid and 
hypolophid are neomorphs (Tedford & 
Woodburne, 1987). 6-cuspid; anterior portion 
narrower than posterior. Trigonid triangular; an- 
terolingual border inclined posterolingually; an- 
te robuccal border of trigonid inclined 
posterobuccally; both these inflections originat- 
ing from an anteromedial position of the anterior 
cingulum, at termination of preprotocristid. 
Talonid wider than trigonid. Protoconid over the 
posterior portion of the anterior root; posterior 
cuspids aligned over the central portion of the 
posterior root. Preprolocristid (or paracrislid) rel- 
atively wide, generally low, with pocket between 
the buccal margin, the anterior cingulum and the 
anterior face of the protoconid, with smaller and 
less well defined pocket between the anterior 
cingulum, the lingual margin of the pre- 
protocristid and the anterior surfaces of the 
'central' cuspid and metaconid. 'Central' cuspid 
of protoiophid with apex slightly anterior to the 
protoconid and metaconid. Anterior positioning 
of 'central' cuspid or neomorph more exagger- 
ated on the hypolophid. Both 'central' cuspids of 
similar height, lower than main cuspids. 'Central' 
cuspid on protoiophid forming a lingual basin 
with the metaconid, not totally enclosed, with 
small openings anlerioriy and posterioHy. Sim- 
ilar, small basin formed between the 'central' 
cuspid of the hypolophid and the entoconid, with 
comparable openings to its counterpart on the 
protoiophid, with anterior opening much smaller. 
A deep crevice dividing 'central' cuspids from 
the main buccal cuspids, continuous an- 
teroposteriorly, shallower in the central part of 
the tooth. Transverse valley interrupting this lon- 



stylar cusps B, C. D, E 




FIG. 4. Kuterintja ngama. A-B. QMF3O058, RM". A, 
occlusal view; B, buccal view. C-D, QMF30057, 
RM'. C, occlusal view; D, buccal view. 



gitudinal crevice wide, blocked at its buccal ex- 
tremity by a small, posterobuccally slanting cin- 
gulum linking the base of the protoconid to the 
base of the hypoconid. Thin crevice in the trans- 
verse valley preventing symmetrical postproto- 
cristid and prehypocristid (cristid obliqua) and 
postmetacristid and preentocrislid from linking. 
Metaconid and entoconid with apices steeply in- 
clined, rather than conical, with lingual surface of 
each much taller than the buccal. Entoconid 
higher than metaconid higher than 'central' cus- 
pids, with slight gradient descending from lingual 
to buccal. A thin posterior cingulum and a small 
pocket in the posterolingual corner of the tooth; 
pocket bordered by the lingual end of the poste- 
rior cingulum, with 2 crests from the postero- 
lingual and posterobuccal sides of the apex of the 
entoconid, respectively. 

In QMF31301 protoconid and hypoconid with 
lingual surfaces slightly more vertically orien- 



3«4 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 5. Kuterimjii ngoma. A. SAM ^2453*^, holoivpe. 
LM\ B. QMF24604. RM\ C. QMF?! 299, RM^ 



latcd. pcissibiy due to less wear Ihan observed in 
QMF233(Xi. 

M2 Like Mi cxcepiianleriorent! resting upon ihe 
poslerior cingulum of Mi; trigonid subreciangu- 
lar raihcr ihan triangular, due to the antenor cin- 
gulum being more Iransversely linear, anterior 



pockets formed with Ihe anterior cingulum 
smaller; trigonid and lalonid equal in transverse 
width. 'Central* cuspid on the protulophid in 
direct transverse alignment with the proloconid 
and meiaconid; *centrar cuspid on the hypo- 
lophid more to anterior; hypoconid slightly more 
poslerior. Crevice between the linked lingual cus- 
pids and the buccal cuspid shallower {possibly 
due to wear on protoconid and hypoconid ). Cus- 
pid height gradient from lingual to buccal much 
steeper (possibly due to wear). Pocket at the 
buccal end of the transverse valley larger. Cristid 
obliqua (or prehypocristid) and hypocrislid 
(posthypocristid) more developed. Lingual 
pocket on the proiolophid less well defined, with 
the openings between the mctaconid and 'central' 
cuspid larger. Small posierolingual pocket bor- 
dered by posteniocrisiid, hypocristid and a small 
posterior cingulum, with most of the latter hidden 
by M.v 

M3. Same as M2 except: anterior cingulum 
rounded. Crown height reduced; with height gra- 
dient, Metaconid and 'central' cuspid not closely 
linked, separate entities with a crevice between 
the two cuspids. Crevice of variable depth. 
'Central' cuspid on the protolophid larger, high- 
lighting an increase in size from Mi to M3. Crev- 
ice between 'central' and buccal cuspids 
shallower, decreasing in depth down the tooth 
row. Despite damage to the talonid. VenlraF 
cuspid on the hypcjlophid much reduced. E:n- 
loconid sub-equal in height to the 'central' cuspid 
on thehypolophid. transversely compressed. Pos- 
terior cingulum and posterobuccal basin much 
shorter. 

Juvenile M3 with an anterolingual basin bigger 
than in Mi or M:. paracristid terminating in lon- 
gitudinal alignment with U)e buccal side of the 
meuiconid. 

M4. Shortest and narrowest molar, with lowest 
crown, rounded subrectangular. with a very 
rounded anterior cingulum Same as M.^ except: 
protolophid and hypolophid slanting more an- 
tcrolingually. due to the buccal cuspids being 
posterior to the lingual cuspids. 

'Central' cuspid on the protolophid not linked 
to the metaconid; crevice between these 2 cuspids 
deeper. 'Central' cuspid on the hypolophid 
greatly reduced, more so ihan In M.^, further to 
poslerior. Posterior cingulum short, extending to 
the medial line of the tooth. Posierolingual basin 
greatly reduced. Transverse valley closed lin- 
gually and huccally. Lingual end of the transverse 



KVTFMNTJA NGAMA FROM RlVEKSLEIOH 



3&S 



valley curving posierulingually; buccal enclcurv- 
inji posterobuccally. Cristid obliqua and 
hypocristid rcialivcly short. Ail cuspids subcqual 
in heighl, with the meiaconid sli^hUy largci ihan 
Ihc entoconid = proUKonid and hypoconid. 

P-\ Subovatc, tricuspcd, transversely wide. Antc- 
riof poninn nariowci I ban posicrior. Cusps 3, 
larpc. subcqual in hci^'ht. Anterior and poslcro- 
mt\iial cusps lon^Mludinally aligned, separated by 
a. shallow trough. A lar^e crevice separalin*; the 
po:aeromedia! and posterolingual cusps. With 
very small anienjlingual ciagulum and larger 
posterior cingulum. A Ihin rib running from the 
apcK ofihc posteromedial cusp to the postcrobuc- 
cal edge of (he posterior cingulum. 

Upper molars. Sty lar cusps well-developed; gcn- 
cmi selcnodont cusp pattern; high crowned, with 
a general gradient towards Ihe lingual side, with 
4 major cusps (paraconc, protoconc. rneiaconc 
and meiaconule), with a siylar shelf consisting of 
slylar cusps B. CD and li. 

M* (Fig, 4). Buccal cusps of RM' nositinncd 
more posteriorly than in other molars, with 
postcfobuccal slant, wider posteriorly than antc- 
ritjrly. giving an anterolingual slant to the buccal 
border. Stylar cusp B smaller and further anterior 
than in other molars. Stylar cusp C as large as that 
on M% anterior to the postparacrista; crista not 
forming pan of the posterior face of the stylar 
cusp. Stylar cusp D largest cusp, subcqual in 
height (o the ntetacone, larger than in any other 
molar. Slyku^cusp E more developed than in other 
molars, larger than siylar cusp B. Aminorcuspule 
on the anterior of stylar cusp D. buccal lo the 
termination of the posimetacrista. larger than sty- 
lar cu.sp B, but slightly smaller than stylar cusp E. 
All stylar cusps subconical lo irianguUiT. exoeni 
postcrobuccally-aligned ridge, stylar cusp C. 
Buccal miurgm wider than lingual: blocking ere.st.s 
in the transver.se valley absent (some minor par- 
tial hlockages buecally); anterior cingulum curv- 
ing posterobuccally at its buccal extremity, 
preparacrisia orientated less transversely than in 
other upper molars; minor depressjon on the i^os- 
tciiof face of the meiaconule less developed than 
in M'^; buccal basins on ihe paraeoue and 
mctaconc poorly developed comt>iined lo oihet 
molars; posterior emgulum thinner than in other 
molars. 

M* (Fig. 4). Square. Cusp m/.cs: paraconc 
>inciacone> proiuconc = mcliiconule. Siylar 



cusp height: C>r)>B>E. Siylai cu.sp B connected 
to the paraconc by a preparacrista, and stylar cusp 
C via a postparacrista. Stylar cusp D connecting 
to the meiacone by a prcinetacrista, and stylar 
cusp E connected to the metacone by a 
posimetacrista. Buccal basin deep, formed he- 
tween stylar cusps B and C and the piiracone. The 
horiiologoiis basin on the metacone less distirvci, 
enclosed less tightly, slanting steeply 
posterobuccally towards the reduced siylar cusp 
E. Basin on the metacone deepest anterolingual 
lo stylar cusp E. Large transverse valley dividing 
this tooth in half, containing the piu-acone (ami 
associated slyku'cusps) and protoconc anteriorly, 
and the mctaconc and metaconule pi>slcriorly, 
partially blocked buecally by ati inct)mpletc cncsi 
linking prcmctacrista and postproiocrisia, 
blocked centrally by a small crest linking 
postproiocrista and premctaconulc crista, 
stopped at its lingual extremity by a very low crcsl 
linking the lingual sides of protoconc and 
metaconule (lingual cingulum). Buccal faces of 
protoconc and mcliiconule steeply inclined. «U 
most to the point of being vertical; anterior cin- 
gulum well-developed, running buecally fmni 
the pn»ioctista to the anterior side of stylar cusp 
B, and lingually from the anierolmgual corner of 
the base of the pixnocone to the protocrista; pos- 
terior cingulum .smaller than anterior cingulum, 
with the former extenditig from siylar cusp B \o 
join the postmctaconulccri.s(a; small depressioti 
on the Imgual side of the |X)simctaconulccrista 
and medial posterior base of the metaconule iper- 
haps remnant of the lingual portion of the poMe- 
nor cingulum); afl cusps over the mid-line of the 
rixrts; stylar cusps triangular, rather than mund or 
conical: buccal cusps with very I'ounil apices. 

M-^ (Fig. 5). Same as M2 except: crown lower; 4 
major and stylar cusps retaining same relative 
heights; stylar cusp E further reduced, virtually 
non-cxistcnt; posicrior cingulum less dellned; 
small pocket on the posterior side of the 
metaconule on M^ absent; lingual cusps closer to 
the anterior side uf their rcspeciivc roots. Stylar 
cusp C more to posterior than in M^, with 
postparacrista ibrming purl of this stylar cusp; 
buccal basin on the paraconc of M^ larger than in 
M-, triangular, with wider buccal edge. Trans- 
verse valley partially Nocked buecally by acresi 
linking stylar cusps C and L)> but not by a cr^sl 
linking the premetaconulecrisia and postpara- 
crista, with central and lingual blocking creus. 
Crest linking stylar cusp and the mctaconc 
larger and oiorc uniform; stylar cusp E more 



386 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 2. Characters and character states used in the 
ilariid intrafamilial phylogenetic analysis. 


Characters | States | 


1 


Stvlarcusp development 


O=poor; l=v\ell 


2 


Transverse valley on 
lower molars 


0=absent; l=moderate; 
2=well-developed 


3 


Transverse linkages 
between cuspids 


O=none; l=poor; 
2=moderate; 3=well- 
deve loped 


4 


Post protocrisia and pre 
meiaconulecrista 


=strongly developed; 
l=pooriy developed 


5 


Protocone compressed 
longitudinally relafive^to 
metaconule (on M -M ) 


0=absent: l=present 


6 


Separation of sty lar cusps 
C and D 


0=no significant 
separation; l=signiricant 
separationby large trough 


7 


Paraconule on M 


0=well developed; 
I =poorly developed; 

2=abseni. 


8 


Paraconid on Mi 


= ab.sent; l=weak;2 = 
strongly developed 


9 


Proiostylid 


0=present; l=absent 


10 


Metaconid 


O=conical; l=lingually 
convex crest 


U 


Relative heights of the 
anterior cuspids 


= subequal ; 
l=:protoconid larger than 
metaconid 


12 


Overall tooth size 


0=small; l=large 


13 


Lingual cusps 


O=compressed together; 
l=not compressed 


14 


Crown height 


O=low; Immoderate; 
2=high 


15 


Posterolingual cusp on P 


0=absent; 1 = slight 
cusp;2 = moderate;3 = 
well-developed 


16 


'1 ( uiinnk'ail 1 


O=dorsovenirally 
flattened; l=canmiform 
and conical; 2=dorsally 
flattened and distally 
inflected 


17 


Posterobuccal cuspid on 
P^ 


0=absent; l=present 


18 


'Central' cuspid 


0=absent; l=present 


19 


Posterolingual cuspid on 


0=absent; l=present 


20 


lingual closure of 
transverse valley by a 
cineulum (on upper 

molars) 


0=cingulum absent; 
l=incinient cingulum (in 
formoTcuspules); 
2=cingulum present 


21 


Bladed P3 


0=present; l=absent 


22 


Bladed P' 


0=stronely bladed; 
I =weakly bladed; 
2=absent 


23 


Posterobuccal cusp on P 


0=absent; l=present 


24 


Longitudinal valley (i.e. 
distance between lingual 
& buccal cusps / ids) 


O=weil-developed; 

1 =moderately developed; 

2=absent 


25 


p' 


O=bulbous; l=elongale 



anteriorly positioned; buccal basin on the 
metacone narrower, slanting more antcrobucc- 
ally. 

M"^. Sub-triangular, posteriorly compressed. 
Same as M^ except: crown height very small, 
with the protocone>paracone=metacone> 
metaconule. Stylar cusp B>D; stylar cusp C non- 
existent; stylar cusp E extremely reduced or miss- 
ing. Crevice between the paraconc and protocone 
transversely wider. Buccal surface of the 
metaconule and protocone far less vertically in- 
clined. Anterobuccal basin larger; buccal basin 
on the metacone absent; buccal basin on the 
paracone very shallow, slanting posierobuccaily. 
Transverse valley not blocked buccally, curving 
posterobuccally rather than being transverse, 
with lingual end enclosed slightly, by a low crest 
(i.e. the crest does not continue to the base of the 
protocone). Anterior cingulum very small. Poste- 
rior root slants posteriorly rather than vertically. 

REMARKS. Comparing LM' QMF23203 to 
RM^ QMF30057: buccal half of the anterior cin- 
gulum transversely shorter; stylar cusps B and E 
less developed; cuspule on the anterior face of 
stylar cusp D absent. M^ of QMF24604 exhibits 
variation compared to the M^ of QMF31299 as 
follows: 1 ) stylar cusp D is larger; 2) the distance 
between stylar cusps D and E is greater and 
therefore a bigger buccal basin is found on the 
metacone; and 3) the medial lingual basin is 
divided into two sub-basins at its lingual margin 
by a very small transverse crest. 

QMF24604 highlights the variability in M"^, as 
follows: posterior half not as compressed as in M'* 
of QMF31299, and therefore has a longer and 
wider posterior cingulum. Stylar cusp E is much 
reduced. Therefore the buccal basin on the 
metacone is also present and it is as deep as the 
basin on the paracone. Styiar cusp D is also more 
defined and larger than in M^* of QMF3 1 299. The 
medial lingual basin is smaller. The anterior cin- 
gulum extends further lingually to the base of the 
protocone. The transverse valley is blocked in 
two places rather than one. It is blocked buccally 
by a crest linking stylar cusps C and D, and is 
partially blocked by a small crest linking 
postparacrista and premetaconulecrisla. The crest 
partially blocking the lingual extremity in 
QMF31299 is not present. Large stylar cusp C is 
not present in QMF31299. A well developed and 
enclosed buccal basin on the paracone is absent 
inQMF31299. 



KUTERINTJA NGAMA FROM RTVERSLEIGH 



387 



TABLE 3. Ilariid inlrafamili?J data matrix 
1 &2;b = 0&l;c = 0&2 


as 


usedb> 


'PAUP 


.? = 


: fossil material missing or status uncertain; 


a = 


Taxa 


CHARACTERS AND STATES | 


1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


11 


12 


13 


14 


15 


16 


17 


18 


19 


20 


21 


22 


23 


24 


25 


Madakoala spp. 








1 














2 




















1 


? 










c 

















P. cinereus 












































I 













2 





1 


1 








Koobor notabalis 





7 


? 














7 


7 


7 


7 














? 




? 


7 


1 


7 


1 





1 





llaria illumidens 


1 


2 


2 


1 


1 


1 


1 


2 











1 


1 


2 


2 


1 




1 


1 


2 





1 





2 





Ilaria lawsonl 


? 


2 


2 


7 


? 


? 


7 


1 


1 


1 


1 


1 


? 


2 


? 


1 




1 


1 


? 


1 


2 


1 


2 


1 


Kulerinlja ngama 


1 




1 


1 


1 


1 


2 





1 


1 


1 





1 


2 


3 


2 





1 


1 


2 


1 


7 


7 


2 


7 


wyn yard lids 


1 


1 


3 








1 


2 





1 





1 





1 


1 


a 


2 














1 


2 





2 


1 



PHYLOGENETIC SYSTEMATICS 

Twentyfive dental characters with up to 4 slates 
each (Table 2) were used to develop the data 
matrix following character argumentation and 
polarisation (Table 3) for the intrafamilial analy- 
sis of ilariids. Three outgroups used to determine 
polarities are: 1) the modern Koala, 
Phascolarctos cinereus; 2) Madakoala: and 3) 
wynyardiids. The former is the most derived 
member of a primitive outgroup because the 
Phascolarctidae is the stem taxon from which the 
vombatomorphian radiation diverged (Marshall 
et al., 1990; Aplin & Archer, 1987). Madakoala 
devisi and Madakoala wellsi are employed be- 
cause of the primitive position of Madakoala 
within the phascolarctid radiation (Woodbume et 
al., 1987). Primitive and derived phascolarctids 
were used to determine the relationships of spe- 
cies of Koobor. Wynyardiids include 
Namilamadeta snideri and Muramura sp. and are 
a closer sister group of the ilariids than the 
phascolarctids, ffierefore providing a basis for 
polarising character slates within the Vombato- 
morphia. 

Character optimisation is performed after the 
character analysis has been completed and the 
most parsimonious trees found. The two op- 
timisation algorithms used by PAUP are 
ACCTRAN and DELTRAN. ACCTRAN accel- 
erates the evolutionary transformation of charac- 
ters so that changes occur at the earhest possible 
stage on the optimal tree. As far as homoplasy is 
concerned, this algorithm has the effect of favour- 
ing reversal of character stales over conver- 
gences. DELTRAN delays transformation of 
characters so that changes occur as far up the 
optimal tree as possible. This has the effect of 
favouring convergences over character reversals 
(Wiley et al., 1991). DELTRAN analyses are 
favoured here because of the large amount of 
missing character data in the matrices. 



RESULTS, the optimal tree (Fig. 6) has 50 steps; 
a consistency index (CI) of 0.800; a homoplasy 
index (HI) of 0.260; a retention index (Rl) of 
0.778; and a rescaled consistency index (RC) of 
0.622. Notably the ingroup (Koobor notabalis. I, 
lawsoni^ I. illumidens and Kuterintja ngama) did 
not form a monophyletic clade. Ko, notabalis is 
sisier taxon to the Wynyardiidae, Ku. ngama, /. 
illumidens and /. lawsoni clade. Madakoala and 
Phascolarctos cinereus formed a basal monophy- 
letic clade. 

Bootstrap analysis for the most parsimonious 
tree had the clade excluding phascolarctids and 
Koobor supported 99% of the time. The ilariid 
clades, excluding and including Ku. ngama, oc- 
curred 78% and 95% of the time respectively. 

Removal of the wynyardiids as an outgroup had 
no effect on the topology in the optimal tree. A 
bootstrap analysis on data excluding the 
wynyardiids found the clade containing /. 
illumidens and /. lawsoni to be supported 62% of 
the time, slightly lower than in the previous anal- 
ysis. While the clade including all 3 ilariid species 
was supported on all occasions. 

Another method of testing support for the opti- 
mal tree is to examine the frequency and topology 
of the *next best' trees (Simmons, 1993). PAUP 
evaluated 945 trees and found one optimal U'ee of 
50 steps. Two trees of length 5 1 were observed as 
well as one tree of 52 steps. Neither of the trees 
of 51 steps in length are considered here as the 
phyletic relationships presented by each do not 
represent the phascol^ctids as a monophyletic 
clade. In both cases Koobor is intermediate be- 
tween Madakoala spp. and Phascolarctos 
cinereus. 

DISCUSSION 

Classification of Ku. ngama as an ilariid was 
tentative (Pledge, 1987) and controversy sur- 
rounded ])\2icexncniof Koobor, Comparison of the 



388 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Riversleigh ilariid with species of Nana and 
Kutehntja ngama. confirms that the Riversleigh 
animal is indistinguishable from the latter. Dental 
variation in Phascolarctos cinereus, one of Ku. 
n^awc/'s closest living relatives, suggests that: 1) 
variation in Riversleigh fossil material is in the 
range for vombatiform species, and represents 
only one taxon; and 2) the Riversleigh species is 
Ku. ngama. 

DISCUSSION OF THE PHYLOGENETIC 
ANALYSIS. Kiiterintja ngama as the sister taxon 
of a clade containing llaria iUumidens and Haria 
lawsoni (Fig. 6), and not united with wynyardiids 
or Koobor, reinforces classification of this animal 
as an ilariid. Synapomorphies used by 
DELTRAN to unite ilariids include: 1 ) a well-de- 
veloped transverse valley; 2) poorly-developed 
postparacrista and premetaconulecrista; 3) pro- 
tocone longitudinally compressed relative to the 
metaconule on M'-M^^; 4) large crown height; 5) 
moderately well-developed posterolingual cusp 
on P-^; 6) closure of the transverse valley on upper 
molars by a lingual cingulum; 7) a non-bladed 
and bulbous P-^; 8) a non- bladed P3 and 9) a 
'central' cuspid on the protolophid and hypo- 
lophid of lower molars. These synapomorphies 
only apply to the Ilariidae relative to the other 
taxa used in this analysis, and may prove to be 
symplesiomorphies when all other vombato- 
morphian laxa are included. Some of these syn- 
apomorphies refer to the upper dentition, which 
is unknown for Uaha lawsoni. However, the 
close similarities between the lower dentition of 
both llaria species suggests that these syn- 
apomorphies will be generically significant when 
upper dentition for /. lawsoni is found. Syn- 
apomorphies used by the same algorithm to unite 
species of llaria include: 1) moderately well-de- 
veloped transverse linkages between cuspids; 2) 
weak paraconid; 3) large tooth size; and 4) trans- 
versely compressed, caniniform lower first inci- 
sors. 

In constructing the most parsimonious tree, 9 
characters were found to exhibit some degree of 
homoplasy. According to DELTRAN moder- 
ately well-developed transverse linkages be- 
tween cuspids is due to convergence between 
primitive wynyardiids and species of llaria, with 
Kmerintja ngama with plesiomorphic pooriy de- 
veloped linkages. Conversely, ACCTRAN sug- 
gests that moderately well- developed transverse 
linkages were already a feature of the common 
wynyardiid/ilariid ancestor, possibly before 
Koohor diverged from the vombatomorphian lin- 




FIG. 6. Relationships of the Ilariidae and Koohor. 
DELTRAN Synapomorphies: A-weak longitudinal 
valley; B-well-developed stylar cusps; moderately 
developed transverse valley; separation of stylar 
cusps C and D; weak paraconule on M^; no pro- 
tostylid; protoconid>metaconid; no compression of 
lingual portion of tooth; moderate crown height; dor- 
sally flattened and distally inflected li; no longitudi- 
nal valley; C- well-developed transverse valley; 
poorly developed postprotocrista and premeta- 
conulecrista; protocone longitudinally compressed 
relative to metaconule on M'-M^; large crown; mod- 
erately developed posterolingual cusp on P-^; 'centraP 
cuspid; posterolingual cuspid on P3; lingual closure 
of transverse valley on uppers by a cingulum; non- 
bladed P3 and P^; D-moderately well developed trans- 
verse linkages; weak paraconid on Mi; large leelh; 
caniniform and conical 1|. ACCTRAN Syn- 
apomorphies: A-moderately developed transverse 
valley; moderately developed transverse linkages; no 
protostylid; protoconid melaconid; dorsally flattened 
and inflected Ii ; no posterobuccal cuspid on P3; mod- 
erately developed longitudinal valley; B-well-devel- 
oped stylar cusps; separation of stylar cusps C and D; 
no paraconule; uncompressed lingual portion of 
tooth; moderate crown height; moderately developed 
posterolingual cusp on P-^; no longitudinal valley; C- 
well-developed transverse valley; poorly developed 
postprotocrista and premetaconulecrista; protocone 
longitudinally compressed relative to metaconule on 
M' - M^; metaconid a lingually convex crest; high 
crowns; 'central' cuspid; posterolingual cuspid on P3; 
transverse valley closed lingually by cingulum; non- 
bladed P3 and P; bulbous P^ D-poorly developed 
paraconule; weak paraconid on Mj; large teeth; can- 
iniform, conical li; posterobuccal cuspid on P3; and 
Pi 



KUT^iNTJA NGAMA FROM RJVERSLEiGH 



389 



cage. Poorly-dcveioped transverse linkagCvS in 
Ku^ ngama would ihcrcfore be the result of a 
reversal to the phascotarctid stale. The 
ACCTRAN model appears preferable, although 
discovery of a lower dentition for Koobor would 
help resolve its classification. 

Hie poorly-developed paraconulc on M' of //a- 
ria iUuniidens (Tcdford & Woodhurne, 1987). is 
^ther a plesiotnorphy dating fri)in some time 
aflcr divergence of A'oojbor (DELTRAN), or (he 
result of a reversal (ACCTRAN). The former 
hypothesis implies that loss of the paracunule is 
convergent between wynyardiidsand A'm. ngcmw, 
while the latter, and possibly more piu^simonious, 
hypothesis suggests that the paraconule was al- 
ready lost from Ihe vomhatomorphian lineage 
before the wynyardiids and ilariids diverged. 

The paraconid on Mi is another homoplasic 
character, Por both algorithms character Iransfor- 
niation suggests that a well-developed paraconid 
is convergeni between /. illunjidens and 
Mihiokoala. A poorly developed paraconid is 
deemed to be convergent between species onia- 
tio and primitive Madakoala. with absence of a 
paraconid being the plcsiomorphic phascolarciid 
Characiei' stale. However, a more likely solution 
is: 1) that a well-developed paraconid is the 
plesiomorphic condition; 2) that absence of a 
well-developed paraconid in P. ctneretis is a de- 
rived condition; 3) that the paraconid was grad- 
ually reduced or lost before or after Koohftr 
diverged; 4) that the paraconid in species of liana 
represents a reversal to the plesiomorphic ^tate; 
and 5) that loss of a paraconid is convergeni 
between F. cinerem, wynyardiids and possibly 
Koobor. Knowing whether there was or was nol 
u paraconid in Koobor would help clarify ihis 
situation. 

Both algorithms suggest thai a proloslylid on 
Ml oi llaria iHumidens represents a reversal to 
the plesiomorphic phascolarciid condition. The 
only discrepancy between the (wo chantcier 
iran.sformati<ui paihwjiys is the point ai which ihc 
prolostylid was losi. DELTRAN dela)s loss of 
ihc prolostylid until after the divergence of 
Koohof, while ACCTRAN mainiajns thai loss 
occiiried l>efore the divergence. An identical 
character transformation occurs for 'relative 
heights of the anlenor cuspids' (character I I), 
such that cuspids which arc suhequal in height 
represent a reversal to the plesicnnorphic condi- 
tion for /. illumidens. Possessing a prolocomd 
larger than the mciaconid is therefore a syn- 
fflpamorphy uniling wynyurdnds. / Uiwaottii 
Kiaviinrja ngama and possibly Krft*liftr. A 



poslcrobuccal cuspid on Pi of species of lUiria li 
deemed to be a reversal la Ihc plesiomorphic 
phascolaattid condition by ACCTRAN. while 
DELTRAN suggests thai abscivce of this slnic 
ture is convergent bet\s'cen wynyardiids and 
Kuierintja ngatna. Again. ACCTRAN seems li> 
be the most parsimonious, implying thaJ Ihc 
posierobuccal cuspid was lost before 
wynyardiids and ilariids. and possibly KooboK 
diverged. 

For lingual closure of the transverse valley on 
upper niularb fcharaciet 20) the pathway for char- 
acter transformation is unclear due primarily to 
the variable narutx; of this ^\xy^cxwr\i\wMadakoala. 
However, the suggested inmsrorniaiion sequence 
is: 1) a partial cinguluin. in the form of 2 cuspules 
on the iintea^lingual and posierolingual bases of 
the mclaconule and protocone respectively, was 
present in the ancestral koala; 2 ) the two cuspulcs 
eventually joined, convergenlly forming ihc de- 
rived lingual cingulum in f nnereus, some spec- 
imens nf Madakoala and ilariids; and 3) other 
Madakoaia and wynyardiids developed in the 
opposile direcii*m, convergenlly losing the 
cuspulcs altogether- The two cuspulcs occur in 
Koobor noudndis but not in Koobor jimbarotii 
(Archer. 1977). perhaps suggesting thai ihc latter 
is more derived than the former Aliernaiively, a 
lingual cingulum may be a plesiomorphic 
phascotarctid condition, implying thai lh<; 
cuspules in Ko. m^uihidis ane an apomorphic ves- 
tige. In this case, absence of a lingual cingulum 
would be a mure derived condiiion. convergent 
between some specimens of Madakoala^ Kit, 
jimbaraui and wynyardiids. 

The final character trarxsrormaiion found to 
contain some dega-e ol honxtplasy involved a 
cuspon (he poMor^^buccal margin of P^ (character 
23). The sequence of change suggested by the 
algorithms is that absence of such a cusp is ihc 
plcsiomoqihic condition, and th;it species of 
Madokoola, llona Hiiimiilens and possibly /. 
tow'ionj convcrecntly share a derneil posieir>huc- 
C3l cusp. This iransfornialjon ^rqucnce s^-enis 
unlikely because Ma<kikoala arc overall nKW 
plesiomorphic phascolaiciids (Woodbumc el at.. 
1987) and a poMerohuccal cusp on P'' is more 
likely lo be ihc ples^ionvorphic condition. If so. 
los> of this structure is a Avnapomorphy for 
Koobor. wynyardiids ;snd Kit ft^ama. and is con- 
vergent on the condiiiun in Phitsccftarctoi^ 
cinereus. Absence of the ctt&p ifl this context i% 
yet aninher potential chanicler state ^pai^atiirg 
Kf*f^b4*r rrv>n> pKascoIairtid.'L 

According to Simmons (1*^93) iJk cji|»?<?tcj 



390 



MEMOIRS OF THE QUEENSLAND MUSEUM 



value for the consistency index of a tree with 7 
taxa is: 

CI = 0.90 - 0.022 (7) + 0.000213 (7)^ = 0.736 
(3 s.O. 

The observed CI for the optimal intrafamilial 
tree is 0.800, 0.064 from the expected value. The 
observed CI value implies slightly less homo- 
plasy for the intrafamilial analysis than would be 
expected for seven taxa. Similarly the retention 
index (RI) and the reseated consistency index 
(RC) are reasonably large, emphasising the low 
degree of homoplasy and potential for homoplasy 
respectively. 

The optimal tree which includes all outgroup 
taxa is reasonably well-supported by bootstrap 
analysis and by the lack of significantly different 
trees, of plausible topology, within a few steps of 
the most parsimonious. The low bootstrap result 
for the Ilaria clade is almost totally due to the 
amount of missing data for /. kiwsoni. This study 
supports the notion that Ku. ngama is an ilariid 
and forms a monophylctic clade with Ilaria. 

CLASSIFICATION OF KOOBOR 

Pledge (1987) discussed the possibility that 
Kuterintja ngama is more closely related to 
Koobor than Ilaria. The lower dentition and 
upper molars in addition to M"* demonstrates that 
Ku. ngama is an ilariid. One of the few similari- 
ties between Koobor and Ku ngama is the smooth 
rounding of the hngual faces of the Hngual cusps, 
a character slate previously thought to unite the 
taxa phylelically (Pledge, 1987), However, 
smooth and rounded lingual faces on lingual 
cusps are also a feature of wynyardiids, and to a 
lesser extent Madakoala, suggesting that it is 
plesiomorphic. The ambiguity of this character 
state also increases the possibility of homoplasy. 
Pledge (1987) hypothesised that Ku. ngama may 
be ancestral to Koobor. Our study does not sup- 
port this view. 

Koobor notabalis appears to be the primitive 
sister-group of wynyardiids plus ilariids. We 
havenoclearsupporl, however, for /£'(9(9/7or being 
in the Phascolarctidae. This may be indirect sup- 
port for the suggestion that Koobor represents a 
distinct family of vombatiform marsupials. 
DELTRAN found only one synapomorphy po- 
tentially uniting Koobor with the wynyardiids 
and ilariids: the less well-developed longitudinal 
valley on the molars. ACCTRAN found 7 syn- 
apomorphies Ibr a A'(9o/7c>r,wynyardiid and ilariid 
clade. This should not be taken at face value, 
however, as 6 of these character slates refer to the 
lower dentition which is unknown (or Koobor. 10 



synapomorphies were found by DELTRAN to 
unite wynyardiids and ilariids to the exclusion of 
Koobor (Fig. 6). 

BIOCORRELATION OF RIVERSLEIGH 
AND THE ETADUNNA FORMATION 

Ku. ngama occurs in the White Hunter Local 
Fauna at Riversleigh and in the Ngama Local 
Fauna in the upper Etadunna Formation. Ilaria 
lawsoni occurs in the Ditjimanka Local Fauna in 
the lower Etadunna Formation. Ilaria illumidens 
occurs in the Pinpa Local Fauna of the Namba 
Formation, at Lake Pinpa. 

Woodbume et al. (1993) suggested at least 6 
magnetic reversals within the Etadunna se- 
quences, correlated them with a biostratigraphic 
zonation and the MPTS (Fig. 7) and suggested 
24-28 Ma for the base of the Etadunna Formation. 

Woodburne et al. (1993) correlated Zone D 
with the Ngama and Tarkarooloo Local Faunas. 
Correlation of magnetic polarity and 
biostratigraphic zones places zone D in lower 
magnetozone R3, which in turn correlates with 
Chron 7n. 1 r of the MPTS, or 24.7 - 25.0 Ma. Ku, 
ngama therefore, correlates White Hunter Site 
with the Ngama Local Fauna at 24.7 - 25.0 Ma 
providing: 1 ) that ilariid material in White Hunter 
Site has not been reworked from older deposits 
(which, given the lack of evidence for weathering 
or transport, does not appear likely); and 2) that 
the apparently short temporal range o\Ku. ngama 
in the Etadunna Formation is the full range of this 
species. 

ACKNOWLEDGEMENTS 

Vital support for research at Riversleigh has 
come from the Australian Research Grant 
Scheme, the National Estate Grants Scheme 
(Queensland), the University of New South 
Wales, the Commonwealth Department of Envi- 
ronment, Sports and Territories, the Queensland 
National Parks and Wildlife Service, the Com- 
monwealth World Heritage Unit, ICI Australia, 
the Australian Geographic Society, the Queens- 
land Museum, the Australian Museum, the Royal 
Zoological Society of New South Wales, the 
Linnean Society of New South Wales, Century 
Zinc, Mount Isa Mines, Surrey Beatly & Sons, the 
Riversleigh Society, and private supporters in- 
cluding Elaine Clark, Margaret Beavis, Martin 
Dickson, Sue & Jim Lavarack and Sue & Don 
Scott-Orr. Vital assistance in the Held has come 



KUTERINTJA NGAMA FROM RIVERSLEIGH 



39: 



MPTS MAGNETOZONE 



Etadunna Etadunna Namba Riversleigh 
Mammal Formation Formation Assemblage 
Zone Assemblage Assemblage 



24.2 Ma 



6Cr 



24.7 Ma 



7r 



25.5 Ma 



Tan 



R4 



Vn.ln 


N3 


1 




7n.lr 


R3 


7n.2n 


N2 



R2 



Nl 



7ar Rl 

25.7 Ma ? 



D 

7 

C+B 

A 

7 



"Treasure/ 
Lungfish" 












Ngania 


Tarkarooloo 


White 
Hunter Site 








Ngapakaldi 
Ditjimanka 


Enemas 




Wynyardiid" 


Pinpa 





FIG. 7. Geochronology and biocorrelalion of the Etadunna and Namba Formations, Lake Palankarinna and Lake 
Pinpa, S.A. and lower Riversleigh faunas, northwestern Queensland. (Modified from Woodbume et al., 1993, 
figs 2 and 15). N = Normal magnetic polarity; R = Reversed magnetic polarity; MPTS = magnetic polarity time 

scale. 



from many hundreds of volunteers as well as staff 
and postgraduate students of the University of 
New South Wales. Skilled preparation of most of 
the Riversleigh material has been carried out by 
Anna Gillespie. TJM acknowledges the assis- 
tance of Prof. Alberto Albani, Karen Black, Jenni 
Brammall, Henk Godthelp, Steve Salisbury, 
Anne Musser, Mary Knowles and his family. 

LITERATURE CITED 

APLIN, K.P. & ARCHER, M. 1987. Recent advances 
in marsupial systematics with a new syncretic 
classification, p.xv-lxxii. In Archer, M. (ed.), Pos- 
sums and Opossums: studies in evolution. (Surrey 
Beatty & Sons and the Royal Zoological Society 
of New South Wales, Sydney). 

ARCHER, M. 1977. Koobor notabalis (De Vis), an 
unusual koala from the Pliocene Chinchilla Sand. 
Memoirs of the Queensland Museum 18; 31-35. 



ARCHER, M. 1984. The Australian marsupial radia- 
tion. Pp. 633-808. In Archer, M. & Clayton, G. 
(eds), Vertebrate zoogeography and evolution in 
Australasia. (Hesperian Press: Perth). 

ARCHER, M., GODTHELP, H., HAND, S.J. & 
MEGIRIAN, D. 1989. Fossil mammals of 
Riversleigh, northwestern Queensland: prelimi- 
nary overview of biostratigraphy, correlation and 
environmental change. Australian Zoologist 25: 
29-65. 

ARCHER, M., HAND, S.J. & GODTHELP, H. 1994. 
Riversleigh. 2nd ed. (Reed: Sydney). 

CREASER, P. 1997. Oligocene-Miocene sediments of 
Riversleigh: the potential significance of topogra- 
phy. Memoirs of the Queensland Museum 41(2): 
303-314. 

FLOWER, W.H. 1867. On the development and suc- 
cession of teeth in the Marsupialia. Philosophical 
Transactions of the Royal Society, London 157: 
631-641. 

LUCKETT, W.P. 1993. An ontogenetic assessment of 
dental homologies in therian mammals. Pp 1 82- 



392 



MEMOIRS OF THE QUEENSLAND MUSEUM 



204. In Szalay, F.S., Novacek, M.J. & McKenna, 
M.C- (eds), Mammal phylogeny: Mcsozoic differ- 
entiation, multituberculaies, monotremes, early 
therians and marsupials. (Springer-Verlag: New 
York), 

MARSHALL, L.G., CASE, J.A. & WOODBURNE, 
M.0. 1990. Phylogencticrelalionships of the fam- 
ilies of miirsupials. Pp. 433-505. In Genoways, 
H.H. (ed.). Current mammalogy, vol. 2. (Plenum 
Press: New York). 

PLEDGE, N.S. 1987. A new genus and species of 
primitive vombatoid marsupial from the medial 
Miocene Nsama Local Fauna of South Au.stralia. 
Pp.419-422'1 In Archer, M. (ed.). Possums and 
Opossums; studies in evolution. (Surrey Beatly & 
Sons and the Royal Zoological Society of New 
South Wales: Sydney). 

SIMMONS, N.B. 1993. 11)e importance of methods: 
archonlan phylogeny and cladistic analysis of 
morphological data. Pp. 1-51. In MacPhee, D.E. 
(ed.). Primates and Iheir relatives in phylogenetic 
perspective. (Plenum Press: New York). 

TEDFORD, R.H. & WOODBURNE, M.O. 1987. The 
llariidae, a new family of vombatiform marsupials 
from Miocene strata of South Australia and an 
evaluation of the homology of molar cusps in the 



Diprotodontia. Pp. 401-418. In Archer. M. (ed.). 
Possums and Opossums: studies in evolution. 
(Surrey Bealiy & Sons and the Royal Zoological 
Society of New South Wales: Sydney). 

WrLEY. E.O., BROOKS, D.R., SIEGEL-CAMSEY, 
D. & FRANKS. V. A. 1991. Thecompleatcladisi: 
a primer of phylogenetic procedures. University 
of Kansas Museum of Natural History Special 
Publication 19. 

WOODBURNE. M.O., TEDFORD, R.H.. ARCHER, 
M. & PLEDGE, N.S. 1987. Madakoala, a new 
genus and two species of Miocene koalas 
(Marsupialia; Phascolarciidae) from South Aus- 
tralia, and a new species of Ferikoala, Pp. 293- 
3 1 7. In Archer, M. (ed.). Possums and Opossums: 
studies in evolution. (Surrey Beatty & Sons and 
the Royal Zoological Society of New South 
Wales: Sydney). 

WOODBURNE, M.O.. MACFADDEN, B.J.. CASE, 
J. A., SPRINGER. M.S., PLEDGE, N.S., 
POWER, J.D., WOODBURNE, J.M. & 
SPRINGER, K.B. 1993. Land mammal 
biosiratigraphy and magnetostratigraphy of the 
Etadunna Formation (late Oligocene) of South 
Australia. Journal of Vertebrate Paleontology 13: 
483-515. 



NANOWANA GEN. NOV. , SMALL MADTSOIID SNAKES 

FROM THE MIOCENE OF RIVERSLEIGH: S YMPATRIC SPECIES 

WITH DIVERGENTLY SPECIALISED DENTITION 

JOfrN D SCANLON 



ScatiluTi. J.D. 1997 06 30: Nanowana gen. nov.. small madtsoiid snakeS from Ihc Miocene 
of Rivcrslcigh: svmpainc species wuh divergemlv specialised denlilioTi. Memoin of the 
Qiwvnslantl Museum M{IY 393-412. Brisbane. ISSN 0079-SS35- 

Twu sinal 1 early Miocene madtsoiid snalvcs from Riversleigh, N W Queensland are described 
as Nanowana goiirhelpi gen. et sp. nov. and N. uhrenki gen. et sp. nov. Jaw elements t)f the 
former arc depressed, lack ankyloscd icelh, and have alveoli of nearly uniform size, Ihcsc 
features arc interpreted as signs of a coadaptcd character complex ( "anhrodonty ' ) where (he 
leeih are attached (o the jaws by a fibrous hinge. This condition is associated with a diet of 
hard-scaled scincid li/ards The latter species retains ankylosis, and has strongly enlarged 
teeth on the anterior demary and middle maxilla indicating a distinct method of subduing 
prey, but extant analogues are also predominantly scincivorous. Departure in ejch species 
from the nearly hoinodonl. iinkylosed condition in other madlsoiids is interpreted as adapta- 
tion loadtcl of scincid lizards. These divergent, but lunctionally panllel specialisations are 
likely to be independently derived from the ancestral condition 

John D. Scanhn. School oj Biological Sciences, University of Nrw Soi4lh Wales, New South 
Wales 2052, Australia (email: johns@ gekonet, an); recet^'t^i 7 Febtvan 1997, 



Madtsoiid snakes in Tertiary faunal assemblages 
ofRiversleigh(Scanlon 1992, 1993, 1995, 1996) 
have been lefen'ed to Yurlim^^iur Scanlon, 1992 
and Wonambi Smith, 1976. Other Rivcrslcigh 
rnadisoiids cannot be included in previously 
known genera. Two small species, estimated to 
reach Im It^ng, are represented by upper and 
lowerjaw elements from System B (Archer eial., 
1989, 1994) o{\ Godthelp Hill. Some are associ- 
ated with vertebrae, but the two species cannot be 
distinguished unambiguously on vertebra! char- 
acters. I include them in a single genus which 
possibly unnatural treatment allows generic iden- 
liiication of isolated vertebrae from other sites. 

This paper provides descriptions of the two 
species including some ontogenetic stages. WTiile 
analysis of phylugeny o\' madlsoiids awaits de- 
tailed comparisons with other primitive snakes, 
some functional and evuluiionaiy points are 
noted by analogy with extant forms. 

MATERIALS AND METHODS 

Material is housed in the Queensland Museum 
(QMF). Australian Museum (AMK), Northern 
Territory Museum of Arts and Sciences (NTMP), 
Museum olVicloria (NMVP), and South Austra- 
lian Muscuit) (SAMP). ...(SMNR> specimens ck- 
ammed in Paris by courtesy of J.-C. Rage. 

Teeth or ;dveoli are numbered beginning from 
the anterior on complete jaw clcmenis; on frag- 
rnenls where ilie \ooih row is or may be inconv 



pletc anterioriy the numbers are spelled out in 
words. In illustrating cranial bones, views of the 
same specimen are usually arranged parallel to 
each other, in lateral, dorsal, medial, ventral as- 
pects. Figures of vertebrae have left lateral, ante- 
rior, posterior, dorsal, and ventral views of each 
clement in a vertical row. If more than one verte- 
bra arc shown in an illusiralion. they are arrangeil 
(I to r) in order Irom anlenor to posterior. 

5YSTEMAT1CS 
Family MADTSOriDAE Hoffstetter, 1961 
Nanowana gen. nov. 
TYPE SPECIES. Nanowana godthefpi sp. nov. 
OTHER SPECIES. Nanowana schrenki sp. nov. 

ETY.VIOLOGY. Greek nonos. a dwarf and Warlbiri 
(Tanami Desert, central NT) Wana. Rainbow Serpent 

of Aboriginal mythology, 

DIAGNOSIS. Small, upto 1.5m long; neural 
spine low to moderately high, not extending close 
10 anterior edge of zygosphene; /ygosphene shal- 
low, with anierodorsal edge straight, slightly con- 
vex or concave in dorsal view; subccniral ridges 
well-defined, straight or slightly concave or con- 
vex in ventral view; haemal keel relalively nar- 
row, wuh 'paired liyt>apophyscs' In (loslerior 
trunk dclined laterally, hut not projecting ven- 



394 



MEMOIRS OF THE QUEENSLAND MUSEUM 




dorsal process with steep an- 
terior edge; dentary with at 
least 2 mental foramina. 

COMPARISON. This genus 
is distinguished from all 
madtsoiids other than .4/- 
amitoph is by the z y g o- 
sphene in dorsal view 
frequently (but not always) 
having a convex anterior 
margin; the convexity is 
broad rather than a distinct 
median tubercle as in Al- 
amitophis. It is distinguis- 
hed fro m Yurlimggur, 
Wonambi, Rionegrophis, 
Gigantophis and Madtsoia 
by being smaller. Its neural 
spines are lower, at corre- 
sponding positions in the 
trunk, than in Madisoia^ 
Rionegrophis, Wonambi and 
Alamiiophis, but higher than 
Fatagoniophis or Giganto- 
phis. It is distinguished from 
all genera except Fatagoni- 
ophis by the less steeply 
converging subcentral 
ridges (relatively more elon- 
gate centrum in ventral 
view). Maxillae resemble 
Madtsoia sp. (SMNR 2879, 
Itaborai) and are distin- 
guished from Wonambi and 
Yurhmggur by prefrontal 
process having a steep ante- 
rior edge; distinguished 
from each of these by devel- 
opment of the sepiomaxill- 
ary process (condition 
unknown in other 
madtsoiids). 



DISCUSSION. Vertebrae 

can be distinguished from 
FIG. I. A'flnoM'cfnfl gi^*f//7^//>isp. nov.. QMF3 1379, hololype. upper jaw bones other madtsoiids, but their 
(right (A) and lefi(B) maxillae, palatines (C.D) and pterygoids (E.F) of a single common feaVures'(including 



individual) in palatal view, CS Site. Scale bar=5mm. 

trally or separated by a median concavity; neural 
arch in pfjsterior trunk depressed, its lateral por- 
tions strongly concave dorsad. Anterior tip of 
maxilla with medial expansion (septomaxillary 
process) contributing to floor of narial chamber; 



small size) may be sym- 
plesiomorphic; the concept 
of Nanowana containing these 2 species can be 
described as a ^marriage of convenience'. The 
phylogenetic relationships of these with other 
madtsoiids remain unknown, but they arc treated 
as a unit because their vertebrae (which provide 



NEW MIOCENE MADTSOIID SNAKE FROM RIVERSLEIGH 



m 



the only taxonomically use- 
ful material in most depos- 
its) are unable to be 
distinguished in many cases. 

In a number of aspects of 
the vertebrae, including size, 
Nanowana is comparable to 
Patagoniophis sp. cf. P. 
pan'us from the early 
Eocene Tingamarra Local 
Fauna (Scanlon 1993); dif- 
ferences include Ihe higher 
neural spine (m adults), nar- 
rower haemal keel in the 
posterior trunk, frequently 
convex anterior edge of the 
zygosphenc, and dorsolat- 
eral concavities of the poste- 
rior neural arch. It differs 
from Alamitophis, which 
also occurs in the Australian 
Eocene (Scanlon, 1993): the 
anterior edge of the 
zygosphene, when convex, 
is broadly so rather than 
forming a distinct promi- 
nence; paradiapophyses do 
not project anieriad; zyg- 
apophyses are more steeply 
inclined at equivalent posi- 
tions within the column. The 
lower neural spine, broader 
zygosphene. and features of 
the haemal keel or 
hypapophyses differentiate 
Nanowano from Wonambi 
Smith, (1976) {Wonambi is 
known from Rivcrsleigh, 
much smaller than W. nar- 
acoorrensis but larger than 
Nanowano; Scanlon, 1996). 

The only other known 
Australian madtsoiid is 
Yurlimggur, at least 2 spe- 
cies of which occur al 
Riversleigh as well as the 
type species from Bulhx'k 
Creek (middle Miocene; 
Scanlon 1992). That genus 
exceeded 5m and thus in- 
cluded only 'giant' snakes, though not as large as 
Gigantophis garstini or Madtsoia bai. However, 
size is rather variable in many snake genera (e.g. 
the pythonid Morelia, sensu Underwood & Stim- 
son, 1990, includes species with maximum 







FIG. 2. Nanowana godthelpi sp. nov. QMF31379, holotype. upper jaw bones 
(left maxilla (A-C), left palatine (D-F) and right pterygoid (G-1)) in lateral, 
dorsal and medial views, CS Site. Scale=5mm. 



lengths from under 1 m to over 7m), and need not 
be considered an essential part of the diagnosis. 
The vertebrae of small and large forms arc rather 
similar except in features which may be directly 
related to size (neural spine height is variable 



396 



MEMOIRS OF THE QUEENSLAND MUSEUM 



within Yurhmggur, and is proportionally similar 
to Nanowana in some), but Nanowana differs 
from Yurlunggur in the shape of the /ygosphene, 
and the haemal keel of posterior trunk vertebrae 
being narrower and lacking a median concavity. 
Comparisons with non-Ausiralian fonns do not 
suggest any links closer than thai with Yurlun- 
ggur, and will not be pursued here. The rib-heads 
o{ Nanowana have not been considered in detail, 
but appear to be similar in shape to those of 
Yurlunggur v^nd W'onnm^/ (Scanlon, 1993). 

Nanowana godtheipi sp. nov. 
(Figs 1-8, Table 1) 

ETYMOLOGY. For Henk Godthelp, University of 
New South Wales, in recognition of his contributions 
to Australian palaeontology. 

MATERIAL. Hololype QMF31379, associated ele- 
ments of a single individual compnsing partial to com- 
plete maxillae, palatines and pterygoids of both sides. 
Paraiypes QMF 3 1 383, 3 1 384 associated dentaries and 
compounds of a single individual; dentaries 
QMF20892, 23052, 23053, 23054, 23056); maxillae 
QMF31380, 31382, 31386, 31387; palatine 
QMF3 1381; pterygoids QMF23058, 31393. All types 
from early Miocene (System B) Camel Sputum Site, 
Godthelp Hill. Other material: Camel Sputum Site, 
trunk vertebrae QMF19741. Upper Site, deniary 
QMF31389; palatine QMF23066; maxilla fragment 
QMF31390; pterygoids QMF23067. 31385; series o^ 
cloacal vertebrae. Mike's Menagerie Site, anterior 
fragmentofpterygoidQMFI 9742. Greaser's Ramparts 
Site, dentary QMF23076. 

DIAGNOSIS. Palatine lateral process about as 
long as two alveoli (nearest to 4th and 5th), ven- 
tral concavity of process with obtuse angle ac- 
commodating posterolateral angle of palatine 
process of maxilla. Maxilla with 23 tooth posi- 
tions, palatine 1 1 , pterygoid 9, dentary 1 6. Teeth 
not ankylosed to alveoli; maxillary alveoli vary 
only slightly in size, dentary alveoli largest in 
centre of tooth row (4-8 or 5-8). Posterior part of 
maxilla strongly depressed. Dentary tooth row 
curved in dorsal view. Two or 3 menial foramina, 
all anterior to the 7ih alveolus. 

DESCRIPTION OF HOLOTYPE. Upper dentig- 
erous elements in a single block (without verte- 
brae or other elements) are complete on one or 
both sides, missing bilaterally only the posterior 
(quadrate) processes of the pterygoids (Figs 1 ,2). 
Maxillae long and fiat posteriorly, supporting a 
high lizard-like prefrontal process anteriorly; pal- 
atines with *alethinophidian' features; pterygoids 



with prominent, also lizard-like, ectopterygoid 
processes. Proportions of jaws indicating a rela- 
tively long postorbilal skull and moderately short, 
rounded snout. 

Palatine: Left more complete than right, both 
well-preserved. Eleven alveoli forming a sigmoid 
tooth row, convex laterad anterior to an inflection 
and lateral concavity (slight, but definite and 
angular) between 7th and 8th. Dorsolateral crest 
arising above 3rd alveolus, bifurcating above 4th 
to form anterior edges of maxillary and choanal 
processes. Maxillary process with an oblique an- 
terior edge (near 45° from sagittal plane), longi- 
tudinal lateral edge and transverse posterior crest 
on its ventral face, level with the 5th alveolus on 
the left palatine (4th-5lh on right side); process 
not perforated or notched for the maxillary nerve. 
Anterior edge of the choanal process smoothly 
concave anteriad for its full width, reaching be- 
tween level of 4th and 5th alveoli; then curving 
strongly anteroventrally, extending to front of 
2nd alveolus. Vertical anteromedial part of the 
choanal process bilobed anteriorly, a dorsal lobe 
curved mediad, the other laterad (forming articu- 
lations with the parasphenoid and vomer); third, 
posterolaterally pointed, lobe on the ventral edge 
deflected laterad, contributing (along with the 
vomer and ectochoanal cartilage, presumably) to 
the floor of the choanal passage. Lamina of choa- 
nal process strongly arched anteriorly, flatter pos- 
terioriy. and ventrally deflected part of lamina 
reducing in depth posterioriy. Posteromedial cor- 
ner of process level with rear of 9th alveolus, 
posterior margin sinuous so that posterior process 
not sharply demarcated (as in some specimens); 
margin concave medially, convex posterioriy. 
Posterior extremities of choanal process and 
tooth row extending back level with each other, 
both with lateral margin parallel to tooth row, and 
separated by a distinct triangular notch extending 
forward to middle of 1 Ith alveolus (thus, poste- 
rior edge W-shaped); on dorsal face this notch 
continued as a tapering trough extending to rear 
of 9th; ventrally a step-like groove running from 
the apex of the notch anteromediad to between 
9th and lOth, with a shallow trough posterior and 
partly medial to the groove. Small foramen 
dorsomedially on the dcntigerous process, just 
below the ridge continuous with the anterior edge 
of the choanal plate; a large foramen medial to 
the 8lh alveolus, piercing the plate and emerging 
dorsally as a posteriorly widening foramen be- 
tween 8th and 9th; another small foramen an- 
teromedial to 1 0th alveolus. Dorsomedially on 
the anterior dentigerous process with lip of a tooth 



NEW MIOCENE MADTSOIID SNAKE FROM RIVERSLEICH 



197 



emerging from the bone 
(ihis is ihe only tooth associ- 
ated wilh jaws of this spe- 
cies). 

Right and left palatines al- 
most identical; spacing of al- 
veoli slightly different on 
different sides; alveoli 2-5 in 
the right shifted posteriorly, 
relative to the left (alveoli 1 
and 2 on the left, 5 and 6 on 
the righl. confluent). Lateral 
(maxillary) process with 
small but distinct angular 
concavity marking the lon- 
gitudinal (lateral) and 
oblique (anterolateral) sec- 
tions of the margin. 

Pterygoid. Nine alveoli 
(complete row), anterior tip 
(length of approximately 1.5 
alveoli) edentulous. Tooth 
row curving medially poste- 
riorly, following inner edge 
of bone; ventral face nar- 
rowing to a point anterior to 
tooth row, point interlocking 
with posterior notch of pala- 
tine. Dorsal surface forming 
a longitudinal trough, with 
foramen above 1st alveolus 
(opening anleriad), lateral to 
a dorsomedial ridge. Lateral 
margin smoothly convex, 
diverging gradually from 
tooth row; anterior edge of 
eciopterygoid process di- 
verging at about 120"' from 
this margin, level wilh 7th 
alveolus. Process nearly as 
wide as rest of bone at this 
point, about as long as wide; 
its anterior and lateral edges 
at 90^^ in dorsal or ventral 
view, lateral margin inclined 
strongly posteroventrally, 
with posterior extremity 
produced as a knob-like ex- 
tension, and posterior edge 
strongly concave. No part of 
the ectopterygoid facet ex- 
posed dorsally. Concave 
posterior surface of Ihe pro- 
cess continuous wilh the ventrolateral face of the 
posterior lamina (quadrate process), bounded 





);?i5BXitt5£- 







FIG. 3. Nanowana godthelpi sp. nov. QMF3 1383. 3 1384, paraiype lower jaws. 
A, B. left dentary in lateral and medial view (upper posterior process broken 
and slightly displaced). C, D. E, righl denlary in medial, dorsal, and lateral 
view. Scale=5mm. 



medially by a narrow extension of the ventral 
(occlusal) surface. Quadrate process broken dIT 



398 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 1 . Measurements (mm) of jaws of Namowana godthelpi sp. nov. C 1 , C2, etc.=single individuals; L=lefl, 
R=right. Alveoli were selected as landmarks for some measurements because they could be identified in 
fragments, but there is variation in the position of anterior alveoli (even between sides of an individual). Values 
in brackets are minima for measurements affected by damage. 

Palatine (ventral view): ptl=length of palatine from anterior tip of dentigerous process to posterior tip of tooth 
row spine or choanal process; pcl=base length of choanal process from intersection of anterior edge with 
dentigerous process to apex of posteriornotch; pi I l=length from anterior tip to anterior edge of 1 1th alveolus; 
ptw=width across choanal and maxillary processes; pcw=width in same line of choanal process; prw=width in 
same line of tooth row bar; pmw=width in same line of maxillary process. 

Pterygoid (ventral view): ttl=length from anterior spine (in plane of alveoli, not dorsal lappets) to rear of 9th 
alveolus; trl=tooth row 1 st-9th alveolus; tte=from anterior spine to furthest point of ectopterygoid process; ll5= 
length across most posterior 5 alveoli (5-9); taw=width between near-parallel edges anterior to ectopterygoid 
process; tpw=width from basipterygoid facet to intersection of ectopterygoid process and dorsolateral edge of 
posterior lamina; ttw=width from basipterygoid facet to furthest point of ectopterygoid process. 

Maxilla: mtl=length; map=length from anterior tip to posteromedial angle of palatine process; m 712=lenglh 
from anterior edgeS of 7th- 1 3th alveolus; mpw=width across palatine process; mph=depth at prefrontal process. 

Dentary: mff=number of mental foramina; dtl=straight-line length; dl 1 5=length to anterior tip of 1 5th alveolus; 
dlf=length to lateral fossa; d4t= posterior edge of 4th alveolus to posterior extremity; d4 15=posterior edge of 
4th to anterior edge of 1 5th; d4f= posterior edge of 4th to lateral fossa; dl 7=anterior tip to anterior edge of 7th 
alveolus; dmd=depth from dorsolateral to ventromedial edge in middle part of bone; dpp=depth of upper 



posterior process. 






















QMF 


31379 


31381 


23058 


31386 


31380 


31382 


31393 


23066 


23067 


31385 


Ind. 


CI R 


CI L 


C2L 


C2L 


C2R 


C3R 


C3L 


C4R 


UlL 


Ul R 


UIL 


Pti 


(7.6) 


7.7 


(8.0) 


- 


- 


- 


- 


- 


- 


- 




pel 


4.4 


4.2 


4.0 


- 


- 


- 


- 


- 


45 


- 


" 


pill 


6.6 


6.7 


6.8 


- 


- 


- 


- 


- 


- 


- 




ptw 


3.9 


(3.8) 


40 


- 


- 


- 


- 


- 


3.8 


- 


- 


pew 


1.9 


1.9 


2.0 


- 


- 


- 


- 


- 


2.0 


- 


- 


prw 


0.9 


0.9 


1.1 


- 


- 


- 


- 


- 


1.0 


- 


- 


pmw 


1.1 


(I.O) 


(0.9) 


, 


- 


- 


- 


- 


(0.8) 


- 




ttl 


5.7 


- 


- 


- 


- 


- 


- 


- 


- 


- 


- 


tri 


4.9 


- 


- 


- 


- 




- 






- 


- 


tte 


6.8 


_ 


- 


- 


- 


_ 


- 


- 


- 


- 


- 


tl5 


2.6 


2.5 


_ 


3.2 


_ 


- 


_ 


1.6 


- 


3.1 


3.0 


taw 


1.7 


1.6 




1.7 


- 


- 


- 


(1.0) 


- 


1.6 


1.6 


tpw 


2.4 


2.4 


- 


3.1 




- 


- 


- 


- 


2.9 


3.0 


ttw 


4.2 


4.1 


- 


(3.9) 


- 


- 


- 


- 


- 


4.3 


(3.9) 


mtl 


- 


(16.3) 


_ 


. 


- 




- 


- 


- 


- 


- 


map 


- 


7.4 


- 


- 


- 


- 


- 


- 


- 


- 


- 


m7i2 


- 


5.4 


- 


_ 


5.7 


4.6 


5.2 


- 


- 


- 


- 


mpw 




2.3 


- 


- 


(2.8) 


2.8 


2.8 


- 


- 


- 


- 


mph 


- 


(3.3) 


- 


- 


3.9 


3.6 


3.3 


- 


- 


- 


- 










Dentaries 










QMF 


31383 


31384 


20892 


23052 


23053 


23054 


23056 


31389 


23076 


Ind. 


CIR 


CIL 


C2L 


C2R 


C3L 


C3R 


C4R 


UlL 


CRL 


mff 


3 


2 


2 


3 


2 


- 


2 


3 


- 


dtl 


- 


- 


- 


15.1 


- 




- 


- 


_ 




dli5 


15.6 




14.1 


13.3 


- 




- 


- 


- 


dlf 


12.3 


- 


11.4 


11.2 


- 


- 


- 


12.4 


- 


d4t 


- 


13.6 


- 


11.9 


- 


. 


. 


- 


16.3 


d4i5 


11.3 


12.3 


11.1 


10.2 


. 


_ 


- 


- 


14.2 


d4f 


8.0 


9.5 


8.2 


8.2 


- 


7.7 


4.9 


9.6 


11.0 


dl7 


7.8 


- 


6.5 


6.3 


8.0 


- 


- 


- 


- 


dmd 


3.2 


3.0 


2.7 


2.6 


3.3 


(2.4) 


1.6 


2.9 


(2.6) 


dpp 


1.0 


0.9 


0.9 


0.9 


- 


- 


- 


- 


1.4 



NEW MIOCENE MaDTSOIED SNAKE FROM RIVERSLJBIGH 



JW 



posieriorly uboui ha! r the length of (he tooth row 
behind the cclopierygoid process. Basi pterygoid 
ftrticularsurlncc opposite cctoptcrygoid process, 
an oval facei fiicing dorsully and slightly medi- 
ally, beginning level with from of 8ih alveolus 
and extending to beyond 9th, only slightly dis- 
tinct in outline? fann the rest of the medial edge. 
Apart from the anterior foramen mentioned 
above, 3 foramina dorsally, antciior, lateral and 
posterior to the facet; anterior 2 near the midline 
of the hone, posterior foramen close to the medial 
ctlge. A shallow but distinct transverse gnx)veofl 
the dorsal surface of the ectopterygoid process. 

I.eft pterygoid retaining posterior H alvci>li. 
which are slightly smaller and moie closely 
Spaced than on the right: pi»ssibly a lOlh alveolus 
or longer edentulous gap anteriorly. 

Maxilla. Alveoli 23, varying only slightly in 
si/e, row curved medially anienof ly. straight pos- 
teriorly. Anterior alveoli elongate anterolaieral- 
po-steromedially; anterior of raaxdla wider than 
deep, with dorsomedial edge forming a crest 
alx»ve 1 st-3rd alveoli, with slight concavities dvjr- 
sal at)d medial to it. In lateral view, ventral margin 
slightly convex up to U^th alveolus, nearly 
Mraighi posteriorly; dorsal edge rising srnoothly 
and increasingly steeply from the anterior tip to 
between 6th and 7ih iilveoli. highest part of the 
dorsal process (7th lo 9th) forming thedorsomed- 
ial surface for articulation with the prefrontal. On 
die posterior slope oi the process, a low promi- 
nence above the 1 1 ih alveolus probably the inser- 
tion site foi the posiorbital ligament, but may also 
mark the anterior extent of the jugal; by 1 3th bone 
very shallow, continuing so lo the posterior ex- 
tremity. Large lateral ( trigemmal) foramen open- 
ing anteriorly above the 5ih-6th alveoli, two 
smaller foramina, equally close to ventral edge, 
above 7th-8lh and 9ih I0ih.and3small foramina 
higher on the prefrontal process. Medial edge 
forming a shelf-like 'septomaxillary' process 
from 2nd lo 4ih alveolus, separated Irom the 
palatine process which widens gradually from 7th 
and then sharply at 10th. then gradually ap- 
proaches maximum width ;ii a sharply obtuse 
pnsteromediiil angle between Ilihand I2lli Me- 
dial shelf narrowing steeply from this |X)ini. then 
very gradually, but with a step-like inllcxion at 
level oi 1 Sih alveolus (marking location of ante- 
rior tip oiectopierygoid). Large foramen entering 
maxilla at broadest part of the palatine process, 
above 1 1 th alveolus, and a smaller foramen exits 
at the same level above the 7lh. Tooth row fol- 
lowing lateral margin closely from I si- 1 1 ih alve- 
oli, then gradually crossing over with I9ih-23rd 



closer to medial edge; lateral edge forming a low 
dorsolateral crest (possibly honxilogous with 
n)ore promii*eni cresii or bulges in snakes such 
as Difulysiii and pyihons). Lateral as well as 
medial pans of posterior maxsHa apparently over- 
lapped by the e^:top<;rygoid. forming slight con- 
cavities on cither side of a slight dorsal crest. 
Between ectoptcrygoid facet and prefrontal pro- 
cess, die suborbiiiil Mirfuce with a shallow longi- 
tudinal groove which probably either was, or 
bounded, a (acet for lJ>e jugal ^an element lost in 
extant snakes but probably retained in Dimlywi 
and madisoiids. including Wofuimhi\ Estcs el aL. 
!970.Scanlon, 1996). 

PARATYPES. Right and left mandibles 
tQMF31383. 31384). each compound and den- 
tary. in loose ariicvilation, lacking the splenial. 
angular and coronoid of each side (Figs 3, 4). 

Right TiXilh mw incomplete posteriorly, bn> 
ken through 1 5th alveolus; no sign of ankyloscd 
teeth. 4th to 8th largest alveoli, subequal. size 
reducing posterioriy and anterioriy. In laseral 
view, dorsal edge convex dorsad I'rom I si \o 3lh 
;ilveolus, concave dorsad for resi o( length Ven- 
tral edge slightly concave anterioriy. remainder 
convex but somewhat worn. Three mental foram- 
ina, below 3rd, 4ih and bih alveoli, openmg an- 
Icrodorsad. Posterior lateral fossa (compound 
notchj extending to between 10th and I lih Lat- 
eral face smooth hut with dorsolateral ridge de- 
fined by slight longitudinal concavity through 
foramina. In dorsal vievr, it>olh a>w concave inr- 
diad. slightly more so anterioriy; alveoli round or 
squarish except Hrsi two which are somewhat 
elongate transversely. 15th alveolus on a niurrow 
process distinguished by an angular concavity 
from the expanded dorsomedial shelf. The medial 
ridge forming the upper edge of Meckel's groove 
overhanging the grwwe distinctly up to the J<th 
alveolus; the overhanging edge of the upper hcci 
ior the splcnial beginning below the 8th bul mure 
dorsally, forming with a slightly acute, pointed 
posierovcniral process separated by a right-angle 
notch (in medial view) from the dorsal shelf. 
Meckclian groove narrowing anieriad, anterior 
end slightly expanded, conuviunicatirig by a fora- 
men with alveolus of I st lv)olh. Smooth bulb-like 
swcllingoverhangingthcgroovc medial to (he Isl 
and 2nd alveoli 

Left: Two mcntiil foramina, between 3rd and 
4lh. and 5lh and 6lh. Posterior lateral fossii ex- 
lending tti between 1 1th and !2ih alveoli. 

Right vompouixl. Elongate, shyllow. IR.Kttim 
long, 1 6.8mm from anterior tip lo dorsal exucm- 



400 



MEMOIRS OF THE QUEENSLAND MUSEUM 





B 




ily of articular facet. Sur- 
angular lamina low bui con- 
cave above, forming low 
coronoid process posterior 
to articulation with denlary, 
about 1/3 of length from an- 
terior tip. Maximum depth 
of compound less than depth 
of denlary at articulation 
(suggesting thai the 
coronoid extended dorsal to 
compound, forming most of 
the coronoid process by it- 
self). Ventral edge, and lat- 
eral in dorsal view, nearly 
straight, but posterior end 
(below articular facet and 
retroarlicular process) de- 
flected slightly ventrad and 
mediad from main shaft. Ar- 
ticular facet dorsal and me- 
dial in position, not 
extending to lateral face, 
reaching to middle of medial 
face, and as far an ten ad as 
ventrad from dorsal extrem- 
ity; facet defined posteriorly 
by a raised transverse lip, 
followed by a groove ante- 
rior to the sigmoid dorsal 
edge of the retroarticular 
process. Slight ventrolateral 
and deeper ventromedial 
concavities defining a ven- 
tral ridge on the retroarticu- 
lar process. Shaft of 
compound nearly cylindri- 
cal just anterior to articular 
lacct; a small dorsolateral 
foramen in this region. Man- 
dibular fossa narrow, begin- 
ning posteriorly at level of 

foramen, curved sli^^htlv FIG.4.A'a/;oHWfago^///e//j/sp.nov.,QMF31383,3l384,paratypes.compound 
mediad and extending to '°^^^ i'*^ bones, CS Site. A-C, Icfi compound in lateral, dorsolateral, and 
half way belwccn posterior "'if ^"i^ '""'^ missing articular). D-F, right compound in med,;.). dorsal, 

, / _! r . 1 and dorsolateral Views. Scale=5mm. 

edge ot coronoid tacet and 






top ot coronoid process. 
Fossa partly suirounded by the facet for the 
coronoid anteriorly; anterior half opening below 
into mandibular foramen, Surangular lamina 
curved, overhanging the mandibular fossa for 
most of its length; reducing in height anterior to 
coronoid process in two steps, reaching a hori- 
zontal or somewhat dorsally concave shelf re- 
ceiving the posterior part of the dentary; lateral 



edge expanded anterodorsally, for anterior 1/3 of 
length anterior lo the coronoid process. Surangu- 
lar foramen, opening anteriad, not exposed later- 
ally or medially, in a shallow dorsal trough 
between lowest point of surangular lamina and 
edge of coronoid facet. Facets for coronoid and 
angukir meeting at a very small angle below this 
point; their line of contact nearly horizontal, only 



NEW MIOCENH MADTSOIID SNAKE FROM RIVERSLEIGH 



401 





B 



D 




FIG. 5. Nanowana godthelpi sp. nov.. paratype. maxillae, CS Site, A-D, 
QMF3I386, in ventral, medial, dorsal, and laleral views. E-H. OMF3I380, in 
ventral, medial, dorsal, and lateral views. Scale=5mm. 



a short section preserved on either side. In lateral 
view the anterior edge of the compound rounded 
dorsally, separated by a right angle from a deeper 
ventral concavity. In medial view, a long, taper- 
ing notch enclosed in the facet for the angular, 
nearly reaching its posterior end (just posterior to 
middle of length of compound ). A medial antenor 
process (defuied by dorsal and ventral longitudi- 
nal fissures) hcanng the continuation of facets for 
the coronoid and angular, and probably also con- 



tacting the splenial and den- 
tary, broken on both sides. 
Left compound similar to 
the right, but broken posteri- 
orly through the articular 
facet. 

When placed in articula- 
tion, the right compound and 
deniary foiming a smoothly 
curved structure, with total 
straightlinc length appro-xi- 
maicly 29.5mm. 

Other paratypes and re- 
ferred jaw elernent.s (pariial 
dentaries, maxillae, pala- 
tines, pterygoids) show 
some individual variation 
(Figs 5, 6) and probably on- 
togenetic changes of propor- 
tions (allomeiry): the 
smallest deniary, QMF 
23056, is relatively deeper 
than larger specimens 
(Table 1 ). while the largest, 
QMF23076, is relatively 
slender except for a particu- 
larly deep upper posterior 
process. 

Vertebrae In shape and 
proportions, vertebrae sim- 
ilar lo. and intemiediate be- 
tween Yurlnnggur and 
Patagoniophis and differ 
conspicuously from Al- 
amitophis, Womimhi and 
Madisoia. Typical anterior, 
middle and posterior trunk 
vertebrae recognised (cf. 
LaDuke, 1991, Scanlon, 
1992, 1993); most anterior 
vertebra possibly 3rd cervi- 
cal {ci\ y. camfieldensis 
Scanlon, 1992. fig. lA). 
Centrum in ventral view rel- 
atively long, similar in pro- 
portions to Patagoniophis sp. but with the sub- 
central ridges nearly straight rather than strongly 
concave. Colyle slightly wider than the 
zygosphene, which is wider than the neural canal 
(all about equal in the most anterior vertebra); 
condyle and cotyle wider than deep, ventral mar- 
gins flattened in anterior and middle trunk, 
rounder posteriorly. 

Zygapophyscal facets inclined at about 20° 
from the horizontal (at mid-trunk; flatter anlcri- 



H 



402 



MEMOIRS OF THE QUEENSLAND MUSEUM 




orly. slightly steeper posteri- 
orly), defining planes pass- 
ing through the iniernal 
lateral ridges of the neural 
canal and intersecting jusi 
above its base. Facets 
broader and more angular in 
outline (especially the pre- 
zygapophyses) in the largest 
inidlrunk vertebrae, with 
long axes inclined at about 
45^" from the sagittal plane 
(somewhat more longitudi- 
nal in most anterior and pos- 
terior elements). Prezygapo 
physcal accessory processes 
lacking, oiiter face of the 
prezygapophysis with a but- 
tress-like ridge extending 
anicrolaterally to or slightly 
beyond the edge of the facet. 
Zygosphene shallower 
than the neural canal, with 
facets defining planes inter- 
secting below the iloor of 
the canal: dorsal edge in an- 
terior view flat, slightly 
aixlied or arcuate; below it 
are shallow concavities de- 
fining a dorsal ridge and lat- 
eral lobes, with sharp ridge 
separating the anterior face 
of the zygosphene from the 

mternal roof of the neural P*^- ^- ^anoyvanu godibelpi sp. nov.. referred elements from Upper Site 
canal h\ dorsal view the an- possibly from a single individual. A-D, left palatine. QMF23066 in lateral. 




^^1,.' 



^^y\y 



teriorly convex dorsal ridge 

and lateral lobes distinct in 

mid-trunk vertebrae, but in 

the most anterior and posterior elements median 

prominence less developed and zygosphene 

broadly concave. 

Paradiapophyses similar to Yuriwiggw or 
Patagoniophis, extending laterally beyond the 
zygapophyses only in the most anterior and most 
posterior vertebrae. 

Roof of zygantrum horizontal, either uniform 
in depth or thickening laterally, demarcated from 
the concave lateral parts of the neural arch by 
angular 'shoulders', with concavity directed 
more dorsally than laterally in the most posterior 
vertebrae because of the shallower neural arch 
and steeper postzygapophyscs. 

One or two small paracolylar foramina on ei- 
ther side of the cotyle. usually 2 lateral foramina 
on either side posterittr to the diapophyses. Sub- 



dorsal, dorsomediak and ventral views. E-H, right pterygoid, QMF23067. in 
ventral, lateral, dorsal and medial views. ScaJc^mm. 



central foramina usually smgle on each side, 
small. Parazygantral and zyganirai foramina 
larger, usually single on each side, frequently in 
distinct fossae. Some vertebrae with small foram- 
ina on Uie anterior face of the prezygapophysis 
below the facet. 

Ventral face of centrum concave between the 
haemal keel and subccniral ridges. In the anterior 
trunk hypapophysis projecting well below cen- 
trum from its poslerit)r half, with eitberan angular 
or sinuous anteroventral edge, and near- vertical 
posterior edge; in more posterior vertebrae the 
keel weakly sinuous to nearly straight in lateral 
profile. Haemal keel with median, kcel-likc 
hypapophysis reducing in depth from ihe cervical 
to mid-trunk regions; lateral ridges on the keel 
(initially just pt")Sterior to lite subcentral foram- 



NEW MIOCENE MADTSOIID SNAKE FROM RIVERSLEIGH 



403 






^^•^ 










m fmh^ yM^ 







5^ 



^1^,^> s'^'^-^ 
W 



979 






rSvO 




^^ 



\ii' „y 



wM am m 




FIG. 7. Nanowana godthelpi sp. nov,, QMF1974I. series of vertebrae from CS Site, possibly from Ihe same 
individual as ihc holot>pe (OMF31379). 



ina) from the approximate location of the largest 
venebrae in the skeleton, ridges increasing in size 
in more posterior vertebrae and posterior point of 
Ihe median keel fading away, leaving the ridges 
as paired hypapophyscs, venirolaieral swellings 
of (he keel. Haemal keel defined by smooth de- 
pressions in the anterior trunk, these becoming 
belter defined more posteriorly and approaching 
the cotylar rim. More posterior vertebrae with 
distmct channels between keel and subcenU'al 
ridges (subceniral paramedian lymphatic fossae, 
LaDuke, 1991). 

Most vertebrae from all regions of the body 
with swellings on Uie neural arch root on either 
side of the spine, forming short longitudinal 
ridges. Similar features in some Wonambi from 
Riversleigh are associated with small foramina 
(not the case here). Vertebrae similar to these and 
referred lo Nanowana sp. ( most of them probably 
N. godtheipi) from numerous sites at Riversleigh, 
including well-preserved examples from 
Wayne's Wok, Wayne's Wok 2, Mike's Me- 
nagerie, and Upper Site. 

Vertebrae of the cloacal region (Fig. 8) proba- 



bly from a single individual with short centrum, 
broad zygosphene. and condyle smaller than neu- 
ral canal (regional features allowing increased 
flexibility in this region). Haemal keel smooth 
(lacking the median ridge of Wonambi spp.). not 
or barely projecting below the centrum posteri- 
orly. Two largest vertebrae with paradiapophyses 
indicating articulated ribs, but on one side of one 
of ihcni the articular surface is expanded and 
roughened suggesting an immobile ciutilaginous 
attachment (i.e. transitional to fixed lymph- 
apophyses), Three others with lymphapophyses 
(broken disially); another with stumps of cylin- 
drical fixed ribs, possibly forkmg more disially. 

Nanowana schrenki sp. nov. 
(Figs 9-12, Table 2) 

MATERIAL. Holoiype QMF31395, a right palatine 
from early Miocene Upper Site, Godthclp Hill. Other 
Material: Upper Site; Maxilla fragments QMF31390, 
31391. 31394. Mike's Menagerie Site: Dentary 
QMF31392 and vertebra QMF23043. Camel Sputum 
Site: Dentary QMF23051; maxilla fragments 
QMF23082, 31388. 



4U4 



MEMOIRS OF THE QUEENSLAND MUSEUM 



T.\BLE 2. Mcasuichicnis of ETYMOLOGY- For 

Nanowana schrmki i>p.no\ . Fnedemann Schrenk, 
holoiype and referred jaw cl- Hcssisches 
emeuis. Abbreviylions as in Landesmuseum. 
Table I . wUh addition of dd8 Dannsiadl. for his en- 
=depthordenlar)'ail!!lhaIve- couragemenl and t\- 
olus. nanciai assistance for 

paiaeoniological co- 
operijliun hcli^fen 
Germany and Aus- 
tralia. 

DIAGNOSIS. Lm- 

eral pi\xcss of pala- 
tine ahout as long as 
4 alveoli (3-6), with 
dorsolateral margin 
strongly notched; 
ventral ridge of pal- 
atine maAillary pro- 
cess without distinct 
angular concavity, 
matching smooth 
edge of nnuillary 
palatine process. 
MaxilUi estimated to 
have about 19 tooth 
positions; palatine 
with IK pterygoid 
unknown, dentary 
18 (or 17-18). Teeth 
ankylosed normally; 
2nd to 4lh of den- 
tary. and 4th to 7rh 
or 8th of maxilla, 
mucti larger ihan 
oihcfH. Dentary 
touih row nearly 
straight in dorsal 

view. Three mental foramina, die third posterior 

to the 7lh tooth. 

DESCRIPTION. Holoiype. Alveoli 1 1, teeth an- 
kylosed in U 3, 4. 5. 6. 8.9. IfK 1 l;only ». 10 and 
1 1 complete Teeth with a simple curve, directed 
posteriorly. Tooth row deflected slightly medi- 
ally anieriorly, laterally posteriorly. Maxillary 
process slightly w ider than the tooth-bearing bar, 
extending from between 2nd and 3rd to between 
6lh and 7th teeth, with an anterioriy sharp lateral 
notch, and sharp posterolateral angle. Ventral 
surface of the process with a diagonal ridge from 
the rear of ;he 4ih tooth to the posicrolaieral 
atigie, defining an anterolateral ly concave facet 
to articulate with the palatal process of the max- 
illa. Anteriorly, the edge of the lateral process 



QMF 


31395 


31394 


23082 1 


INI), 


V\ 


Lf 


M 


pit 


4.7 


. 


. 


pel 


2.5 




- 


pill 


3-8 




- 


ptw 


2.5 






pew 


1.3 


- 


- 


prw 


0.6 


- 


- 


pmw 


0.6 


- 


- 


mpw 


^ 


1-7 


- 


mph 


_ 


2.0 


(1.0) 




Dem 


aries 




QMF 


23051 


31392 




Ind. 


CI 


MM 


mO 


3 


3 


dlt 


(16.7) 


17-1) 


dl)-. 


13.7 




dl)7 


15.6 




dlt 


12.6 




d4i 


- 


53 


d4i5 


9.3 


4.1 


a4,7 


11.2 


46 


d4f 


8.3 


3.3 


di? 


6J 




ddg 


2.3 


1.0 


dpp 


dJ 


0.4 



continuous wjih a dorsolateral ridge extending to 
the anterior tip of the tootlvbcaring prtxrcss. A 
second ridge diverging medially from the anlofo* 
lateral corner of the process, hearing a distinct 
knob above the tooth row and continuing onto the 
anterior edge of the choanal process, level s^-ith 
die rear of the 4th alveolus. Antcromcdial comer 
of choanal process (to articulate with posterior 
process of vomer and possibly parasphenoid) 
missing. Medial edge intact, and smoothly con- 
vex, from level of 7th alveolus to rear of tooth 
row. but posterior process broken off. Cusp dc- 
Hning lateral edge of choanal trough diverging 
posieromedially from the 6th tooth, disappearing 
level with the 8lh: 2 foramina close together' in 
the space between and medial to 7th and 8ih 
alveoli, one of them piercing the choanal plate to 
emerge dorsally in a more medial position and 
opening medially. Toolh-bearing bar pointed 
posrerioriy, tapering from the 9th tooth, a broad 
parabolic surface for the retractor ptcrygoidei on 
the ventral face with its apex beside the 9th, 
becoming less distinct poslcrolaieraliy. Deep 
notch U) aniculate with the pterygoid on the dor- 
sal side between the tooth row and posterior 
process, extending ro al>ovc the anterior edge of 
the 1 0th tooth. Distinct growth lines through the 
translucent cht>anal plaie parallel to its curved 
medial edge. 

Referred material. Maxilla represented by sev- 
eral fragnieniary specimens from different sized 
individuals (Fig. 10). Tooth row curves mcdiad 
anteriorly (QMF23082), with a so-ong gradient of 
increasing alveolar diameter from 1 to 5; 5 and 6 
suhcqual. Dorsal edge is a sharp, concave 
dorsomcdial crest, extending to a high dorsal 
pR»cess, jevellmg off above 6ih alvcoius; this 
crest divides anieriorly, enclosing a shallow 
trough above the first two alveoli (thus, maxilla 
partially flooring narial cavity). Lateral face 
mostly convex, with a shallow longitudinal 
trough including a large foramen (opening an- 
teriad and ventrad) above rear of the 4th rooth; a 
smaller foramen near the dorsal edge at>ovc the 
5th. Medial face concave, with a u-ough jast 
below the dorsomcdial ridge containing a small 
foramen just anterior to the medial one. Middle 
part of maxilla (QMF3i3';)4) with distinct knob- 
like posterior pan of prefrontal process and slop- 
mg suborbital portion, becoming more rod-like 
and wider than high poslcrioriy. Tooth si/c de- 
creasing sharply, with increased alveolar spacing, 
just behind pi-efronial process; longest (7lh or 
8th?) 2.2mm long, curved at middle but straight 



NEW MIOCENE MADTSOIID SNAKE FROM RIVERSLEIGII 



M\5 



distally, with medial and lat- 
eral cutting ridges (like 
loneest tooih of dentarv 
QIvrF31392. see below)"; 
more posterior teeth (broken 
before drawing) with simple 
curve, about half as long. 
Palatine process diverging 
from tooth row at last large 
looih and reaching maxi- 
mum width between the next 
2 alveoli. Medial edge of 
Ihc palatine process quite 
smooth, matching the con- 
cavity of Ihc maxillary pro- 
cess in the holotype; large 
opening on dorsal face of 
process for palatine nerve 
and blood supply through 
several foramina on lateral 
surface. Teeth on posterior 
part ofmaxilla(QMF3 1391) 
still reducing in size from 
anterior to posterior, and 
with slight double curve. 
Posterior part triangular in 
section, with near vertical 
lateral and oblique 
dorsomedial faces both 
slightly concave, meeting at 
a dorsolateral ridge. Lateral 

edge straight, medial edge FIG, 8 ..V£jAjoivartft^c>£///?t7/>/sp. nov.. series of most posterior tmnk and cloacal 
produced as rldiie with con- vertebrae from Upper Site, possibly from the same individual as jaw elements 
vexily probably marking an- 




in Fig- 6. Lateral, posterior, dorsal, and venu*al views. 



lerior Umii of ectopterygoid. 

Dentaries. Two right dentaries, differing con- 
siderably in size (Fig, 1 1). represent the lower jaw 
in this species. 0MF3 1 392 w ith complete row of 
18 alveoli, teeth ankylosed in 1 (possibly), 3, 6. 
8- 10, II. 13, 15. 16. and IS; lOlh broken, other 
Tccih in tact, and a replacement tooth apparently 
in sun behind 15th. QMF2305I has 17 alveoli, 
but another may have been present posteriorly: ! . 
4,5,6.7,9, 11, 12, 13. 14 and 15 ankylosed, but 
all teeth broken near base (the jaw has also been 
broken tlirough 3rd alveolus and subsequently 
healed in life), 1st alveolus approximately same 
size as 5lh, but 2nd to 4th considerably enlarged; 
3rd nearly twice diameter of 5th, size decreasing 
gradually more posteriorly; in the small speci- 
men, lengths of teeth from anterior edge of base 
to tip (mm) -, -, 1 .26, - -, 0.61, -, 0.63, -, -, 0.55, 
-, 0.52, -, 0.40, -, 0.37. 0.28. Anterior alveoli 
(J-3)dencctcd vcntrad and mcdiad relative to rest 
of tooth row, which is moderately concave dorsad 



but only very weakly concave mcdiad. ThitxJ 
tooth directed slightly laterad as well as posie- 
riad; other teeth mediad, more strongly towards 
the rear of the tooth row. Each tooth with a weak 
lateral and medial cutting edge near the lip Den- 
tary deepens gradually from anterior to posterior. 
Three mental foramina open anteriad below alve- 
oli 4, 7 and 9 (QMF31392) or 3, 6-7 and R 
(QMF23051), decrea.sing in size posteriorly. A 
shallow dorsal trough medial to 3rd and 4lh alve- 
oli defined by a dorsomedial crest. Lateral fossa 
extends as far anteriorly as the rear of the 13lh 
tooth, blunt in outline; posterior edge of the ver- 
tical iniramandibular septum smoothly concave, 
extending forward to between the 14ib and I5lh 
teeth. Differences between the two include shape 
of Meckel's groove (tapering more strongly in the 
small jaw, dorsal edge composed of two shaqMy 
defined sections separated by a short gap below 
8ih-9th dlveoli. but no gap in the larger speci- 



406 



MEMOIRS OF THE QUEENSLAND MUSEUM 





FIG. 9. Nanowaria schrenki sp. nov., hololype, 
QMF31395 from Upper Site, palatine in ventral (A), 
dorsal (slightly lateral) (B). dorsomedial (C), and 
lateral (D) viewi>. Scale bar=2Tnni. 

men), upper facet for splenial (with posteromed- 
ial angle in the smaller, but a free-ending prcx;ess 
in the larger), and lateral fossa (constricied in the 
smaller by deepened upper posterior process); 
both narrowest in the region of the 6ih to 8lh 
alveoli: but the larger specimen is relatively 
broader posteriorly. 

Vertebra (Fig. 12) from mid-trunk of a juvenile, 
witii short broad centrum, large neural canal, and 
condyle and colylc much wider than deep. 
Weakly defined subcenlral ridges narrow only 
slightly behind the parapophyses, posterior half 
of centrum nearly parallel-sided except for a shal- 



low, short precondyjar constriction. Blunt haemal 
keel extending from just behind the cotybr rim, 
postenorly forming a slightly prominent single 
hypapophysis extending below the condyle. Keel 
defined laterally by broad shallow depressions. 
Comparisons with Yurlunggitr or Patagoniophis 
would imply that a haemal keel of this fonn 
indicates a vertebra from close to the cardiac 
region (transitional between prominent single 
hypapophysis anteriorly and llallened or double 
keel posteriorly), and would thus be among the 
largest in the skeleton. Condyle and colylc about 
twice as wide as deep, slightly oblique in lateral 
view; cotyle wider than the neural canal bui not 
as wide as the zygosphene. Zygapophyscal facets 
inclined at less than 20"" above the horizontal, 
defining planes which intersect near the middle 
of the neural canal. Prezygapophyseal facets ob- 
ovale, with transverse anterior edge; ptxstzyg- 
apophysea! facets more smoothly oval, and 
somewhat prominent poslerit^rly in dorsal view. 
Both pairs of facets arc elongate anteropost- 
enody, with long axes at about 35^ to the sagillal 
plane (as in anterior, but not middle trunk verte- 
bra of Patagoniophis sp. cf. P. paryus; Scanlon, 
1993). No prezygapophyseal processes. 

Paradiapophyses directed ventrolaterad, 
slightly wider than prezygapophyses, not extend- 
ing ventral to colylar rim. Interzygapophyseal 
ridge smoothly concave laterally, only slightly 
wider than the centrum, and weakly defined in 
lateral view. 

Zygosphene tfiin. slightly arched; anterior edge 
smoothly but weakly concave (again, like ante- 
rior rather than middle vertebrae of Paiai^oni- 
opliis). Zygosphenal facet (preserved on left 
only) dorsovenirally shallow, with dorsiilly ctm- 
vex upper and lower edges, inclined at about 45*^ 
from vertical; a plane tangent to the facet would 
pass close to the centre of the neural canal. 

Neural canal arched, about as high as wide, 
lacking internal lateral ridges. Neural arch low, 
with shallow concavities above and belou the 
level ol the zygosphene and extending to the 
posterior edge. Zygantral roof arched, thickness 
uniform across its width. In dorsal view, rear of 
neural arch forming a broad concavity above (he 
zyganirum, interrupted by the neural spine. Low 
neural spine fomied by a narrow, but sharply 
defined anterior lamina rising from the rear of ihc 
zygosphene and applied to a higher, columnar 
ponion posteriorly, overhanging the zygantrum. 
Dorsal surface of column broken off. with a sinus 
wisible within the neural arch. Lateral and sub- 



NEW MIOCENE MADTSOIID SNAKE FROM RIVERSLEIGH 



407 



centra) foramina preseni, 
any olher obscured by den- 
drites. 

TROPHIC 

SPECIALISATIONS 

OFNANOWANA 

N. godthelpi sp. nov. The 
homogeneity in size, mor- 
phology and approximate 
siratigraphic position of 
these toothless but otherwise 
well-preserved jaws makes 
it appear probable thai the 
lack of ankylosed teeth is a 
natural (and apomorphic) 
characteristic. To quote 
Owen's (1840) conclusion 
on the 'dislocated' lail of 
ichthyosaurs, the toothless 
condition '... is too uniform 
and common to be due en- 
tirely to an accidental and 
extrinsic cause'. Variation 
in the shape and size of alve- 
oli along the tooth rows, and 
the presence of iVothy' bone 
similar to bone of attach- 
ment in some cases, indi- 
cates that different stages of 
replacement are repre- 
sented, so that absence of 
teeth is not explained by 
synchronised replacement. 
Some of these specimens are 
practically intact, preserving 
delicate processes, and not 
worn in such a way as to 
account for the absence of 
even stumps of teeth; in 

most other specimens from f . jg ^ j 

the same deposits, parts of ' ~ "*"*' 

teeth are typically retained 

even after heavy wear. The 

alveoli are shallow, rather ^*^- 10. A^a/ioH'flnajr/irpnA-(Sp. nov.. maxillary fragments. A-C,QMF3 1394, 

rprnnaiilqr nits 'in rh^f q ^pper Site, middle part of right maxilla in lateral, dorsal, and medial views. 

kI .Tl. , i^l .V !i« 1*1.. D-G, QMF23t)82. CS Site, anterior right maxilla in lateral, dorsal, medial, and 
Ihecodoni type ol implanta- ^,^^^^^, ^.^^, ^_j QMF3I391. Upper Site, posterior left maxilla in medial, 
tion IS not mdicated as an ventral, and lateral views. Not to same scale, 
alternative to ankylosis. 

Failure of teeth to anky- 
lose at any stage is rare among squamaies, first 
reported by Savitzky (1981). Anomochilus 
weheri, a small fossorial 'anilioid' (An- 
omochilidae is possibly the sister taxon to iHher 




m^^ 



;^5»feSO^ 



^^^^ 




living Alethinophidia; Cundall et al. 1993), ap- 
parently has fibrous tooth attachment rather than 
ankylosis (Cundall & Rossman, 1993). There are 
also several lineages of snakes, and one genus of 



408 



MEMOIRS OF THE QUEENSLAND MUSEUM 







^^^.ff:^^ ^^^^o,. 




./: 



^^it^ T'WTSsc- 



i^... 



■ut^^^^^^^- 




^^-^T^^amz<;i¥^^^^ 



x^"^'^^^ 



FIG. 1 1. Nunowana schrenki sp. nov- righl dcnlnries. 
A-C. QMF2305I, CS Site, in medial, dorsal, and 
lateral views. D^F. QMF3 1392, MM Site, in medial, 
dorsal, and lateral views. Scale bars=2 mm. 



lizards, where the attachment is not only fibrous 
but forms a funclional hinge allowing each tooth 
to told posteriorly under pressure and return up- 
right when released (Savitzky, 1981, I9S3; 
Paichell & Shine 19S6c; cf.Edmund, 1969:141). 
This hinge mechanism has been interpreted in 
each case as an adaptation lo feeding on scincid 
or gerrhosaurid li/ards in which the scales are 
underlain by osieodcrms; the hinged teeth are 



thought to act as a ratchet mechanism, folding 

back rather than penetrating the dermal armour, 
and locking in an upright position against the 
edges of the scales when the prey is oriented 
head-first for swallowing. In extant snakes other 
functionally associated apomorphies also occur; 
the teeth are sinall and numerous, often with a 
spatiilate rather than conical tip. and lack enamel 
on the posterior surface; and the levator anguli 
oris muscle (inserting on a long upper posterior 
process of the denlary) is enlarged (Savii/ky, 
1981). In the pygopodid Lialbtceih are tif similar 
form, and instead of increased intramandibular 
kinesis there is pronounced kinetic ability at the 
frontoparietal joint (me.sokinesis: Patched & 
Shine 1986b). Both types of kinesis allow the 
jaws more effectively to surround and compress 
a cylindrical prey item, immohihsing or even 
asphyxiating it. An equivalent adaptation for 
prey-holding (without hinged teeth) is seen in the 
largely scincivorous bolyeriid snakes, in which 
the required kinesis is provided by the uniquely 
derived inlramaxillary joint (Cundall & Irish, 
1989). 

Savitzky (1983) described this set of adapta- 
tions to feeding on skinks. which has evolved 
independently in several lineages, as an instance 
of a 'coadapted character complex', among other 
cases of 'durophagy' (feeding on hard-bodied 
prey). Other durophagous snakes have distinct 
specialisations, and feed on other kinds of 'hard* 
prey such as snails (pareine and dipsadine colu- 
brids) or crabs (the homalopsine Fordnnia)^ 
'Durophagy* is thus a broad concept. I introduce 
'arihrodonty'to refer specifically to the "hinge- 
toothed' modeof durophagy. 

While soft-tissue structures such as fibrous 
hinges cannot be observed in fossils, absence o^ 
ankylosis implies that attachment was fibrous and 
potentially ficxible. M godthelpi ydw material is 
similar to that of extant arthrodonr species alter 
maceration, especially Xenopehis (Savitzky, 
pers. comm.). Hutchinson (1992) demonstrated 
that scincid lizards were abundant and diverse in 
the Tertiary at Riversleigh; skinks today repre- 
.sent a major food source for small terrestrial 
predators, including most extant Australian snake 
species (Shine, 1991). As funclional aiihrodoniy 
has evolved in several lineages in association 
with predation on skinks, its presence in N, 
godihelpi is a plausible explanation for the lack 
of ankylosis. 

N. godihelpi appears to be less specialised than 
each of the extant anJimdont snake lineages in 
some respects. The high number of ncariy uni- 



NEW MIOCENE MADTSOIID SNAKE FROM RJVERSLEIGH 



409 







FIG. 12- Nanowana schrenki sp. nov.. vertebrae. 
QMF23043. MM Site, probably Same juvenile as 
deniary QMF31392 (Fig. II) in anlcrior(A}. postc- 
rior(B), lateral(C), dorsa](D) and ventml (E) views. 

lorm maxillary alveoli is typical of an arlhrodont 
species, bul a similarly long lotuh row is present 
in Wonawhi naracoortensis (Barrie I99U). and i^i 
therefore likely to be a retained ancestral condi- 
tion rather th:in a specialisation. The overlap be- 
tween dentary and compound is moderate, 
without any great elongation of the tooth-bearing 
posterior process; the extreme condition in extant 
arthrodonl lineages may be precluded by ihc 
probable insertion oi'm. levator an\iuH oris ox\ the 
iclaiively large ctironoid rather than the dentary, 
but the overlap is actually shorter in this species 
ihan m Wonambr 

The dentary alveoli of M godjhelpi are consid- 
erably larger, especially in the middle part o{ the 
row, than those of the maxilla, so the lower leeih 
may have functioned differently, and possihiy 
lacked a functional hinge. In Xenopeltis, rcla- 
lively laige teeth are present on the middle pari 
of the palate (posterior palatine and anterior pter- 
ygoid), hui these appear to be fully hinged. While 
fibrous attachment of 'sessile' leelh has been 



reported only in one highly unusual extant (axon, 
Anomochiius, il is possibly a necessary precursor 
or incipient stage of arthrodonly (see below), and 
a mixe^j or * semi -arthrodonl* condition in N. 
godtlu'lpi seems possible. 

Nanowana schrenki sp. nov. In the absence of 
articulated or strongly associated material, refer- 
ral of jaw elements described here to a single 
taxon can only be provisional. In particular, ihe 2 
near-coniplele dentarics differ in several respctls 
which make their assignment to the same species 
doubtful: in QMF23l)5i the upper edge of the 
Meckclian groove is a continuous ridge and ex- 
tends posteriorly as a free-ending process, while 
in QMF31392 it Is interrupted at the '^ih alveolus, 
and appears (o end abruptly. (Additionally, the 
larger specimen broadens more posterii)rly . while 
the small one is widest at the 3rd looih. but this 
difference may be allometric. ) 

The teeth of snakes play several roles in the 
capture, subdual puncturing or laceration, and 
swallowing of prey; in general they will he 
adapted fi)r a combination of functions, but often 
either a single runt:iion is di)minant. or ceiiain 
stages arc cither not required (e.g. because inac- 
tive or defenceless prey is taken) or carrietl out 
cxlra-orally (e.g. constriction). Teeth specialised 
for different functions are often separated be- 
tween the front and rear of the mouth, in some 
cases with diastemaia between leelh of different 
morphology (Fraz/ella. 1966: Scanlon & Shine, 
l9Sfi,Cundall& Irish. 198^). 

Numerous terms have been mtroduced fordif- 
fercni patterns of loolh si/.e and fang location 
(Smith 1952). Primitive snakes {DtnUysia. an- 
ilioids) are isodont or mesodont. with relatively 
few, stout teeth; while also capable of consiric- 
tionv they use a powerful 'crushing' hiie in sub- 
duing prey (Fra/./.etta, 1 970; Greene. 1983). Such 
a 'crushing' method seems possible for Mcuitsoia 
cW M h<ii, in W'hich the dentary is heavily built 
and bears relatively lew teeth (Hoffstetter, 1960). 
but not for Australian madlsoiids. Different pat- 
terns ol loolh-sizc variation in upper and lower 
jaws are known in each of the 4 best- represented 
taxa: 

In Wonanibi naracoortensis the very numerous 
leeih (25 m the dentary. 22 or 23 in the maxilla) 
are proierodont, sharp and strongly inclined ptxs- 
teriorly and medially (Barrie, 1 990); the jaws are 
shallow, suggesting a limited role in subduing 
prey, and more emphasis <in holding and swal- 
lowing functiiins. This implies thai an extra-oral 
method of subduing prey (probablv cvnisIficlioiM 
was well-dcvciopcd. When the upper and lower 



410 



MEMOIRS OF THE QUEENSLAND MUSEUM 



jaws are both proterodonl, teeth often have a 
sigmoid curvature with the lips directed some- 
what anteriorly as in many pythons (Frazzelta, 
1 966), and seems to be associated with relatively 
soft-bodied prey such as mammals, birds, earth- 
worms (McDowell, 1969) and eels (Smith, 1926; 
Cogger etal., 1987). 

Nanowana godrhelpi apparently had a nearly 
isodont marginal dentition. No complete tooth 
crowns have been reported for this species, but 
based on alveolar sizes it was weakly proterodonl 
on the maxilla and mesodont on the dentary (Figs 
1,3). 

The condition in Yurlunggur is less clear but 
apparently the opposite; a dentary with well-pre- 
served teeth (Archer et a!., 1991 :71 ) is proterod- 
ont, while the maxilla was apparently mesodont 
(Scanlon, 1996). 

N. schrenki can be described as megadont 
(Smith, 1952), having regions of distinctly en- 
larged teeth. Otherwise it has the same pattern of 
enlargement as Yurlunggur, opposite to that of M 
godthelpi, being mesomegadont on the maxilla 
and promegadont on the dentary. The dentary is 
relatively longer and less robust than in Madtsoia 
or Dinilysia, but not depressed as in Wonambi\ 
the teeth are intermediate in number and in mor- 
phology (stouter and more erect than Wonambi, 
but not so much as in Dinilysia or anilioids); and 
the enlarged teeth are a uniquely derived condi- 
tion within Madtsoiidae (albeit convergent with 
many other lineages of snakes). 

Many snakes share this pattern of enlarged 
teeth at the front of the dentary and the part of the 
maxilla below the prefrontal articulation, 
whether or not they are set off by diastemala or 
local minima of tooth size. On the basis of occur- 
rence in scincivorous colubroids such as 
Lycodon, Glyphodon, Demansia, and Hemiaspis 
signata (but not the anurophagous //. dcimeli; 
Boulenger, 1896; Worrell, 1961; Shine, 1991; 
Cundall & Irish. 1989; pers. obs.), this is here 
tentatively considered an adaptation to hard-bod- 
ied prey, often skinks. Snakes with enlarged teeth 
offset between upper and lower jaws are able to 
trap hard, cylindrical prey items between a notch 
in one tooth-row and one or more enlarged fang- 
like teeth (sometimes true fangs) in the other 
(Cundall & Irish, 1 989). As well as this 'trapping' 
function, having only a few long teeth in each jaw 
maximises the probability of hard-bodied prey 
being deeply punctured, whereas this is avoided 
in arthrodont forms. 



EVOLUTION OF TEETH 
AND ATTACHMENT 

Snake teeth are slender compared to other ver- 
tebrates; they break frequently during normal use 
and are quickly replaced (Edmund, 1969). The 
have reduced occlusal area (sacrificing strength) 
to increase sharpness and depth of penetration. 
Tooth form is a compromise betwen competing 
selective forces defining a 'fitness landscape' 
over attainable phenotypes (Wright, 1932), and 
local optima will be attained only if intermediate 
states are evolutionarily stable. If the rate of 
breakage is too high, prey capture or swallowing 
efficiency (and consequently fitness) will be low. 

During the stages of feeding on a given range 
of prey types with given neuromuscular reper- 
toires, forces on the tooth come from particular 
directions with greater or lesser frequency and 
magnitude, so it will generally be favourable for 
the tooth to be asymmetrical rather than a simple 
cone. The orientation of 'cutting ridges' (which 
function as buttresses as well as blades), curves 
in the shaft, and the shape of the tooth base, will 
confer maxima of resistance in one or more direc- 
tions, at the expense of minima elsewhere. 

Horizontal components of pressure (shear 
stress) at the tip ofan approximately conical tooth 
are converted to bending stresses at the base, i.e. 
compression at one side and tension at the other. 
The magnitudes of these forces will depend on 
base diameter, but only tension and shear will 
tend to either break the shaft or disrupt the attach- 
ment of tooth to bone. Bone of attachment can 
apparently withstand such stresses within a wide 
range of values of the ratio of tooth length to basal 
diameter. A fibrous connection will remain stable 
at low values of this ratio (short, broad teeth as in 
Anomochilus), and at intermediate values will 
have enough elasticity to return the tooth upright 
after displacement (functional arthrodont condi- 
tion). At high values (longer, slender teeth) a 
fibrous attachment would merely bend passively, 
without developing enough tension to right the 
tooth; the orientation of the teeth would then not 
be precisely controllable, and during prey capture 
and ingestion they would more often encounter 
shear stresses at unfavourable angles, leading to 
rupture. Such a condition (elongate, slender teeth 
with fibrous attachment) is unknown in any living 
snakes, and would presumably be evolutionarily 
unstable for most diets and feeding methods. 

This consideration of the forces applied at the 
tooth tip and base suggests that arthrodonty and 
elongate teeth are mutually exclusive conditions. 



NEW MIOCEKEMADTSOIID SNAKE FRO^f RIVERSLEIGH 



411 



Thus ihc specialisations of dcniition and jaw mor- 
phology in Ncmowivui arc most likely to be inde- 
pendently derived from ihe nearly isodont. 
ankyloscd condition ul' other madlsoiids. and ap- 
parently represent allernaiive solutions to the 
problern of feeding on hard- scaled lizards. 

Healed breaks ofthe jaw elements (pnicularly 
dentaries) are noi uncommon in snakes (pcrs. 
obs.). and presumably result in niosi cases ra>m 
auempis lo capture or subdue relatively large and 
powerful prey. Sublethal irauma associaied wiiii 
particular morphological specialisations may be 
an indicaiorol'mcchanismsorselcction; there arc 
upper limits to prey si/e and slrcnglh for every 
species of prcdau^, and both prey selection and 
behavioural aspects of prey-handling, as well as 
morphology, will be subject lo selection. The 
break through the third dcniary alveolus ol 
QMF23051 {N. schrenki sp. nov.) would have 
occurred most easily (i.e. greatest stress would 
occur) while the 3rd alveolus was unoccupied, 
and while a prey item was held by the enlarged 
2nd tooth, hut not the smaller posterior teeth. 
Fractures of ihis kind c<JUld be expected to be less 
tnmmon (all else being equal) with a more uni- 
form dentition, but this possible disadvantage of 
raegadonty may have been outweighed by an 
increased rate of capture success, or of reienlion 
once a prey item was secured behind (or impaled 
on) the enlarged dentary teeth. 

The ribbon-like posterior maxilla of A'. 
godthelpi presents an even more Iragiic appear- 
ance, but no .specimens suggest brcaksduring life. 
While this is negative evidence, the rarity of such 
breaks would tend to support the presence ul a 
jugal in the suborbital region. Prcsetu'cof ajugal 
in Wonatnbi naracoonensis can similarly be in- 
ferred from the oblique trough crossitvg the max- 
illa (Barric, 19*^0; Scanlon. 1996) which would 
otherwise be an obvious point of fragility. 

SYMPATRY OF RELATED SPECIES WITH 
SIMILAR DIETS 

The two Species of A/iancnnfl/wi occur together 
in at least 3 Sites, existing sympatrically for a 
significant period. They are tlioiighi to have had 
siniilardie[s(skinkij),andsimilaradullsi/e,They 
ihus occupied quite similar niches, and were 
slriclly equivalent ecologically. They may have 
differed in aspects of behaviour which would not 
be di.sccinible in the fossil fecofd> but at least a 
difference in habitat can be suggested. 

The diffcreni representation of the two species 
when found together (minimum number ot indi- 



viduals, number of identifiable elements, and 
quality of preservation) implies that A', godthelpi 
was more abundant close to the sites of dcpisi- 
tion, whereas M sclttenU may have been less 
abundant locally, and the more damaged rettuins 
transported from further allckl (cf. LaDukc, 
I yy I ). Thus N. godihelpi lived near water (possi- 
bly riparian, probably closed forest), whereas ,V. 
achrenkf may have lived further Irom water, pos- 
sibly in more open or drier area.s .such as cicaringii 
or Ricky hills. 

Most sues where Nanowana vertebrae ha^'C 
been found have not produced jaw elements di- 
agnostic in species. The genus as dellncd here, 
therefore provides a convenient level of descrip- 
tion which can be applied to a larger set of !»itcs, 
but as yet all specimens relerred to Nonowana arc 
fromRiverslcigh 

ACKNOWLEDGEMENTS 

f thank Mike Archer for the opportunity lot 
study Riversligh foSMis under his supervision* 
and Henk Godthelp, Sue Hand, Anna Gillespie. 
Jcaneltc Muiihcad, Sy|) Pra.soulhsouk, and Sic- 
phan Williams forprcparalion and other activities 
in field anil lab which made ihis work possible. I 
also Ihank Mike Archer, John Barric, Dino Frcy, 
Mark Huichmson, Mike Lcc, Ralph Mt^lnar, 
Jean-Claude Rage. Alan Savitzky, Rick Shine. 
Zbignicw S/yndlar, imd Paul Willis, tor insights, 
discussions, and access to material; and Wighardl 
von Koenigswald and Fricdemann Schrenk for 
facilitating visits lo Germany where much of the 
paper was written. Support for research ai 
Rivcrsleigh has come Irom the Australian Re- 
search Grant Scheme; the National Estate Grants 
Scheme (Queensland); the University of New 
Soud) Walcb. the Commonwealth Department of 
Environment, Sports and Temfories; the Queens- 
land National Parks and Wildlife Service; the 
CoiDrrionwealth World Ueritago Unit; ICI Aus- 
tralia; the Australian Gci>graphic Society; the 
Queensland Museum; the Australian Museum; 
the Royal Zoological Society of NSW; the 
Linncan Sncicty of NSW; Century Zinc; Mount 
Isa Mines; Surrc) Bcatly & Sons; the Rivcrsleigh 
Society; and private supptxters including I:laine 
Clark, Margaret Bcavis, MtUtin Dickson, Sue & 
Jint Lavarack and Sue & Doii Scoit-Orr Vital 
field assistance came from many hundreds of 
volunteers as well as staff and postgraduate stu- 
dents of the University of NSW. Skilled preivim* 
tion of most of the Riversleigh material has been 
carried Out by Anna GiUcsi?ic. 



4): 



MEMOIRS OF THE QUEENSLAND MUSEUM 



LITERATURE CITED 

ARCHER. M.,GODTHELP. H., HAND. S J & MEG- 
IRIAN, D. 1989. Fossil mammals olRivLTNleigh, 
norihweslcm Queensland' prclin>inary nverviLw 
lit biu.slraligraphv, correlalion anU enviionmcutal 
change, Australimi Zooloei^l 25: 27-65. 

ARCHER, M.. GODTHELP. H. & HAND. SJ. 1991. 
Riversleich 2ncJ Edition. (Reed Books: Sydney). 

BARRIE, D.r 19W Skull elements and a.ssociaied 
rcniains dI ihe PIci.slOL-cnc hold snake Wonambi 
na racoon en si.K. Memoirs tjf the Queensland Mu- 
seum 28: 139-151, 

BOULENGER. G.A. 1 8%. Caialugue oi ilic .snakes in 
the British Museum (Natunil History. III. O'aylor 
& Francis: Londtin). 

COGGER. H.G.. HEATWOLE. H.. ISHIKAWA. Y., 
MCC0Y,M,TTAM1YA,N &TERUUCHI.T. 
1987. The status and natural tiisrory of tlie RcnncI 
Island sea kniit, LLiiUttudit crachsri (ScqKniCs; 
Laiicifudiducj Joumiil of Hcrpclology 21: 255- 
266 

CUNDALL D.& IRISH. F.J. 1 989. The function of ihe 
intrarnaxiilary joint in Ihe Round Island bua. 
Casuren thissimtieri. Journal of Zoology, London 
217:569-598. 

C13NDALL. D »t ROSSMAN. D S- 1993^ Cephalic 
aiialomy of the rare IndiJne.sian snttkc Anomochils 
^ehef. Axilogical Journal of the Linnean Society 
109: 235-273. 

CUNDALL. D.. WALLACH. V. Hl ROSSMAN. U.S. 
1993 The systematic relationships of the snake 
genus Anomochilus. Zoological Journal of the 
Linnean S(Kicty K)9- 275-299. 

EDMUND. A.G. 1969. Denlilion. Pp. i 17-197. In 
Gaas, C & Parsons^ T S (ed.s). Biology of the 
Rcptilia I. (Academic Press: London). 

ESTES. R.. FRAZZETTA. T.H. & WILLIAMS, E.E. 
1970. Studies on the fossil snake Dinilysk 
putaj^ouiai Woodsvaid. Part 1. Cranial morphol- 
uey. Bulletin of the Museum of Comparative Zo- 
»:iTogy.nnrvnrd 140:25-74. 

FRAZZETTA, T.H. 1966. Sludics on ihe morphology 
and function of the skull intheBi)i(lac(Serpenlc.s), 
Part li. Moq^hology and lunclion uf the jaw appa- 
nttus in Pyihoti Krhat' and Pyrhon tnohirus. .roumal 
Of Morphology J 18: 217-296. 
1970. Studies on the fossil tinixke fJhnlysia 
fiOfa^ontcii Woodward. II. Jaw machinery in the 
earliest snakes. Fotina el Funciiu3: 205-221. 

GREENC, H.W. 1983. Diclary correlates of the origin 
.and radiation of snakes. American Zoologist 23: 
431-441. 

HOFFSTETrCR. R. I9C*0- Un dentaire dc Mudtsoio 
(.serpent giJant du Palt-oeene de Paiagcmia). Bulle- 
tin du Mu5(5um national dTlistoire naiurelle. Paris 
(2)31:379-386. 

laiTCmNSON. M.N. 1992. Origins of the Australian 
seincid ti/ards: a prehminary report on (he skinks 
of Riverslcigh. The Beagle 9: 61-69, 



LADiKR TC. 1991 The fossil snakes of Pit 91, 
Raiidu) La Brea. Caliiomia. Natural History Mu- 
seum of Los Angeles County, Cuntnhulions in 
Science 424; 1-28. 

MCDOWELL. SB. 1969. ToxicocaUtmus, a New 
Guinea genus of snakes of Ihe fiimily Flapidae. 
Journal of Zoology, London L59: 443-51 1. 

OWEN. R. 1 840. Note on the dislocation of the tail at a 
ceriam point observable in the skeleton of many 
ichthyosauri Trtmsaclions of the Geological St>- 
cieiy K^i London 5: 5 1 1 -5 1 4 

PATCHBLL. F & SHINE. R, 1986a. Feeding mecha- 
nisms in pygopodid li/ards: how can Liali.s swal- 
low such large prev':.' Journal of Herpeiology 20: 
59-64. 

PATCHELL. F C ik SHINE, R. 1986b. Hinged iccih 
lur hard- bodied prey, a case of convergent evolu- 
tion bclwern snakes and legless li/^rds. Journal of 
Ziulogy, London 208: 269- 275. 

SAVITZKY. A.H, 1 981 Hinged teeth in snakes: an 
afldaplation for swallowing hard-bodied prcv, Sci- 
ence 212: 346-349. 
1983 Coadapted character complexes among 
snakes: fos&oriality, piscivory, and duaiphagv- 
American ZvH>logisl 23. 397-409. 

SCANLON. J.D. 1992. A new large madl.soiid snake 
from ihe Miocene of the Northern Temlory. Tlie 
Beagle: 9: 49-60. 
1993- Madtsoiid snakes from the Eocene Tinga- 
mtirra Fauna of eastern QuecnsUind. Kaupia. 
Darmsiadier Beitrage zur Natnrgeschichle 3: 3-8. 
1995 First records from Wellington Caves, New' 
South Wales, oi the extinct madtsoiid snake 
\^(mambi naracooriensis Sv(\\i\^. 1976. Proceed- 
ings of the Linnean Society of New South Wales 
115:233-238. 
1996. Studies tn the palaeontology and systematics 
of Australian snakes. PhD thesis, University of 
New South Wales. (Unpuhl.). 

SCANLON,J.D.&SHINE. R.l988.DcntilHmanddicH 
in snakes: adaptations to oophagy in the Austra- 
lian elapid genus Sbnoselapa. Journal of Zoology, 
London 216: 519-528. 

SHINE. R. t99L Australian Snakes: a natural history. 
(Reed: Sydnev). 

SMITH. MA. 1926 Monograph of the Sea-snakes 
(Hydrophiidae). (British Mu.seum (Natuial His- 
tory), London), 

.SMITH. M J, 1976, Small fossil vertebraie.s from Vic- 
lonaCave, N;aacoorte. South Australia. fV. Rep- 
tiles. Transactions o\ the Royal Society of South 
Australia UX): 39-51 

WORRELL, E. 1 96 1 . A new generic name for a nomi- 
nal species of Ocnisotna, Proceedings of Ihe 
Royal Zoological Society of New South Wales 
1958-1959-54-55 

WRIGHT, S. 1932. The ailes uf nmiaiion. inbnccdmg, 
cmssbrceding and selection in evolution. Procccd- 
■»iebolihe6ih Inicmalional Congress of Genetics: 
35"6-366. 



CAINOZOIC TURTLES FROM RIVERSLEIGH, NORTHWESTERN QUEENSLAND 

A- W. WHITE 



While, A.W. 1997 06 30: Cainozoic turtles from Riversleigh, norlhweslem Queensland. 
Memoirs of the Queensland Museum 41(2): 413-421 . Brisbane. ISSN 0079-8835. 

The Chelidae and Meiolaniidae occur in the Oligo-Mioccne ill Riversleigh with the former 
dominant and shorl-necked Elseya/Emydura accounting for over 90% of turtle material. 
Chelodina and Pseudemydura are present with several undcscribed forms. Chelid diversity 
increased from the Oligocene to the Miocene, while average size decreased, suggesting a 
change in the aquatic habitats. I propose that large late Oligocene river and overflow systems 
were replaced by smaller, slower-flowing waterways thai by the mid-Miocene had developed 
numerous small, stationary aquatic habitats, some occupied by dwarf turtles. 

The Miocene appearance and radiaiion of the large terrestrial meiolaniid turtles and their 
presumed thermal and dietary requirements suggest that Riversleigh's gallery forests were 
heterogeneous and punctuated by open clearings. Absence of trionychid turtles and dearth 
of long-necked chelids may be related to the general unsuitability of habitats at Riversleigh, 
particularly during the Oligocene. D Ausiralia, Riversleigh, turtles, palaeoecology. 

A. W. White. School of Biological Sciences, University of New South Wales, New South Wales 
2052: received 24 March 1997. 



Riversleigh's Oligocene and Miocene turtles 
(White, 1988, 1990, 1992; Gaffney et al., 1989; 
White & Archer, 1989) represent the highly 
aquatic side-necked Chelidae and the large, ter- 
restrial, horned Meiolaniidae. 

Chelid remains are common in Systems A, B 
and C of Archer el al. ( 1 989, 1 994) and are known 
from Pleistocene gravels (Terrace Site) adjacent 
to the Gregory River (While & Archer, 1994). 
Living specimens of the presumed exUnclElseya 
lavarackonan, have been collected from nearby 
Lawn Hill Creek and subfossil material from near 
Ihe Gregory River (Thomson et al., 1997). 
Meiolanid remains are conilned to System B and 
one site in System C (Gaffney el al., 1992). 

Riversleigh turtle material including skulls is 
well preserved (White & Archer, 1993) but no 
articulated remains have been found. Relation- 
ships of extant chelids have been based almost 
exclusively on cranial anatomy (Gaffney, 1977, 
1979) but biochemical techniques are establish- 
ing alternative phylogenies (Georges & Adams, 
1992) and derived shell features have been used 
to define fossil species (White & Archer, 1994). 

FAMILY CHELIDAE 

The extant chelids Pseudemydura, Emydura/ 
Elseya and Chelodina have been reported from 
the Oligocene-Miocene of Riversleigh (While, 
1988; Gaffney el al., 1989; White & Archer, 
1989). 



Celid taxonomy is based heavily of features of 
the skull (Gaffney. 1977, 1979). There are no 
derived skull characters known that can be used 
to distinguish Emydura Elseya despite these gen- 
era being electrophorelicaily distinct (Georges & 
Adams, 1992). Shell features are m^l normally 
used because they have been presumed lo be 
variable in chelid lineages (Gaffney, 1977) but 
some diagnostic shell features are now known 
(White & Archer, 1994; Thomson & Georges, 
1996). I adopt the more conservative position of 
using skull features lo distinguish shorl-necked 
chelids. Gaffney et al. (1989) reported on 
Emydura/Elseya skull material from Riversleigh 
although shells and shell pieces were available. 
Scute features are used to identify modern taxa 
but these have little or no phylogenetic value 
(Gaffney, 1979). 

Riversleigh Oligocene-Miocene turtles are 
dominated, both in abundance and diversity, by 
short-necked chelids of which Emydura/Elseya 
were most common (White & Archer, 1989), 
representing over 90% of identifiable material, 
and are known from 10 fossil sites. These two 
genera contain nearly 60% of extant chelid spe- 
cies (Georges & Adams, 1992). Two mandibular 
pieces from Riversleigh's Bob's Boulders (Sys- 
tem C) are the only remains assignable directly to 
Elseya. They have alveolar ridges typical of the 
Elseya dentata species complex. 

A single skull fragment and partial plastron of 
Pseudemydura is known from Ringtail Site (Sys- 
tem C; Gaffney el al., 1 989). This genus contains 



4l:V 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. I . Sku)l of Chelodinasp.. QMF3 l303.Qucnlin*s 
Quarry, Rivcrsieigh. A, dorsal. B. ventral. 

a single extant species, the Western Swamp Tor- 
toise (P. umhrina), an endangered species con- 
fined 10 iwo small swamps north of Perth 
(Cogger. 1992). The Riversleigh specimen is the 
first fossil record of the genus and indicates 
P senile my dura was once much more w idesprcad. 
Pseudemydura is the most derived extant chelid 
(GalTney. 1977, 1979) and is unusual among 
pleurodires in having little temporal or posterior 
eniargination of the skull. This results in a skull 
with a near complete roof above the cranium. The 
Riversleigh skull fragment consists of a mas- 
sively expanded supraoccipital indicating that en- 
closure of the hindmost portion o^ the skull 
occurred betore the Miocene. Similarity of the 
Riversleigh specimen to the modem species sug- 
gests that Pseudemydura diverged from other 
short-necked chclids early in the Australian radi- 
ation, a conclusion supported by cladistic analy- 
sis of modern chelid skull features (Gaffney. 
1977). Elcctrophoretic data (Georges & Adams, 
1992) did not include Pseudemydura and 
Gaffney 's (1977) hypothesis thai Pseudemydura 
is the sister-group lo all i)lher Australian chclids 
remains (but see Manning Sl Kofron, 1996). 

Long-necked chclids arose before the 
Oligocene (Manning & Kofron, 1996). Georges 
& Adam's (1992) indicate that Clielodmo is the 
sister group to all Australian short-necked turtles 



(excluding Pseudemydura). An almost complete 

plastron and some carapace bones of a small 
long-necked Chelodina are known (Gaffney ct 
al., 1 989). A species oi Chelodina fromQuentin's 
Quarry Site is described below. An almost entire 
carapace of a diminutive chelid from Melody's 
Maze Site (System C) and a partial skull from 
CMP Site (System C) White (.1992, 1993) arc 
described below. 

SYSTEMATICS 

Order TESTUDINES Linnaeus. I75H 

Infraorder PLEURODIRA Cope, 1K63 

Family CHLLID^VE Gray, 1825 

.Subfamily CHELINAE Gray, 1825 

Chelodina sp. 
(Fig. I) 

MATERIAL. QMF31303, dorsal skull elements con- 
sisting of a fused frontal, paired parictals and prootics 
and supnioccipiial bones from Quenitn^s Quarry Site* 
middle Miocene. System C. 

DESCRIPTION. Frontal fused, detached from 
parietals, with suture undamaged. Posterior tip of 
supraoccipital missing. Parietals joined, sutured 
to theirrespective prooiics and the supraoccipital. 
Frontal Hat, roughly triangular, with maximum 
width behind rim of the orbit near the suture for 
the postorbilal bones, with long and tapered an- 
terior aspect, with the suture for the prefrontal as 
well as the rim of the orbit. Suture with the 
parietals horizontal. Parietals wider than frontal 
at their suture, widest in the posterior section of 
the orbit near their sutures with the postorbital 
bones, with dorsal aspect drawn into a tapering 
sagittal crest terminating with the intru.sion ol the 
supraoccipital to fonn The very tip of the crest. 
Supraoccipital forming roof of the posterior cra- 
nium; parietals forming roof for the majority of 
the vault. Cranium widest near the parietal- pro- 
otic suture. Prooiics large, inilated, with suture 
attachments for the quadrate and basisphenoid 
intact, with latter suture uniting a broad foot-like 
plate of the quadrate with the basisphenoid. 

DISCUSSION. This specimen is referred to 

Chelodina on the basis of the fusion of the frontal 
hones, absence of temporal skull roofing and lack 
of contact between the parietals and squamosals 
(GalTney. 1977)- Goodc (1966) and Legler 
( 1985) divided Chelodina into: the C lan^irolUs 
group (A) containing C. longicollis, C. 
steindachneri and C novaeguinea and the C 



CAINOZOIC TURTLES FROM RIVERSLEIGH 



415 




FIG. 2. Carapace of small turtle, QMF31304, Melody's Maze, Riversleigh. A, dorsal. B, ventral. 



expansa group (B) containing C. expansa, C. 
rugosa, C. oblonga and C. parkeri. Goode (1966) 
separated short-necked species (Group A) where 
neck length is less than shell length, the posterior 
skull is not disproportionately extended, the 
basiocciptial is large and expanded anteriorly, 
while the squamosal is reduced and lacks a pro- 
truding lateral process. Group B long-necked tur- 
tles were longer-necked where neck length is 
often longer than shell length, the posterior skull 
is markedly elongated, the basioccipital is small 
and squarish, and the squamosal has a prominent 
lateral process. Electophoretic data (Georges & 
Adams, 1992) support these groupings but not 
their generic status. In particular, they disagree 
with the placement of C oblonga and its relation- 
ship to the C. longicoUis group. 

The Quentin's Quarry shell cannot be readily 
allocated to either group although the skull shows 
no marked posterior elongation. The skull is 
unique in: I. Contribution of the frontal bone to 
the orbit. In living Cheiodina frontals are fused, 
forming a flat, dorsal plate behind the orbit with 
a thin medial process extending to the nasal re- 



gion. The anterior process is intimately fused to 
the prefrontals which make up most of the dorsal 
orbit. The Quentin's Quarry skull has a thin me- 
dial process but the suture points with the prefron- 
tals are minimal and anterior. The frontal process 
bears the rim of the orbit behind the prefrontal 
sutures and makes up the majority of the dorsal 
orbit. This means that the eyes were relatively 
closer together and directed more upwards than 
sideways. 2. Expanded anterior parietals. In mod- 
ern long-necked turtles the anterior parietals are 
flat dorsally but curve steeply down to form the 
walls of the anterior cranium. In the Riversleigh 
specimen, there is a parietal shelf formed by the 
extension of the parietals. The postorbital bones 
are fused to this shelf which forms part of the roof 
of the post-orbital canal). 3. Triangular sagittal 
crest. In extant long-necked turtles the parietals 
are flattened dorsally before being drawn into a 
narrow, elongate mid-cranial crest that extends to 
the rear of the skull. This leaves a massive canal 
for the neck and jaw musculature. In the 
Quentin's Quarry skull, the dorsal parietals taper 
evenly to the back of the skull creating the most 



416 



MEMOIRS OF THE QUEENSLAND MUSEUM 



robust crest uf any known Chehdina. 4, Enlarged 
prooiic bones. In modem Chehdina the prooiics 
are fused laterally to the parietais. As such, they 
form a horizontal beam connecting the cranium 
to the external ear. In the Queniin"s Quarry spec- 
imen the prootics ai'e inflated producing a gradual 
sloping from ihe side of the skull down to the 
quadrate. The temporal canal is therefore triangu- 
lar in profile with the majority of ihe canal space 
being more lateral and closer to the quadrate. 5. 
Restricted posterior extension of the skull. All 
modern long-necked turtles have low, elongate 
skulls. This is achieved by the extension of ihe 
mid and hind skull regions in the horizontal plane. 
In pariicuUir. the supraoccipital, quadrates, squa- 
mosals and opisioiics are markedly elongated. 
The Qufntin's Quarry skull has an almost rectan- 
gular supraoccipital that would scarcely have ex- 
tended beyond the exiernal ear. 6. Shape of 
cranial vault. In modern Cheiodina Ihe cranium 
is widest near the fronlal-parielal suture. In the 
Riversleigh long-necked turtle, the vault is widest 
in the mid-parietal, close to the prootic contact 
/.one. 

Genus indel. A 
(Figs 2, 3) 

MATERtAl- QMF31304, an almost entire carapace 
and nght femur IVom Melody's Maze. Gag Plateau. 
Riversleigh. System C. middle Miocene ( Archer efaL, 
1989). 

DESCRIPTION. Carapace elongate, oval, 
lOOmni long, 75mm wide across pleui'als 4. Shell 
without keeling along the midline, lacking ser- 
rated posterior peripherals, relatively low and flat 
in profile, with a longitudinal vertebral depres- 
sion running the length of the shell, without fe- 
nestra, Pyga! without division or indentation. 
Pleurals 1 1. relatively small. Peripherals increas- 
mg in size towards the posterior, with peripherals 
8 and 9 the largest; peripherals 3, 4 and 5 and pait 
of 6 curved ventral ly, formmg rounded shell mar- 
gin associated with the bridge; anterior and pos- 
teriorio the bridge, peripherals flattened, forming 
lateral platforms around (he shell. On the inside 
of the carapace, rib heads on each of iJ-ic pleurals. 
Pelvic scars well-devetopcdon pleurals 8 and the 
suprapygal bones. Transverse ridge running 
across Ihe Hoor of pleural 1 from the raised rib 
head io the recess for the bridge Recess curved 
anteriorly, meeting the peripherals near the suture 
between peripherals 2 and 3. 




FIG. 3. Scute boundaries (dorsal) of QMF3 1304. turtle 
carapace from Melody's Maze. 

DISCUSSION. The carapace has almost all of the 
right side intact, lacking only the third peripheral 
and nuchal bones. The left side is damaged and 
lacks peripherals I to 7 and portions of pleurals 
L 2 and 3. 

The Melody's Ma/e carapace may be tenta- 
tively assigned lo Emydura on the curved bridge 
recess and raised transverse ridge running across 
the Hoor of the first pleural bone (White & 
Archer, 1 994). However, the pelvic scars on pleu- 
rals 8 and the suprapygals are more typical of 
Eheya. The shell is very small although it is from 
an adult animal. Its adult features include the 
closure of ihccarapacial fencstral, the light fusion 
of the peripherals to the pleurals and the absence 
of features such as keeling of the carapace and 
expansion and Ilanng of the peripherals. The 
shell shows signs of advanced age: the extended 
growth of the pleurals during adult life has cre- 
ated a vertebral groove running the length of the 
carapace. 

This shell cannui be placed m any of the known 
species of Emydnra because of its small adult size 
and the unusual formation of the mid-peripheral 
bones- Among extant Emydnra, the smallest 
known adults are in E. signata (Cogger, 1992) 
from the coastal rivers of northern NSW. Adult 
E. si^qnata from the MacLean River show similar 



CAINOZOIC TURTLES FROM RIVERSLEIGH 



417 




FIG. 4. Tunic skull, QMF3 1 305, CMP Site, Riversleigh. A, dorsal. B, veniral. C, right lateral. D, posterior. 



features to those of the Melody's Maze shell with 
carapace lengths of approximately 150mm. 

Curving of the mid-peripheral bones associated 
with the bridge occurs elsewhere only in Elseya 
denfam-Vikc turtles. In modem short-necked tur- 
tles, the peripherals are flattened and project lat- 
erally to form distinct ledges around the shell 
margin. The posterior peripherals are often flared 
to produce 'wings' that cover the hind limbs. 
There is virtually no Ilaring of the peripherals in 
the Melody's Maze shell; the peripherals are 
small compared to the pleurals (an adult growth 
trend in all chelids exaggerated in this species). 
Peripherals make up only ]4% of the dorsal sur- 
face of the mid-carapace whereas Emydura (e.g., 
E. krejfti) the peripherals account for 20%. 



The flattened shell is unlike most Emydura 
species which have domed shells. Only in 
Psettdemydura, Chelodina, Elseya latistenmm- 
like turtles and some Elseya denfata-iikG species 
are the shells ilattencd and the central crest lost. 

Genus indet. B 

(Fig. 4) 

MATERIAL. QMF3I305, a partial skull comprising 
the paired frontals, parieials, posi-orbitals, quadrates, 
right squamosals and supraoccipilal, with paired pter- 
ygoids, quadrates, prooiic bones, opislilic bones, ex- 
occipitals, basisphenoid and basioccipilal from CMP 
Site. Gag Plateau. Riversleigh. middle Miocene, Sys- 
tem C. 



418 



MEMOIRS OF THE QUEENSLAND MUSEUM 



DESCRIPTION. Medium-sized, with extensive 
dorsal roofing, lacking temporal and posterior 
emarginations that typify most chelids (Gaffney, 
1979a). Parictals expanded, forming the bulk of 
the dorsal roof; squamosals broad, forming the 
temporal bridge; supraoccipitals only contribut- 
ing to the mid-dorsal section of the skull roof, 
weakly expanded in the dorsal plane. 

Frontals paired, with small, anterior, media! 
projections dividinge the prefrontals, contribut- 
ing minimally to the hind orbit. Poslorbitals 
mainly in the dorsal plane, with a descending strut 
forming the posterior wall of the orbit. Sutures tor 
the jugal evident. The descending strut from the 
postorbital and ascending process from the 
pteryoid in broad contact. Alary process of the 
pterygoid reduced, may not descend vcntrally 
below the level of the palate. Postorbital canal 
correspondingly more obvious in lateral view. 

Dorsal skull sloping downwards anteriorly. 
Squamosal broad, forming a broad temporal arch 
between the quadrate and the parietals. Quadrate 
large, with a deep angular ventral base continuous 
with the articulation facet for the lower jaw. 
Quadrate with a deep posterior vertical groove. 

Floorof the skull widening markedly where the 
parietals sweep laterally to unite with the quad- 
rates. Articulation facet for the lower jaw well 
below the level of the palate, with a broad area 
formed by the lateral extension of the pterygoid 
and the quadrates. Pterygoids with a transverse 
suture with the palantines. Basisphenoid triangu- 
lar, with a broad sulural contact with the basioc- 
cipital. Occipital condyle below the level of the 
foramen magnum. 

DISCUSSION. The right side of the skull is rel- 
atively intact whereas the left parietals and squa- 
mosals are broken. The skull is heavily 
impregnated with a dark mineral. 

The CMP skull cannot be assigned to any 
known infraorder or genus. It is particularly un- 
usual in the structure of the skull roof. The only 
other chelid that lack posterior and temporal 
emarginations on this scale is Pseudemydura in 
which the roof is extensive, the posterior emargi- 
nation is replaced by a posterior dorsal extension 
of the supraoccipital, and the temporal emargina- 
tions are replaced by the extension of the squa- 
mosal. In the CMP skull, the supraoccipital forms 
relatively little of the skull roof and some poste- 
rior emargination is evident. The squamosal is 
expanded, as in Pseudemydura. As a conse- 
quence, the temporal roof is broad. 



In Pseudemydura y expansion of the supra- 
occipital causes the hindmost portion of the skull 
to be lower than the parietals. In the CMP skull, 
the parietals extend back to the posterior margins 
of the skull and overiie most of the supraoccipital. 
This results in the skull having an anterior slope 
with its highest point behind the level of the 
quadrates. 

The quadrates have an unusually deep ventral 
footing that forms a thick foundation for articula- 
tion with the lower jaw. The articulation facet is 
dislocated laterally and lies beneath the most 
lateral edge of the quadrate. These features, com- 
bined with the reduction of the alary process of 
the pterygoid indicate an unusual distribution of 
muscles between the upper and lower portions of 
the skull. The ventral portion of the cranium and 
the exoccipital regions are very similar to the 
arrangement found in all Chelininae. 

PALAEOECOLOGY 

Living Australian freshwater turtles have a va- 
riety of life history strategies and survival mech- 
anisms (Kennett et al.,1993; Grigg et al.,1986; 
Georges, 1982; Georges et al.,1986; Georges & 
Kennett, 1989; Heaphy,1990; Kennett & 
Georges, 1 990; Georges, 1 988; Thompson, ! 988). 

Several major trends which may have ecologi- 
cal significance are apparent in Oligocene- 
Miocene turtle assemblages at Riversleigh. Only 
2 families, the Chelidae and Meiolaniidae are 
present and of these, chelids make up about 98% 
of materia! recovered. Chelids dominate these 
fossil assemblages in the same way that they 
dominate modern Australian freshwater systems 
(Legler, 1985). Only in a few, far northern local- 
ities are non-chclid freshwater turtles present; 
Carettochelys insculpta, the Pig Nose Turtle (a 
carettochelid), is found in a few NT rivers 
(Georges & Kennett, 1989; Heaphy, 1990). The 
earliest Chelids known are from the Cretaceous 
of Patagonia (de Broin, 1994) and the family is 
thought to have evolved in southern Gondwana 
(South America and Australia. A sister-group, the 
pelomedusids, evolved at the same time in north- 
ern Gondwana and fossils occur in Africa, Mad- 
agascar and South America. 

Other turtles lived in Australia during the 
Oligocene-Mioccne including the giant horned 
meioianiids and soft-shelled trionychids 
(GalTney. 1979. 1981). 

Another feature of the Riversleigh fossil turtle 
fauna is the dominance of the plesiomorphic 
short-necked chelids, Emydura/Elseva turtles ac- 



CAINOZOJCTLfRTLES FROM RIVERSLEEGH 



419 



couni for over 85^- of turtle remains. Mudcm 
EmyduralEheya arc predominantly herbivorous 
and ocLur in touslal and inland rivers, creeks and 
!ap.oon.s cspcciMlIv those with a wcll-devclopcd 
.-Kiuaiic flora (Cann, 1978: Lcglcr, 1985). They 
arc not abundant in nnuldy or .stagnant water. 

Cheiids in System A arc typically large with 
shells up to 5(MJrntii U>ng and equivalent in size to 
the largest extant chclids. Chclid fossils from 
S>stcms B and C arc smaller (shell lengths 200- 
3CX)ninO and thinner-shelled. System C turtle is a 
dwarf with adull shell irK)mm lon^. Tlie large 
System A turtles occur at Site D (Archer et al., 
1994) which yields many broken sections ot un- 
usually ih>ck turtle shell. Carapacial plates 10- 
2()mm thick arc typical. Frotii the larger shell 
pieces I estimate shell lengths of 35()-450mm. 

The largest exlanl cheiids are the northern snap- 
ping turtle {Elseya dm^aui) and gulf snapping 
turtle (Elseya iavarackorum) which inhabit large 
flowing rivers or deep still water bodies (Cogger. 
1992). The only other fossil chclids from 
Riverslcigh that approach the dimensions of the 
Systems A turtles are late Pleistocene Terrace 
Site (While & Archer. 1994). 

Species diversiiy increases fromSysicms A to 
C with 1 chclid species in System A, 2 cheiids and 
2 nieiolanids in Systetri B and 6 chclids and 1 
nieiolanid in System C. The increase in diversity 
suggests an increase in small, shallow or slow- 
flowing aquatic habitats from late Oligocene to 
middle Miocene. 

Maximum vurtle MfA is a ustrful indicator of 
water depth and How rate. Riverine species that 
occurin deep or relatively fa>i- Ilowing water are 
typically large and capable of sustained swim- 
ming. Smallci species are excluded from such 
siies and often confined to fringe water bodies 
such as side-streams, overflows or ponds ( Pritch- 
ard. 1979). The range of small-shelled cheiids in 
System C sites suggests that a vaiicty of shallow 
water or slow moving habitats were available at 
the time. System C tunles typkally had shell 
lengths 150-250mm. 

The smallest extant turtle is the hog lurile, 
Clemmys mtthlfngergi, an emydid fPritchard, 
1979) which is 76-M4mm long. These tunics 
occur in extremely shallow, still water habitats; 
in some cases free water is not available. None of 
these sites are necessarily clear water sites 
(Behler&King, 1979). 

In the Gregory River at Riversleigh there are 5 
cheiids; these, in order of abundance, arc 
Emyttitni sp, ajf. subglobosa, Emydura sp. nff. 
ikumae^ Elseya latisternum, Elseya Iav- 



arackorum and Cheiodirta m^osa. The latter iwx> 
are uncommon. This level of diversity is reasun- 
ahly high for modem freshwater habitats in Aus* 
tralia. Species diversity increases as mean 
temperature aiki habitat variation increase {ObsU 
1986) 

Only Systems B and C contain large, teniesirial 
turtles (meiolaniids) with shell lengths up to I m 
long Using Mriokmia pUttyceps as u model, 
these creatures would have bad average bcxly 
masses of l50-2(.)Okg. 

There arc a number of large, land turtles aJivc 
today. The best known ate the varioits Gala(Ugo.s 
tortoises (Geofhelarte elphantopus) itnd the Al- 
dabran tortoise [Geocfielone ^iganrea). Both 
reacti txidy sizes and masses consideralily grc;Uer 
than that calculated for Riverslcigh \s Miocene 
meiolaniids. They arc regarded as examples of 
island endemism leading to gigantism (Pritchard. 
1979). The majority of the extant large terrestrial 
tunles inhabit hot savanna regions of the wx^ld 
For example, both species of large African land 
turtles {Gi'orlielone sulcata and 6', pardalts) 
occur in northern and central Africa. Neither >]>e- 
cies is found in dense forest. C. stdcara's disU^i- 
huiion is along the southern Sahara into central 
Africa (Priic'nard, 1979). Savannah habitat verg- 
ing onto treeless plains is the preferred habitat ior 
all African land lortoises. 

During U>c Mi<vcnc. Rivxrrslcigh was covered 
by wet iWc'sl communittes (Archer et uL, 1994) 
but the large land turtles seem incongruous m this 
habitat. I.aiul lunles such as the S)iuih Ainctican 
G. dfntkukva and C carbonariu are generalisi 
herbivores and live in a range of habitats, includ- 
ing open savanna, closed woodland and rainforest 
(BjorndalJ9S9), MosK()vits, l985;Moskovits& 
Kiester. 1987). While both spectes will venture 
deep into rainforest and feed on fungi, fallen fuiit 
and herbs, they do not permanently reside in the 
closed forest. Both prefer to reside near the for- 
est-savanna interface (Moskovits & Bjorndal, 
1990). Flowers and grasses fonn a major part of 
the dictof South American land turtles in savanna 
and swamp habitats. They consu!i»e sand which 
is thought lo a.s.sist digestion of coarse fibrous 
matter; they are hinil-gut fermcniers and neetl a 
certain amount of fibre in their diet. In iherainlor- 
cst, the turtles get fibrous foodstuffs in the form 
of vines and shoots, but these arc usually not 
plentiful. In addition, the tortoises seek out clear- 
ings (e.g. sites of recent tree falls) to bask. Hind- 
gut digestion requires fairly high ami constant 
temperatures (of 30"^ or more) and the lurties need 
Ko have periodic access lo dircci sunjjghi. Youn 



MO 



MEMOIRS OFTWB QUEENSLAND MUSEUM 



gcr and smaller lurtlcii necti access lo sunhglii on 
a more regular basis than large adults (MoskoviLs. 
1985). 

Ifihcsclimitaiions for giani land tonoiscs apply 
also lo meiolannd turtles living in or near forested 
habitats, then ihcre musi have been clearings 
where the unimals could get sufficieni voluitte of 
high fibre food and have direct access to sunlight. 
The rotund body form of these lanti turtles means 
that large anurmis would have a hi^h ihermid 
inertia. At air icmpcraturcs abtuc ?{)^ they may 
not need to seek additional evlenial hcai sources. 
However, should their body temperature lall 
below this level they would need to reach ex- 
posed sites quickly lo restore core lernperalures 
(Swingland & Fazier, 1979). 

Riveislcigh'sineiolaniidsarclow in abundance 
despite their high diversity; 4 species in 2 genera 
arc known from 5 individuals. Chelids. in eon 
Irasl. itre represented by hundreds of specmiens. 
Nothing is known abuui dieiary itquirements t»i 
horned turtles. L^rgc exiant terrestrial turtles are 
opportunistic lictbivores and have rcquifemcnis 
lor librous matter intake (Pntchard. 1976). The 
unusual structure ol the jaws ofmeiulaniids indi- 
catcs that son^c dietary selectivity nuiy have been 
possible, but the nature of their diel is unknown. 

The absence of soft-shelled uionychids from 
Riversleigh's Oligocene Miocene coupled with 
the very low occurrence of long-necked turtles 
may be ecologically significant. Bcnh types ol" 
turtles are *ambush predators* (Pritchard, 1976). 
Both groups flourish in situuiions where iliey 
cannot be easily seen and ihis usually means 
turbid, muddy or dark water. Their absence and 
the preponderance of EmyduratElseya suggests 
relatively clear water aquatic environments. 
Chekniina in System C coincides with ihe reduc- 
tion in average chelid shell size, indicative of 
shallow, turbid lagoons during this period of Ihe 
Mit-Kcne. The complete absence of trionychids 
suggests ihai ihc lagoons did noi have sihy bot- 
toms and were unsuitable for concealment. 

ACKNOWLtDGEMENTS 

The Rivcfsleigh project is supported by the 
AuNlralian Kesearch Council, the LX-parlinent of 
the Environment, Spori and Territories, National 
Estate Programme Grants (Queensland). Queens- 
land National Parks and Wildlife Service, the 
Australian Geographic Society, the Waanyi pe<v 
plc and Carpentaria Land Council, the Linnean 
Society of New South Wales, ICI. the Queens- 
land Museum, the University of New South 



Wales and the RiverMeigh Society. I thank Anna 
Gillespie who prepared the specimens and took 
the photographs. 

LITERATURE CITED 

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ARCHER. M.. HAND. S.J.. & GODTHELP. H. 1994. 
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BEHLER, J.U, & KING, F W. 1979. The Audihtm 
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BJORNOAL, K.A. 1989. Flexibility of digeslive rc- 
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de HROIN, F. 1994. Examples ot parallel evolution in 
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GAFFNEY, E.S. 1977- The side-necked tunic Family 
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1979.1. Comparative cranial morphology ul Kceeta 
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1981 A review of the fossil turtles of Au.stralla. 
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1991. The fossil turtles of Australia. Pp 701-720. Irt 
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GAFFNEY. E.S.. ARCHER, M & WHITE, A.W. 
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1992. Warkalartia^nDCV^ meiolaniid turtle iVoni the 
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GEORGES. A. 1982. Diet ot the Australian Freshwater 
iiirlle Emydum kreffti (Chelonia.Chelidae^, \u an 
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3.^1-336. 
1988. Sex deiemiinanon is independent of incuba- 
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GEORGES. A & KENNETT. R. 1989. Dry season 
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GEORGES,A.,NORRIS.R,H &WENSING.L. 1986. 
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GEORGES. A. & ADAMS. M. 1992. A phylogeny for 
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GOODE. J. 1976. Freshwater tortoises of Australia and 
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•GR1GG,G.C . JOHANSSEN.K.. HARLOW, P.. 
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HEAPHV. L. 1990. The ecology of the Pig-Nosed 
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KENNETT, R'& GEORGES, A. !99(K Habitat utilisa- 
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THOMPSON, M.B. 1988 Population ul the Mumty 
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NEW CROCODILIANS FROM THE LATE OLIGOCENE WHITE HUNTER SITE, 
RIVERSLEIGH, NORTHWESTERN QUEENSLAND 

PAUL MA. WILLIS 

Willis, Paul M.A. 1997 06 30: New crocodilians from the late Oligocenc White Hunter Site, 
Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41(2); 423-438. 
Brisbane. ISSN 0079-8835. 

Four new species of crocodilian are identified from the late Oligocene White Hunter Site, 
Riversleigh, northwestern Queensland, one of which is also found in other System A sites 
at Riversleigh. All four species are assigned to known genera and some revision of two 
generic diagnoses is required. Two different forms of posterior cranium are also identified 
from While Hunter Site and retained in open nomenclature. Palaeoecological significance 
of four crocodilians in a single site are interpreted as a sympatric assemblage because they 
have different head shapes. However, the diversity in these crocodilians could also suggest 
a thanatocenosis involving taxa from different hydrodynamic regimes with differing degrees 
of forest canopy cover. □ Riversleigh, Bam. Qumkana, Mekosuchus, Oligocene. 

Paul Michael Arthur Willis, Quinkana Pty Lid, 3 Wanda Cres., Berowra Hts, NSW, 2082; 
received 4 November J 996. 



The fossil assemblage from White Hunter Site 
at Riversleigh, NW Queensland contains skull 
fragments and postcranial material of crocodil- 
ians and other vertebrates. The fragments repre- 
sent at least 4 crocodilian species. Three different 
maxillae are assigned to new species of known 
genera. Of 4 mandibles identified 3 are assigned 
to 3 of the species identified by maxillae; the 
fourth belongs to a species better known from 
Riversleigh's D, Sticky Beak and Pancake Sites 
(Willis ctal., 1990). Cranial material is described 
but not assigned to any of the 4 new species. 

Mekosuchus Balouel & Buffetaut, 1987 and 
Bant Willis et al., 1990 were previously mono- 
typic and their generic diagnoses require revision 
in the light of new species assigned below. Quink- 
ana was revised by Willis & Mackness (1996) 
and their expanded generic diagnosis (based on 
Molnar, 198 1) encompasses the new species de- 
scribed here. 

Mekosuchus was known from Recent cave de- 
posits in New Caledonia. It has a unique au- 
tapomorphy; the maxilla participating in the 
orbit. M. whitehunterensis sp. nov. is the first 
pre-Pleistocene record of this genus. 

Quinkana has 3 species: Q. fortirostrum Mol- 
nar, 1981 from E Queensland; Q. timara Megir- 
ian, 1994 is more slender-snouted from late 
middle Miocene of Bullock Creek, NT; and Q. 
babarra Willis & Mackness, 1996 is from early 
Pliocene at Allingham Creek, Queensland. 
Quinkana is distinguished by a suite of ziphodont 
features and is unique among mekosuchines 
{sensu Willis el al.. 1993) in being a broad- 
snouted ziphodont, Quinkana meboldisp. nov. is 



the third pre-PIiocene record after Q. timara and 
a species from the late Miocene Ongeva Local 
Fauna, Alcoota, NT (Murray & Megirian, 1992; 
Murray etal., 1993; Megirian, 1993). 

Baru darrowi (Willis et al., 1990) was de- 
scribed from middle Miocene of Bullock Creek, 
NT and Site D, Riversleigh. Two species of Baru 
(Willis el al., 1990) are recognised from While 
Hunter Site. One species is particularly small for 
the genus and the other species is based on mate- 
rial from a number of Riversleigh' s System A 
sites (sensu Archer et al., 1 989), including While 
Hunter Site. Some of the material assigned here 
to the second species of Baru was previously 
assigned to B. darrowi. Baru species arc broad, 
moderately deep-snouled mekosuchines with 
moderately compressed teeth and a distinctive 
ridge on the posterior of the maxilla and the jugal. 

Mekosuchus, Quinkana and Baru can all be 
shown to be mekosuchines. A more detailed phy- 
logenetic analysis of these three genera forms 
part of a more comprehensive investigation of the 
phylogeny of mekosuchines (Salisbury & Willis, 
1996). 

The ecological implications of four crocodil- 
ians in the same deposit invites an investigation 
of the possible structure of crocodilian faunas. 
There would appear to be no ecological conflict 
between the sympatric existence of all four spe- 
cies because their different morphologies suggest 
the exploitation of different habitats. This is con- 
sistent with modern analogies such as some parts 
of the Amazon River Basin and with other fossil 
deposits such as Messel and Gcisalial in Ger- 
many, the Bridger Basin in the U.S.A. and the La 



424 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG- I - Mekoutchus whitrhunjetvnsis n.sp-, QMF3 1 05 1 , 
hololypc. fight maxilla, ventral view. Scale = 5mm. 

Venlu fauna in Colombia. Alternatively, the dif- 
(erenl crocodilians in White Hunter Site may be 
from ditTerent habitats atid have been collected 
together in a Ihanatocenosis. 

This publication was the content of a seminar 
presented at the Conference on Australia Verte- 
brate Evt)lulion. Palaeoniulogy and Evolution 
(CAVEPS) in Alice Springs, March, 1991 and 
published as an abstract (Willis, 1992). 

Mekosuchus Balouet & Buffetaut, 1987 

TYPH SPFCIES. Mekosuchiis inexpectalns Balouet & 
Burfelaul, 1987. 

DIAGNOSIS (translated from French). Eu- 
suchians with choanae relatively little displaced 
posleriad; wings of pterygoids strongly devel- 
oped posteriorly: skull deck very broad; maNiila 
participating in lower border of the orbit; external 
narcs opening to the side and the front (an- 
terolateral ly); nasals not reaching external nares; 
palatines very narrow in their posieritir part; 
quadratojugal lacking a spine; snout short and 




FIG. 2. MtkosHchus \\hi(ehim!crensis, QMF3105L 
hoiolypc. right maNiila. Dorsal view with while arrow 
showing portion of the maxilla that participates in the 
orbit. Scale = 2mm. 



FIG 3. Mekosuchus whitehuntercnsi.s, QMF31052, 
partial frontal, dorsal view. Scale =5mm. 

deep; splenial does not participate in the mandib- 
ular .symphysis; posterior crushing teeth; 13 man- 
dibular teeth; lower teeth occlude medial to tipper 
series; vertebrae procoelous with strong neural 
spines m the cervical region; limb bones showing 
strong muscle insertions; presence of dorsal 
scutes. 

My diagnosis includes the following features 
(apomorphies indicated by 'a') are: 1, (a) maxilla 
participating in lower border ofthe orbit. 2. snoul 
short and deep. 3, (a) no conspicuous gap between 
the sixth and seventh maxillary alveoli. 4, high, 
narrow alveolar process. 5, symphyseal region 
very shallow dorsovenlrally. 6, splenial anteriad 
lo the level of the seventh deniary alveolus. 7, 
external mandibular feneslrae strongly reduced. 
8, (a) out-iumed llange on the angular and sur- 
angular. 

The type species diagnosis is: palatal fenestrae 
reaching antenorly to the level ofthe sixth tnax- 
illary alveoli; posterior teeth of rounded, crashing 
form; symphysis reaching posteriorly to the level 
of the seventh dcntary alveoli. 

Some features of the original diagnosis are 
synapomorphies of wider groups and others are 
of uncertain value so they are not employed 
herein. 

The character 'nasals not reaching external 
nares* is equivocal on available material so is not 
employed pending more complete materiul, 

Mekosuchus whitehunterensis sp. nov. 
(Figs 1-5) 

MATERIAL. Holotype. QMF31051. right maxilla 
(Figs I. 2). Paraivpe.s QMF31()52. partial frontal; 
QMF31053, almosi complete mandible; QMF3I()54 
and QMF3I055, anterior portions ot dcntaries. All 
from late Oligoccne White Hunter Site, Riverslcigh. 



NEW CROCODILES FROM THE LATE OLIGOCENE 



425 




FIG. 4. Mekosuchus whitehunterensis, QMF31053, left mandible, lateral view. Scale = Icm. 



DIAGNOSIS. Longitudinal sulcus below the orbit; 
palatal fenestrae reaching anteriorly to the level of the 
seventh maxillary alveoli; posterior teeth compressed 
and blade-like; and symphysis extending posteriorly to 
the level of the sixth dentary alveoli. 

ETYMOLOGY. From White Hunter Site. 

DESCRIPTION. Maxilla broad, deep-snouted, 
with moderately high, narrow alveolar process 
(sensu Molnar, 1981). Lateral wall steeply in- 



clined to the palate, with longitudinal sulcus ven- 
tral to the orbit. Small portion of the maxilla 
participating in the orbit, separating lacrimal 
from jugal. (Full extent to which the maxilla 
participated in the orbit cannot be deduced be- 
cause the posterior portion is missing in this spec- 
imen.). Alveoli ovale, slightly compressed 
laterally, close to each other so excluding the 
lower series from resting between them; fifth 
alveolus largest; first and seventh alveoli small- 




FIG. 5. Mekosuchus whitehunlerensis, QMF31053, left mandible, dorsal view. Scale = 1cm. 



426 



MEMOIRS OF THE QUEENSLAND MbSEUM 




FIG. 6. Quinkana meboldi n. sp., QMF3 1056. holotype. left maxilla, lateral view. Scale = 1 cm. 



est, almost equal in size. Only two pits for recep- 
tion of dentary teeth medial to the upper alveoli, 
between the sixth and seventh alveoli, and a dis- 
proportionately large pit posterior and media! to 
the seventh alveolus. PaJatal fenestra reaching 
level of the seventh alveolus. 

Frontal. Closely resembles frontals of M. in- 
expectatuSy very wide between the orbits; orbit 
margins raised, giving a concave transverse sec- 
tion to the dorsal surface, crania crisiae frontalis 
shallow, close together leaving a wide, thin shelf 
between them and the orbit margins. 

Mandible and dentary fragments. Pseudoheierod- 
ont with an undulating tooth row; tooth row 
shorter with respect to the whole mandible than 
in other crocodiles. Posteriorly, dentary strongly 
compressed laterally and deep dorsoventrally. 
Dentary thick around the base of each alveolus; 
buttressing variable in proportion to size of alve- 
olus, not strongly developed. Slight ridge defin- 
ing a groove lateral to the 1 2th through lo the 1 6th 
alveoli probably received posterior maxillary 



leelh. Spienial extending anteriorly lo 7th alveo- 
lus; symphysis extending to 6lh, Low ridge on 
the external surface of botli deniaries running 
from below the 8th dentary alveolus onto the 
angular, probably strengthened the dentary. 
Teeth 16, with unserratcd anterior and pcistcrior 
carinae, becoming laterally compressed posteri- 
orly so that posterior teeth are blade-like. 

QMF3 1 053 with large, out-turned fiangc on the 
posterior and ventral margins of angular and out- 
turned flange on the dorsal margin of the external 
surface of the surangular; flanges joining at the 
posterior margin of the mandible, marking 
boundary between the sculptured surfaces and the 
smoother surfaces for muscle attachment. 

Articular broad, expanded medially; articular 

portion of retroarticular process shallowly con- 
cave. Retroarticular process short and steeply 
inclined. Medial side of condylar surlace re- 
duced, strongly buttressed ventrally. External 
mandibular fenestra reduced, almost closed. 
Sculpture of indistinct scarring on the external 




FiG.l, Quinkana meboldi n. sp., QMF3 1056, holotype. left maxilla, ventral view. Scale = 1cm. 



NEW CROCODILES FROM THE LATE OLIGOCENE 



427 




FIG. S. Left deniaiy of Quinkana meboldi (QMF3\059). Laierai view, scale = 1cm. 



surfaces of the dentary and a well-developed 
mosaic of pits on the angular and surangular. 

DISCUSSION. This mandible is very derived 
and shows distinct similarities to A/. mexpecrafM^. 
Both are distinguished by: external fenestra re- 
duced or closed; anterior edge of surangular 
forming distinct step dorsal to the dentary; angu- 
lar and surangular flange; posterior portion pro- 
portionally short and deep; symphysis very 
shallow; similar sculpture. Maxillae of M. in- 
expectatiis and M. whitehimterensis exhibit the 
apomorphic condition of contacting the orbit. 
Thus, it is most parsimonious to associate the 
derived mandible and maxilla from White Hunter 
Site, both of which most closely resemble M. 
inexpectatiis. The association of mandibles and 
maxillae of M. inexpectatus is not in doubt 
(Balouet pers. comm.). 

Quinkana Molnar, 1981 

TYPESPECIES. Quinkanaforurosimm Molnar, 1981 

DIAGNOSIS. See Willis & Mackness (1996). 



Quinkana meboldi sp. nov. 
(Figs 6-9) 

MATERIAL. HolotypeQMF31056, left maxilla(Figs 
6, 7). Paraiypes QMF31057, almost complete left 
maxilla; QMF3I058, right maxillary fragment; 
QMF3 1 059, dentary fragment. All from late Oligocene 
White Hunter Site, Riversleigh. 

DIAGNOSIS. Small to moderate-sized, with 14 max- 
illary alveoli; palatal fenestra extending anteriorly to 
the level of the 8th maxillary alveolus: teeth partially 
interlock; snout narrower than in Q.fortirostrum; mild 
festooning; carinae of teeth without serrations. 



ETYMOLOGY. For Ulrich Mebold, 
Institiit fiir Radioastronomie. 



Max Plank 



DESCRIPTION. Maxilla. Teeth 14, compressed 
and blade-like with anterior and posterior carinae. 
Alveoli compressed to varying degrees; anterior 
6 teeth directed slightly posteriorly. 

Alveolar ridge low, mildly undulating, uninter- 
rupted laterally but medial side interrupted by pits 
for the reception of dentary teeth. Palatal fenestra 
extending anteriorly to the 8th alveolus. Midline 
palatal suture straight to the level of the 7th 
alveolus, then diverting laterally to accommodate 
a short, pointed anterior palatal process. 




FIG. 9. Quinkana meboldi, QMF31059, left dentary, dorsal view. Scale = 1cm. 



428 



MEMOIRS OF THE QUEENSLAND MUSEUM 




compressed alveoli and does 
not have a laieraily cornpresscd 
mandibular body. 

Baru Willis, Murray & 
Megirian, 1990 



TYPE SPECIES. 

Willis ctal.. 1990. 



Baru dartayvi 



i 


I '" 




""V" 
e 


'■'"1"""" !'■■"■ 1 



DIAGNOSIS. Broad, moder- 
ately deep snout; reduction of 
the second prcmaxillary tooth 
during growth sometimes re- 
sulting in four premaxiiiary 
teeth in adults; premaxiiiary 
and anterior six maxillary teeth 
directed posteriorly; toolh 
crowns mod e ra te I y c o m- 
prcsscd laterally; looth crown 
and socket dimensions highly 
ditlerentialed along both upper 
and lower tooth row s with cor- 
respondingly wide, deep alve- 
olar processes; conspicuous 
maxillary reception pits corre- 
sponding to dcnlary tooth 
crowns situated medial to the 
upper toolh row; anterior mar- 
gins of the palatal feneslrae ex- 
tending to the level of the 
seventh maxillary tooth; anle- 
FIG. 10. B^rxWrnii^n, QMF3 1060, holotype, snout, dorsal view, with line rior palatine process absent; 
interpretation. F^fronial; J=jiigal: L=lncrimal: Mx=maxilla; N=nasal; splenial lerminatcs anteriorly 




Pf=pre frontal; Pmx=premaxilla. Scale in cm. 

Dorsal surface steep-sided, indicating a deep, 
moderately broad snout. Preorbital or lacrimal 
ridge giving the snout a u*apezoidal cross section 
anterior to the orbits. Margins contacting nasals 
straight. Sculpture of distinct pits anteriorly, de- 
generating to pitted scars posteriorly. 
Alveoli with a sharply defined groove running 
medial to the alveoli. This appears to have been 
derived from the line ot foramina normally found 
in other crocodilianii in a homologous position. 

DISCUSSION. The lateral compression of both 
the dentary and the dentition as well as the lack 
of festooning indicates that the dentary fragment 
(QMF31059) belongs to Q. meboldl The single 
looth is identical to the posterior teeth of 
QMF31056. This dentary fomi differs from M. 
whitehunterensis in which the posterior-most al- 
veoli are interconnected. It also differs from 
dentarics atu^ibuted to Baru which lacks strongly 



at the level of the seventh den- 
tary tooth and does not enter 
symphysis; external nares terminal and broadly 
'apple'-shaped; distinctive bony crest arches pos- 
teriorly from the maxillae and jugals. extending 
to the quadrutojugais, 

REMARKS. The 2 new species are most closely 
related to Miocene B. darrowi from the NT (Wil- 
lis et al., 1990). Some of the material attributed 
here to B. wickeni sp. nov. was previously re- 
ferred to B. darrowi. ^Internal nares with raised 
rim' was included in the original diagnosis of 
Baru but its status is now uncertain. 

Baru darrowi Willis, Murray Sc Megirian, 
1990 



DIAGNOSIS. Snout broad, deep; rounded pre- 
maxillae; 13 maxillary teeth; nasals excluded 
from external nares; anterior termination of nasal 



NEW CROCODILES FROM THE LATE OLIGOCENE 



429 




S tlHllV.' tv* 




lary fragmeni; QMF3I066, maxil- 
lary fragment; QNTF3)067. den- 
tary; QMF31068, demary; and 
QMF31069, pairof dcm;mes with 
splenial fragments. All from late 
Oligocene While Hunier Site, 
Riversleigh. 

DIAGNOSIS. Snoul broad, not as 
deep as in other species: rounded 
premaxillac; 14 maxillary leeth; 
nasals contact external nares; lat- 
eral border of the na:sals without 
angulation at the maxilla-premax- 
ilia boundary: non-serrated cari- 
nae; mandibular symphysis 
extending posteriorly to the 5th 
dentary teeth. 

ETYMOLOGY. For Professor 
Huber, Rektor of (he Fricdrich 
Wilhelms Universitat, Bonn 



FIG. 1 1. Bam huben. QMF31060. holot>pe. snout, ventral view, with line 
interpretation. Mx=maxilla; Pa=:palaline; Pmx=premaxilla. Scale in cm. 



is a short, broad wedge; serrated carinae; mandib- 
ular symphysis extends posteriorly to between 
the sixth and seventh dentary teelh. 

Baru huberi sp. nov. 

(Figs 10, ID 

HOLOTYPE. QMF3 1 060, fragmentary snout (Figs 1 0, 
1 1 ). Paralypes QMF3 1 061 , right premaxilla and an- 
terior portion of right maxilla: QMF3I062, relatively 
complete premaxilla; QMF31063, partial maxilla; 
QMF3i064, maxillary fragment; QMF3I065. maxil- 



DESCRIPTION. Skull anterior 
to orbit. Snout low. broad; pre- 
maxillary alveoli circular, 4lh 
largest, 5 in juvenile, with 2nd 
alveolus reduced and almost 
lost, 4 in adult (2nd lost). Inci- 
sive foramen broad, lear- 
shaped, external narcs 
apple-shaped, with a short an- 
terior process of the premaxilla 
and nasals on its posterior mar- 
gin. Deep reception pit for the 
first dentary tooth not reaching 
dorsal surface. Fourth dentary 
looth reception notch promi- 
nent, with a secondary pit me- 
dial to it on the palate. 
Premaxillary-maxillary suture 
relatively straight, with a slight 
posterior convexity. Maxillary 
alveoli 14, arranged in a typi- 
cally crocodylinc enlargement 
sequence with the 5ih largest, 
laterally compressed particu- 
larly the poslerior-most 5. Low alveolar process 
on the anterior 6 maxillary alveoli. Dentary tooth 
reception pits 5, well-developed, medial to The 
upper series, between 6th- 1 1 Ih alveoli. Anterior 
teeth moderately robust, ovate in cross section. 
Posterior teetli witli low, rounded crowns. All 
teelh with distinct anterior and posterior carinae. 
Palatal fenestra extending anteriorly to between 
the 7th and 8th maxillary leeth; straight suture 
with the palatine forming a short palatal process 
reaching anteriorly to 6lh alveolus. Broad shelf 



430 



MEMOIRS OF THE QUEENSLAND MUSEUM 





FIG. 12. Baru wickeni, QMF 16822, holotype, dorsal view, snout, with line interpretation. F=frontal; J=jugal; 
L=lacrimal; Mx=maxilla; N=nasal; Pf=pre frontal; Pmx=premaxilla. Scale in cm. 



between palatal fenestra and posterior alveoli, 
rounded dorsally into the palatal fenestra and on 
to the internal surfaces of the maxilla. Sharp- 
crested ridge on the external surface of the max- 
illa at the line of the palate, starting above 10th 
alveolus, running off the posterior border of the 
maxilla. 



Nasals broad, contacting the external nares, 
gradually widening to the lacrimal-maxilla-nasa! 
triple junction, then tapering more sharply. Short, 
pointed anterior process of the frontals dividing 
the posterior extremities of the nasals.Lacrimals 
with low, rounded canthi rostrales, about twice 
the size of the prefrontals. 



NEW CROCODILES FROM THE LATE OLIGOCENE 



43 




■ \: T I t i I r 




FIG. 13. Baru wickeni, QMF16822, holotype, snout, ventral view, with line interpretation. Ect=ectopterygoid; 
Mx=maxilla; Pa=palatine; Pmx=premaxi!la. Scale in cm. 



Dentary fragments. Dentary pseudoheterodont, 
with an undulating tooth row. Symphyseal region 
deeper and larger than in Mekosuchus. Dentary 
built up around the base of each alveolus, with 
this buttressing variable in proportion to size of 
the alveolus and not strongly developed. Slightly 
raised area on the dorsal surface medial to the 4th 
and 5th alveoli. Dorsal margin of dentary be- 



tween and lateral to the 2nd and 3rd alveoli and 
between and lateral to the 7th, 8th and 9th alveoli 
with indentations for reception of teeth in the 
upper series, indicating that the upper series oc- 
cluded lateral to the lower series. Splenial extend- 
ing anteriorly to the 7th alveolus; symphysis 
extending to 5th alveolus. Sculpture of indistinct 



43: 



MEMOIRS OF THE QUEENSLAND MUSEUM 





FIG. (4 Bam wickeni , OMF16822, hololype, partial left maxilla and premaxilla, lateral view. Scale = 2cm. 



scarring on the ventral surfaces of the dentary 
merging to well developed pilling dorsally. 

DISCUSSION. This mandible is assigned lo B. 
huheri because ii is the only unassigned mandible 
of appropriate proportions and size range from 
While Hunter Site. QMF3 1 068 is an almosl cxacl 
fit for QMF3 1060. The other 3 mandibular forms 
from White Hunter Site can be shown lo belong 
to other taxa. 

Baru wickeni sp. nov. 
(Figs 12-17) 

MATERIAL. Hololype. QMF16822 (Figs 12-14) as- 
sociated posterior cervical and lumbar vertebrae and a 
calcaneum. Paratypes QMF3 1 070. anterior ponions of 
mandibles; NTM PS738-I, poslcrior nghi skull frag- 
ment and associated right anterior dentary fragment, 
NTM P86S 1-14, left mandible lacking the anicular and 
adjacent angular and surangular posterior lo the lateral 
tbranien and a small portion of the dentary at the level 
ofthe third luolh; NTM P8738- 1 , nghl jugal, pterygoid. 
eclopterygoid and posterior maxilla and an associated 
dentary fragment: QMF16823. jucal fragment; 
QMF16824, prcmaxillary fragments: QMF16825. 
right dentar>'; QMF16826, right dentary. All from 
Oligoccnc (System A) Site D. Riverftleigh. 
QMF31071 and QMF3I072, posterior portions of 
large mandibles and QMF3 1 073, anienor dentary frag- 
ment. All from late Oligocene White Humer Site. 
Riversleigh. 

SAMP27866» right premaxilla from late Oligocene 
Panciikc Site, Riversleigh. 

QMF31074, right maxillary fragment and fragment of 
skull roof from late Oligocene Sticky Beak Sue. 
Riversleigh. 



ETYMOLOGY. For Tony Wicken, University of 
NSW, for supporting the Riversleigh Research Proj- 
ect. 

DIAGNOSIS. Snout narrower than B. huberi or 
B. darrowi; deep; anteriorly pointed premaxillae; 
13 maxillary tecih; nasals entering external niires; 
anterior termination of nasals long and thin, 
strongly constricted hy premaxillae; non-serrated 
carinae; mandibular symphysis extending poste- 
riorly to 6lh or 7th dentary teeth. 

DESCRIPTION. Skull material. Snout narrower 
than B. darrowi or B. huberi, similar in depth to 
B. darrowi Premaxilla poinicd anteriorly rather 
dian rounded as in the other 2 species. Teeth 
similar to B. darrowi, lacking serrated carinae. 
Premaxillary alveoli in adults 4. in juveniles 5, 
with 2nd alveolus lost during growth. Maxillae 
with short posterior process medially invading 
lacrimal (not present in B. darrowi and unknown 
in B. huberi). Maxillary alveoli 1 3; palatal fenes- 
U"a extending anteriorly to the level of 7Lh maxil- 
lary alveolus. Nasals entering premaxillae unlike 
B. darrowi but similarly to B. huberi. Anterior 
nasals distinctive in B. wickeni. being thin slivers 
strongly constricted between the premaxillae. 
Lacrimal with distinct, rounded canihus ro^iralis, 
extending for a short distance onto the maxilla, 
Jugal with well-defined, arched ridge on the ex- 
terior surface. 

Mandibles. Alveoli 1 5, subcircular except for the 
4 slightly laterally compressed most posterior. 
Alveoli 3rd-6th on an alveolar process most 
strongly developed around the 4th alveolus. No 



NEW CROCODILES FROM THE LATE OLIGOCENE 



433 




FIG. 1 5 . Baru wickeni, NTM P8738- 1 , portions of the right side of the skull with associated dentary fragment, 
lateral view. Scale = 2cm. 




FIG. 16. Bant wickeni, NTM P8738-1. dentary fragment, dorsal view. Scale = 2cm. 




FIG- 1 1- Baru wickeni. QMF3 1 070, denlary, dorsal view. Scale = 2cm. 

reception pits for teclh from the upper series but phvsis exlendinu posteriorly to 6th alveolus; 
spacings and a lateral sulcus between 2nd and 3rd ^P^''^"'^'' reaching^anteriorly to 7lh. 



alveoli, 7lh and 8lh alveoli and between the 8th 



External mandibular fenestrae ovate, of moder- 
ate size and inclined posteriorly. Surangular nar- 



and 9th alveoli. Symphyseal region narrow. Sym- row dorsoventrally, inclined posteriorly with 



434 



MEMOIRS OF THE QUEENSLAND MUSEUM 





FIG. 18. Dorsal views of reconstructed snouts of Ban4 
hiiberi (lop), B. wickeni (middle) and B. darrowi 
(bouom) showing differences in sutural relations, par- 
ticularly in the nasal-premaxillae suiures, and general 
proportions. Bam huberi based on QMF3 1 060 (holo- 
type ), B, wickeni based on QMFl 6822 (holotype) and 
B. Harrow/ based on NTM P8695-8 (holotype). Scale 
= 5cnn. 

dorsal margin not parallel to the denlary. Angular 
slender, inclined. Smooth region for attachment 
of theposieriorpterygoideus musculature sharply 
demarcated from the heavily sculptured area.s of 
the angular and surangular by a low ridge. Artic- 
ular and retroariicular process short, broad and 
steeply inclined. 




FIG. 19. Ventral views of reconsimcied snouts oi Baru 
huberi (top), B. wickeni (middle) and B. darrowi 
(bottom) showing differences in general proportions. 
Baru huberi based on QMF31060 (holotype), B. 
wickeni based on QMF16822 (holotype) and B. 
darrowi based on NTMP8695-8 (holotype). Scale = 
5cm. 

DISCUSSION. This new species is based primar- 
ily on material from Site D. 

In describing B. darrowi, Willis et aj. (1990) 
recognised thai specimens from Bullock Creek 
differed from specimens from Rivcrsleigh. How- 
ever, at that stage there was insufficient material 
to separate 2 species. Since then a large portion 
of a snout from Riverslcigh (part of QMFl 6822) 
a fragment of which was in the original descrip- 
tion of B. darrowi has been rediscovered and 



NEW CROCODILES FROM THE LATE OLIGOCENE 



435 



prepared. This and other new material 
allows the material from Riversleigh to 
be allocated to a third species of Baru 
(Figs 18, 19). 

TWO CRANIAL FORMS 

White Hunter Site has produced sev- 
eral posteriors of crocodihan skulls and 
skull decks representing 2 similar fonrts. 
No specimen duplicates portions of other 
specimens so although the cranial forms 
almost certainly pertain lo 2 of the taxa 
described above, they cannot be as- 
signed. 

WHITE HUNTER CRANIAL FORM I 

(Figs 20-22) 

MATER I A L. QMF3 1 075. 3 1 076 posterior of 
skulls; QMr3I077. skull fragment: 
OMF3107S, isolated parietal. 

FIG. 20 
dorsa! 
DIAGNOSIS. Supratemporal fenestrae 
tear-shaped with point directed an- 
terolaterally and with posterior shelf foniied by 
the squamosal; prominent expression of supra- 
occipital on skull deck; postorbital bar slender 
and round in section; posiorbital-frontal suture 
twice the length of postorbital-parielal suture; 
foramen magnum wider than occipital condyle; 
width of supratemporal fenestrae greater than 
width of postorbital; sculpture of more or less 
regular pits closely spaced. 

DESCRIPTION. Wide across the skull deck, 
high with the quadrate tucked under the squamo- 
sals. Supraoccipital prominent on the dorsal sur- 
face, forming a broad triangle almost excludmg 
parielals from posterior margin of skull deck. 
Supratemporal fenestrae with an anterior point, 
teardrop-shaped, with much of the posterior and 
medial portions closed by a iloor fonned by the 
squamosal and parieials inside the supratemporal 
fenestrae. Posterior face of the skull widi pro- 
nounced concavities on exoccipitals and squamo- 
sals for attachment of mandibular depressor 
muscles. Paroccipitai process encroaching ven- 
trally onto the quadrate. Foramen magnum sub- 
triangular, wider than the occipital condyle. 
Basioccipital with pronounced keel ventral to the 
occipital condyle. Quadrates steeply inclined. 
Pterygoids forming large portion of the posterior 
margin of the palatal fenestrae; internal nares. 
although not preserved, must have been well to- 




Cranial form I . QMF3 1075. posterior portion of skull, 
view. Scale = I cm. 



ward the posterior of the pterygoids. Otic meatus 
and foramina for the trigeminal nerve proportion- 
ally large. Laterosphenoids with a pronounced 
longitudinal crest mediallyonttic ventral surface. 
Sculpture on the skull deck distinctive, deep and 
well-defined pits separated by equally distinct, 
uniform walls. Pits close spaced. 

WHTTE HUNTER CRANIAL FORM 2 

MATERIAL. QMF3I079, anterior fragment of sktUl 
deck; QMF3 1 080> right postorbital. 

DIAGNOSIS, Small supratemporal fenestrae lat- 
erally compressed, shallowly floored by squamo- 
sals; postorbital bars inset from skull deck 
margin, robust and triangular in section; posior- 
bital-fronial suture equal in lengih lo posiorbital- 
parietal suture; width of postorbital greater than 
width of supratemporal fenestrae; sculpture of 
irregular shaped pits with irregular distribution. 

DESCRIPTION. WH 2 is described where it 

differs from WH 1. 

Supratemporal fenestrae narrower; squamosal 
Oooring makmg supratemporal fenestrae shal- 
lower posteriorly. Sculpture pits on WH 2 are 
small and irregular, separated by thick, irregular 
walls. Sculptured skull deck overhanging postor- 
bital bar on WH 2 but in WH I postorbital bar 



436 



MEMOIRS OF THE QUEENSLAND MUSEUM 



marginal. Postorbiial bar mod- 
erately robust, with triangular 
cross section. Postorbiial ver>^ 
large compared to the supra- 
temporal fenesirae. Triple 
junction between the postor- 
biial, fronial and parietal dis- 
tant from margins of 
supratemporai fenestrae. 

DISCUSSION. The Irontals 

associated withQMf3 1076 are 
very different from those re- 
ferred to Mekostwhtts 
{QMF31052) in being nar- 
rower and Hal between the or- 
bits. They are also deeper and 
have better dcfmed crania cris- 
tae fromalls than QMF31052. 
Thus cranial fomi 1 can be eon- 
fidently excluded from 
Mekosuchus (but not Bam or 
Qiiinkana). 

Although there are no Iron- 
tals unambiguously associated 
with cranial form 2. ihe difference in sculpture 
(compared with QMF3 1052) and the thickness of 
the orbit margins of the postorbiial make it un- 
likely thai this cranial form represents 
Mekosuchus. 

PAUAEOECOLOGY 

Four crocodilians have not previously been 
found in a single fauna in Australia, However, 
compared with world faunas, this is not an unusu- 
ally high diversity of crocodilians. particularly 
when the 4 species have differing head shapes or 
w hen Ihe site perhaps represents a thanaiocenosis 
collected from 2 or more habitats. 

Among extant crocoth'lians, many species have 

ranges that overlap but true sympalry is not com- 
mon. In parts of South America 5 or 6 crocodilian 
ranges overlap but rarely do 3 or more share the 
same habitat (Gor/^ula, 1987; Magnusson 8i 
Lima. 1991). The range of Crocodyltts porosus 
encompasses the ranges of C. johnstoni, C 
novae filtineae, C. mindorensis. C. siamensis, 
Tomisiowa schle^elii and parts of the range of C 
pafustrts and Gavialis gangeticus (Ross & Mag- 
nusson, 1989; Groombridge, 1987) but rarely do 
any of these species exist in true sympalry. Where 
C porosiis and C. johnstoni have been found in 
sympalry the larger C. porosns tends lo exclude 
C. johnstoni to the margins of the habitat or 




FIG. 21. Cranial form L QMF31075, posterior portion of skull, posieriur 

view. Scale = lem. 



sympatry is restricted by the need for different 
nestmg substrates (Webb et al., 1983). 

Modern studies of crocodilians in the Amazon 
Basin indicate that larger watercourses are occu- 
pied by larger, generalised crocodilians such as 
Melanosuvhus niger and Caiman crocoddus as 
well as Paleosuchus palpebrosus, while smaller 
watercourses in closed canopy forests arc occu- 
pied only by the more derived, deep-headed P. 
trigonatus (Magnusson, 1987, Magnusson & 
Lima, 1991 ). This could suggest that the variety 
of crocodilian head shapes al White Hunter Site 
IS Ihe result of a thanaiocenosis collected from 2 
or more different habitats. 

Thcorelically. 2 crocodilians may live sympal- 
rically when there are differences in head shape 
(implying exploitation of di ffercnl prey ) t)r where 
small, broad-snouted species can evade larger 
broad-snouted species by escaping to marginal 
habitats (Meyer. 1984). In no exlani crocodilian 
fauna, do 2 species share the same head shape 
(Meyer, 1984). 

The most diverse fossil crocodilian fauna is 
from the La Venia fauna in Colombia which 
consisted of 8 species (4 broad-snouted, 1 duck- 
bill, 2 narrow-snouted and 1 ziphodonl; Lang- 
slon, 1965). However, that fauna is a thanato- 
cocnosis from o\er240rn siraligraphically. Sym- 
palry was noi dcmonslraled. 



NEW CROCODILES FROM THE LATE OLIGOCENE 



437 



The Messcl fauna of Ger- 
many is more likely a bioceno- 
sis and has 6 crocodilian spe- 
cies including large and small 
broad-snouled forms, a shori- 
snouied form and two 
zipliodonts. A similar assem- 
blage has been recovered from 
Geisallal (Kuhn, 1938; 
Hauhold. 1983; Haubold & 
Krunibiegel. 1984). 

There are two possible expla- 
nations for the diversity of 
crocodilians in While Hunter 
Site. Barn wickeni is a large 
broad-snouled form while B. 
huberi is a much smaller broad- 
snouted form; Mekosncluts 
whhi'lnmtt'rensis is a small, 
short" snouted form; and 
Qiiinkana meholdi is a ziphod- 
oni. Compared to other fossil 
sites around the world and to 
modern analogues, the White 
Hunter assemblage differ 
enough to be sympatric ex- 
ploiting different niches. Alter- 
natively, they may indicate a 
thanaloccnosis from two or 
more different habitats. 

Arrangements of dift^ring ecomorphs of 
groups of mammals from sites at Riversleigh 
support the hypothesis that these faunas represent 
biocoenoses. A complex fauna of 8 species of 
bandicoots, belonging to clearly dellncd guilds, 
has been recovered from Upper Site C-). Muirhead, 
pers. comm.). Similarly, several species of ring- 
tail possums of differing ecomorphs have also 
been found in many sites at Riversleigh (M. 
Archer, pers. comm.). This pattern is apparently 
repealed in several other groups of mammals 
currently being investigated. That the Riversleigh 
rnanmtalian faunas repeatedly show sympatry be- 
tween several closely related laxa supports the 
hypothesis tliat Riversleigh sites preserve bioce- 
noses rather than thanatocenoces. This supports 
the hypothesis that the While Hunter crocodilians 
were also sympairic. 




FIG. 22. Cranial form 1, QMF31076, posterior portion of skull, dorsal view. 
Scale = Icm. 



niche separation. This is the first record of such a 
diverse crocodilian fauna from Australia but it is 
consistent with the structure and complexity of 
crocodilian faunas known from elsewhere. By 
comparison with other Riversleigh faunas the 
crocodilianfaunaof White Hunter Site was prob- 
ably typical for Oligo-Miocene Australia. 

ACKNOWLEDGEMENTS 

I thank John Scanlon, Mike Archer, Mark 
Norell and Ralph E.Molnarforconstructivecom- 
mcnts. I thank Mike Archer, Suzanne Hand and 
Henk Godihelp for access to specimens. 

This study was part of a PhD Studentship at the 
L^niversiiy of New South Wales. Financial sup- 
port was provided by the University of New 
South Wales, Friedrich-Wilhelms-LIniversitaL 
Bonn and the Rektor of thai institution. 



CONCLUSIONS 



LITERATURE CITED 



The 4 crocodilians from White Hunter Site 
include the first record of Mekosuchus outside 
New Caledonia and demonstrates a suiprising 
morphological diversity suggesting significant 



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43S 



MEMOIRS OF THE QUEENSLAKD MUSEUM 



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BUSCALIONI. AD. SANZ. J L. & CASANOVAS. 
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GORZULA. S. 1987. The managcmeni ofcrtJCodilians 
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GROOM BRIDGE. B. 1987. The dislribulion aiid status 
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UAUBOLD. H. 1983. Wiercliiere (Venebrata) In 
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KtJHN, O 1938. Die crooidilier aus dcm mtttlcren 
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LANGSTON> W. 1965. Fossil crocodilians from Co- 
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cations in Geological Science 52: I - 1 57. 



VI AGNUSSON. WE. 1987. Diets of Amarx»nlan croc- 
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MAGNUSSON. W.C. &. LIMA, A. 1*^9 1 . Tlic ecology 
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MEGIRIAN, D. 1994 A t»ew spccie.s of Quinkana 
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MEGIRIAN, D. MURRAY. P.P. & WELLS, R.T 
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crocodylian from the early Eocene of south- 
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WEBB. G.J W., MANOLIS. S.C (fe SACK. G C I^W 
Croaidylux johnsum't and C. porusus cocxif^lliig 
in a tidal river. Aus(rali;Hn Wildlife Rcseardi 10: 
639-50. 

WQ-LIS. PM.A. 1992. Four new crocfjdilians from 
early Miocene siu;s at Rivcrsleigh Station, norih- 
werrtem Queensland The Bcaule9- 269 

WILLIS. P.M A , MURRAY. PT & MEGIRIAN. D. 
1990. Barn dorrowi j^en. el .*/.». mn-., a large, 
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cfocodylidac) fioin niid-Teiiiary freshwaier lin»c- 
«lOTies in Ntirlhem Au.^tralia- MemoiT^ nf the 
Qucensliind Museum 29: 521-540 

WtLLlS. P.M.A., MOLNAR, K.E. & SCANLON. J.D. 
1993. An early Eocene crocodilian from Murgnii. 
southeastern Queensland. Kaupia; Danivst;idtcr 
B<^lirage /ur Naturgeschichic 3: 25-32. 

WILLIS. P.M A & MACKNESS, B. 1996 Qumkana 
^'fl^rtnw. anew species of ziphodonlmckosuchinc 
crocodile from the early Pliocene Blulf Downs 
Local Fauna, norlheni Australia wilh ;i a'vision o\' 
the genus. Journal and Proceedings of llic Lumean 
SocictyofNcwSoulh Wales 116: 143-151. 



MAYIGRfPIIUS ORBUS GEN, ET SP. NOV.. A MIOCENE DASYUROMORPHIAN 
FROM RIVERSLEIGH, NORTfW^STERN QUEENSLAND 

S.WROE 

Wroe. S. ]997 Of> 30: May i grip husorhus gcn.eisp.nov. a Miocene dasyuromorphian from 
Riversltfigh. northwestern Queensland. Memoirs cfthe Queensland Museum 41 (2): 439-448. 
Bxisbanc. ISSN 0079-8835. 

Mayigriphus orbus gen. cl sp. nov., an enigmatic Miocene dasyuromorphian from 
Rivcrsleigh, is described from dental maieri^i.The. hny Mayigriphus orbus shows a number 
of derived character- states forDasyuromorphia and two of these derived featuresmay .signify 
a special relaiionship with Planigale (Dasyuridae). However, no specialised features shown 
by M . orbus are unique to dasyurids wjihin the order and M , orbus also possesses derived 
characters shown by basal thylacinids. Because previous investigation has indicated that 
Dasyuridae is not currently defined by any dental synapomorphies, caution is demanded 
regarding allocation of M. orbu$ at the family level. Problems associated with the phyloge- 
netic placement of Af. orbus portend a story of growing complexity for dasyuromorphian 
phylogeny — a story progressively being revealed in the Tertiary limestones of Riversleigh. 
\^^^ayigriphus, dasyurornorphan, Miocene, Riversleigh. 

5- Wroe, School of Biological Sciences, Unhferslty ofNew South Wales, Sydney, New South 
Wales 2052, Australia; received4 November 1996, 



The fossil record for Dasyuromorphia is re- 
viewed by Wroe (1996b, 1997b). Uniil recently 
the pre-Pliocenc fossil record for 
Dasyuromorphia was limited to five described 
laxa, all from deposits in central Australia {An- 
kotarinja tirarensls and Keeuna woodburnei 
(Archer, 1976a), Wakamatha tassetli (Archer & 
Rich. 1979), Dasyiurinja kokuminola (Archer 
1 982a) and fhylacmuspotens (>^'oodbiime, 1 967 
). With ihccxccpiioDofThylacinuspoiens, Lnve;5;- 
ligaiion has failed to unequivocally link the^ 
fossil laxa with elements of modern 
dasyuromoqjhian radiations. More recentlyj the 
fossil-rich middle to late Tertiary deposits of 
Riversleigh have yielded six new thylacinid spe- 
cies: Nirnbacinus dicksoni (Muirhead & Archef» 
1990), Thylacinus macknessi (Muirhead, 1992; 
Muirhead & Gillespie, 1995); Wabuiacinus ridei 
(Muirhead, 1997), Ngamatacinus timmuhaneyi 
(Muirhead, \991),BadjcinusturnhulU (Muirhead 
& Wroe, in press), and Munbacinus gadiyuli 
(Wroe, 1996b). However, only two un-named 
laxa have been assigned to Dasyuridae: a possible 
phascogaline taxon known from a single M * or 
M ^ (Archer, 1982a) and an unnamed *An- 
lechinus-like' species from Riversleigh (Van 
Dyck. 1989). Wroe (1996b, 1997b) investigates 
problems with dasyurid phylogeny, concluding 
that Dasyuridae is currently defined by possibly 
three basicranial, but no dental synapomorphies. 
A new dasyiuximorphian descnbaJ hcrcshowsim 
enigmatic combination of features wuhtn 



DasyuTomoTphia and can not be unequivocally 
assigned at the family level. 

Dental nomenclature follows Floww (1867) 
and Luckett (1993). Taxonomic tcrrainolog)' for 
Dasyuromorphia follows Wroe (1996b), wilh 
three subfamilies recognised within Dasyuridae 
(Sminthopsinae, Phascogalinae [including 
Murexia], Dasyurinae [inci\}dir\^SeophascogaU 
BJid Phascolosorex]) and the following taxa con- 
sidered Dasyuromorphia incenae sedis: 4n- 
kotarinja tirarensis^ Keeuna woodburnei^ 
Wakamaiha tasselU and Dasyiurinja kokuminola. 
Higher level marsupial syslemalics follows Mar- 
shall et al (1990). Material is housed in the 
Queensland Museum (QMF). 

SYSTEMATICS 

Order DASYUROMORPHIA Gill, 1872 
Family INCERTAE SEDIS 

Mayigriphus gen. nov. 

TYPE AND ONLY SPECIES. Mayigriphus r^rhus 
|en. etspnov. 

GENERIC DIAGNOSIS. Mayigriphus orbus differs 
from all dasyunds in the following combination of 
features: Premolar row compressed longitudinally. P| 
very small; P3 reduced but with two roots; M\ com 
pressed on long axis with protoconid central on long 
and transverse axes with paraconid liny, Mi-^ 
melaconids and meiacristids reduced; Ml meiaconid 
not differentially reduced relative lo M2-4 meiaconids; 



440 



MEMOJRS OF THE QUEENSLAND MUSEUM 



MngUA* aniericjr(cmtmaiian0<cri«kl oWiquium Mi_4. 
with M3 cfjsiid obltqaa icrminaliug, beneath 
nit'tacrislid camasSial nolch; Mj 4 prou»ci>niil.s lin- 
gually shifted and rccurvcJ; M1-3 entoconids smiill 10 
liny. M4 lalunid reduced with ciituconid abscni. May- 
ighphiix orhus can be distinguished from knc^un thy- 
Incinjds by the following combination of features: Mi 
shijwsaga'utly reduced paracon id hut un!> moderately 
reduced mciaconid; clearly defined hypocuniilid nolch 
in anterior cinguUd of tower molars; very small size, 
reduction orP3 relative to P^; lack ul diaxiema between 
Pi and P^. Mayigriphus orbus differs from known 
bandicoots in po&scshjon of the alxjve conibmaiion of 
chiiraciers, a well-defined posterior cingul id ynd more 
buccaliy shifted hypoconulid. 

ETYMOLCKW. Wanyi mrjvL tooth: Lalin f^ripfiHU 
puzzle; refers to the cnigmauc combination of demaJ 
fcmurc».M«scuIine.. 

Mayignphus orbus sp. nov. 

(Fig. 1) 

ETV'MOLOGY. Latin orhu\, orphim. refers to iu un- 
ccaalu phylo^eneiic posiiiou. 

MATERIAL. Holoiype, QMF237«0. r*ghi dcmary 
wiih partial anterior alveolus of Pj. P| posterior root, 
1*2-3. M|-4; Puralype QMPi279l. rigtii denmry fragment 
containing M3 and'aivex>lu.-5 foir M4 AM from early late 
Miocene Encore Site, Riversleigl*. 

DESCRIPTION. Dentary bmken away anteriorly 
from midpoint of Pi anienor root alveolus and 
posteriorly from about I mm along ascending 
ramus; deniary almost uniform in depih. slight 
tapering anteriorly from beneath P^ prt)toconid; 
iDcnlai foramen beneath Mi hypoconid. 

Pi. Pi crown missing, only posterior half of 
anlvrior root aiveolus and posterior root remain; 
bused on root and alveolus size Pi small, less than 
half P: length; anieHor ulvcolits bua^ally dis- 
placed. 

P2. Nn diastema between Pi and P:; twin 
focned; Pi latgesi premolar in height and length; 
protoconid moderately worn; buccal cingulid 
runs posteriorly from midpoint between anterior 
and posterior roots to small posterior central 
cuspulc on heel; another ciii^ulid circumscrib<?s 
the lingual crown base from this cuspule to ante- 
nor raargm of posterior root. 

P3. no diastema between Pj and P3; twin rooted; 
P3 morphology similar to Ft but differs in posses- 
sion of continuous cingulid circumscribing base 



of entire civ^wn artd s»ialle/ she (Pi around 30 
percent smaller in height an length) 

Mt. no diastema between P_i and Mi; Mi un- 
worn; principal cusps in order of deerciising 
height; protoconid, mctaconid. paraconid. 
hypoconid and enioconid; enioconid tiny, closely 
abutting posterior lace of irigonid adjacent lo 
mctaconid, paraconid damaged, but from basal 
dimensions was clearly small; ineiaconitl small 
and shifted posteriorly; protoconid dominant 
cusp, lingually recurved, occupying almost ccn- 
U"al position on tooth; Mi reduced on the long 
axis; lalonid small, slightly wider transversely 
than Irigonid but shorter on long axis; parucrislid 
paiallel to, atid cristid ohliqua shows slight lin- 
gual inllcction at anterior end; nietacrisiid and 
hyptx'ristid parallel and angled at about 20' to 
Iransvcrse axis oi dcnlary; cnslid oblicjua lenni- 
nates beneath apex of pnUoconid; anterior 
cingulid runs basally from piiracomd a> beneath 
protoconid; posterior eingulid weakly developed. 

M:. Mj differs from Mi as tollow.s; M2 much 
larger; paraconid much larger; metaconid rela- 
tively and absolutely lager Ihciugh still small 
eompared to protoconid; (ulonHl shoitct on long 
axis: eristid obliqua terminates against posicriof 
face of trigonid in more lingual position, wilh 
angle formed between cristid obliqua and 
hypocristid more acute; metacrisiid and 
hypocristid run closer to transverse axis of dcn- 
tary; entoconid relatively larger than in Mt 
though still small; posterior cingulid more 
stR>ng]y developed; M? paraeristid runs at about 
30° to iong axis of dcnlary. with angle between 
paraerislid and mciacristid slightly less than 90^, 

M3. M3 differs from M: as tbilows; protoct)nid 
larger; on transverse axis irigonid wider and 
talonid shorter; entoconid on M3 smaller; cristid 
obliqua terminates m a more lingual position 
against posterior face of irigonid beneaih carnas- 
sial notch of metacrisiid. 

M4.M4similarloM.i except: metaconid smaller 
than paraconid; protoconid less lingually re- 
curved; talonid greatly reduced, entoconid ab- 
sent, hypoconid and hypoconulid small; no 
posterior cingul id. 

Merisiic gradients from M1.4. oricntalion of 
mctacristid and hypocristid to long axis of dcn- 
lary increasingly transverse from M1-2. departs 
away from transverse from M2-1, then back to 
more transverse orientation from M.v^; orienliih 
lion of paracristid to long axis of dentary increa^- 

FIG. 1. Mayigriphus orbus Sp. nov,. QMF237S0. hololype. A, buccal view, B and D. stereo-pair, occlusal view 
of M1-4. C and E. stereo-pair, occlusal view of P 1 posienor alveolus. P2-3, M1.4. 



MAYIGR/PHUS ORBUS, ENIGMATIC DASYUROMORPHIAN 



44 




442 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE L Mayigriphiis orbus. denial measuremcnis {mm). I=anleroposlerior dimension, wl 
verse dimension ol irigonid, w2-niaximum irans verse dimension of Lalonid. 


=nmxinttui2 iraiu- 


Ref no 


P2 P3 


Ml 


M2 1 Ml 


M4 




Icn 


wid 


len 


wid 


1 


wl 


w2 


1 


wl 


w2 1 


w 1 w2 


1 


wl 


*2 


OMF 23780 


0.83 


0.42 


0.67 


0.36 


1.26 


0.64 


0.67 


1.49 


0.84 


0.86 1 L52 


0.84 078 


L33 


0.74 


0.4> 


OMF 22791 


















I L45 


0,86 


JfJ.6S^ 









ingly transverse from Mi-?, termination of crislid 
obliquu against posterior face of irigonid increas- 
ingly lingual M1.4; reverses departing from trans- 
verse for Mv4; protoconid and mctaconid hcigiil 
increases Mi.;*, decreases Mu4; paraconid height 
increases M1.4; talonid width increases Mi.^y de- 
creases M.V4. 

CHARACTER ANALYSIS 

Wroe (1996b. 1997b) discusses^ laxa consid- 
ered appropriate in the reconstruction of a 
da.syuromorphian morphotype (i.e., pcradectids, 
didelphoids, microbioiheriids, peramclc- 
morphians). Siuiilar iiU'lhodology is usoii for 
characters treated here to assess character-state 
polarities (Tables 2, 3). Within f^asyuromorpl^ia 
Ankoiarinja tirarensis is the least derived taxon 
and a possible ouigroup lo reimiiriing 
dasyuruinoqihians. Muribacinm gadiyuli and 
Badjihttis fiir/ihulli aie The IcasT derived Ihy- 
tacinids. For Dasyuridae tlic ibiiowing taxa arc 
considered plesiomorphic for their respective 
sub lam i lies: Murexia longicaudata 
(Phascogaiinae), Neophasco^ale ioreruzU 
(Dasyurinae) and Sniintbopsis IctnopHs 
(Sminlhopsinae). 

P^. A P3 larger than P2 is plesiomorphic for 
ouigroups to Dasyiiromorphui (Wroe, 1996b, 
1997b). Fov A. titarensis P? is slightly smaller 
than P:. Wakormftha tasselli shows P3 larger ihmi 
P2. The basal thylacinid Bcuijcinus turnhulli has 
P3 slightly smaller than Pj. Wroe (1996b) inter- 
prets similar Pj morphology for Murihacinus 
gadiyuli i^n the basis of alveolar dimciisiotis. For 
all remaining thylacinid laxa P3 is larger than P2. 
with the largest P5 in the most derived taxa (Thy- 
lacinus). Within Dasyuridae wide variation is 
apparent lor this leaturo. Taxa treated here its 
basal to their respective subfamilies show a P^ 
larger than P2 {Murexia, Sminthopsis lcitcopi4x)^ 
or slightly reduced iNeuphast-ffsaU'). However, 
within each subfamily some taxa show marked 
reduction or absence of Pi. P3 reduction in May- 
i^riphus orbus exceeds that shown by all taxa 
considered Dasyuromorphia inccrlue sedis, all 



Thylacinidae and basal laxu for dasyurid sub- 
families. 

MI paraconid. The Mi paraconid is nol reduced 
in dasyuromorphian <_»utgroUps and A. tirarensis, 
W. iasselli, moderate reduction is shown by thy- 
lacinids (excepting ft. lurnbidH which shows 
marked reduction 1 and plesiomorphic Sminlh- 
opsinae (Smmthopsis leucopus). Phascogalinac 
(Marexia) and Dasyurinae {Neophascoiiale) M 1 
paiaconid reduction in M otbus is less marked 
than in all Dasyurinae excepting Ncophasco^ali' 
For A/, orbus M] sh4)ws greater reduction of the 
paraconid than for all Pha.scogalinac and 
Sminlhopsinae except Planigale. 

M4 talonid. The M4 talonid is unreduced in basal 
taxa for Dasyuromorphian ouigroups cxccpl ban- 
dicoois (e.g., Yarala burdijivldi. Muirhcad & 
Filan. 1995). Within Dasyiironiorpliia/^. tiraren- 
sis, basal phascogaline {Murexia) and diisy urine 
{Neophascogale) laxa show slight reduction for 
this feature. The IS/Li talonid is greatly reduced on 
ihe plesiomoq^hic dasyuromorphian condition 
for W. tassclli, most dasyurines and 

f)hascogalines, and all sminthopsines. For Iby- 
acinids the M4talonid is unreduced for basal taxa 
[Muribiuinus, Badjcinus), but significantly re- 
duced for derived species [Thylarmus, 
Wabulucinus). Even for specialised Thykuinus 
M4 talonid reduction does not approach that of 
derived dasyurids which show tar greater dimi- 
nulicm on the transverse axis The degree of re- 
duction for this feature in M. orbus is closest lo 
tliat shown in Phascogalc. but less than for most 
Dasyurinae, and all Sminlhopslmu:. 

Crislid obliqua orientation. A buccal position for 
the anterior termination for the cristid obliqua 
relative to the camassial notch ol the niclacnslid 
is common to iitosl dasyunnriorphian outgroup 
taxa. This feature is associated with crislid ob- 
liqua orientation and tormaiion of a right angle 
hotwccn the cristid obliqua and hypocrislid. Most 
ouigfoup taxa lo Dasyuiidac and Thylacinidae 
show u cristid obliqua aligned closely with the 
long axis of ihedcntary and a 9(r angle is formed 
belwcen the cristid obliqua and hypocrislid. In the 



MAYIGRIPHUSORBUS, ENIGMATIC DASYUROMORPHIAN 



443 



character-analysis (Table 2) only M3 is consid- 
ered. Buccal termination is shown by A. tirarensis 
and, to a lesser degree, by K, woodburnei and W. 
tasselli Basal thylacinids show relatively lingual 
anterior termination for the crisiid obliqua, but a 
more buccal position is apparent in Thylacinus. 
Basal dasyurines show lingual termination for the 
cristid obliqua (Neophascogale. Myoictis, 
Dasyunis halhicatus), but more specialised taxa 
show buccal termination (other Dasyurus, 
Dasycercus, Dasyuroides, Sarcophilus) . All 
phascogalines show lingual termination. 
Sminthopsines show buccal termination. May- 
igriphus orbits shows lingual termination. 

Orientation of the cristid obliqua correlates 
with other features of both the upper and lower 
molars. These include the angle between the 
postparacrista and premetacrista (together termed 
the centrocrista), the relative size of the pro- 
loconid and melaconid and, the occlusal surface 
area presented by the prolocone and lalonid basin. 
Scoring of character states for cristid obliqua 
without consideration of these associated features 
may be phylogenetically misleading. For exam- 
ple, derived Thylacinus and some dasyunnes 
show longitudinal alignment for the M3 cristid 
obliqua (unspecialised didelphids, microbio- 
theriids and A. tirarensis), but for these derived 
dasyuromorphians this feature correlates with 
protoconid hypertrophy, melaconid reduction or 
loss, and a linear centrocrista. These character- 
states are all associated with the dominance of 
longitudinally oriented vertical shearing crests. 

The basal position for Sminthopsinae within 
Dasyuridae indicated by molecular analyses 
(Kirschelal, 1990; Krajewski et al., 1993; 1994) 
supports the contention that a buccal point of 
termination for the cristid obhqua is a plesiomor- 
phy for the clade. However, dental features of 
Sminthopsinae are products of a different selec- 
tive regime and transverse rather than longitudi- 
nal vertical shearing crests dominate. Archer 
(1976) noted that a buccal position for the cristid 
obliqua may be associated with reduction of the 
paracone or a lingual shift in the camassial notch 
(of the mctacrislid). Both derived features are 
shown by sminthopsines and it is probable that 
cristid obliqua position represents a correlated 
apomorphic feature. A further derived feature 
shown by sminthopsines is gross reduction of the 
talondis on the anteroposterior axis which may 
also impact on cristid obliqua orientation. For 
sminthopsines and derived dasyurines and thy- 
lacinids, a buccal position for the crisiid obliqua 
is treated as derived relative to that of microbio- 



iheriids, unspecialised didelphids, bandicoots 
and A. tirarensis (Tables 2, 3). A relatively more 
lingual termination for the cristid obliqua, as 
shown by most dasyurids and basal thylacinids, 
is also considered derived. The character com- 
plex associated with most dasyurids (a relatively 
lingual anterior termination point and acute angle 
formed between the cristid obliqua and hypo- 
cristid) is scored as 'a'. Buccal termination and 
formation of 90*^ between the cristid obliqua and 
hypocristid may be associated with increased 
transverse vertical shear (b) or increased longitu- 
dinal vertical shear (c). 

Angle between paracristid and metacristid. For 
dasyuromorphian outgroups an acute angle is 
formed between the paracristids and metacristids 
(mirrored by an equivalent angle fomied between 
the posimelacristae and preprotocristae with 
which they occlude in the upper dentition). Sim- 
ilar morphology is shown by A. tirarensis, W. 
tasselli and sminihopsine dasyurids. All dasy- 
urines, phascogalines, basal thylacinids for which 
a metacristid is retained, and M. orbus show a 
relatively obtuse angle between paracristids and 
metacristids. Widest paracristid-metacristid an- 
gles are in Sarcophilus, Glaucodon and D. 
maculatus among dasyurids and Ngamalacinus 
among thylacinids. This phenomenon is corre- 
lated with carnivory and the development of lon- 
gitudinally aligned vertical shearing crests. 

Hypoconulid notch. Many marsupials have a dis- 
tinct notch in the anterior cingulae of their lower 
molars to receive the hypoconulid of the preced- 
ing tooth. Oulgroup data for Dasyuromorphia 
regarding this feature is equivocal. Some out- 
group taxa (e.g., some peradectids) show a well- 
developed hypoconulid notch, but among other 
possible outgroups this feature is absent (e.g.. 
pcramelemorphs). Within Dasyuromorphia this 
feature is well-developed for Ankotarinja 
tirarensis and Keeuna woodburnei, but poorly 
defined for Wakamatha tasselli (see Wroe 
(1996b) re arguments for possible bandicoot af- 
finities of this taxon). Among thylacinids, a well- 
developed hypoconulid notch is present for 
Muribacinus and Badjcinus, weakly-defined in 
Ngamalacinus, and absent in ail other taxa. A 
well-developed hypoconulid notch occurs in all 
dasyurids excepting Dasyurus maculatus (re- 
duced), and Glaucodon and Sarcophilus (absent). 
Mayigriphus orbus has a well-defined hypo- 
conulid notch. Wroe (in press b) infers that loss 
of the hypoconulid notch is a function of ad- 



444 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 2. CharaciCis and chiiraclcr-siulcs usee! in phylogcnetic analysis. 



CI. Py size rclanvc lo P2. 0. larger. L reduced. 2. intermediate. 3, Uny. 4. ah^enu 

C2. Ml paraconid size. 0. large. I . reduced. 2. tiny. 3. absenL 

C3. M4 lalonid size, 0. large. 1 . moderately reduced, 2. markedly reduced. 3 liny. 

C4. M3 cristid obliqua morphology *. P- plesiomorphic. a. lingual, b. trans, shear, c. long. shear. 

C5. Angle between paracristid-metacristid. 0. acute. L intermediate. 2. obtuse. 

C6. HyptKonulid notch. 0. well developed. 1. intermediate. 2. absent. 

C7. Mclaconid morphology. 0. no clear diJTerenlial between Ml and M2-4. I. clear differential. 

C8. M2-4 inctaconid si2e. 0. large. 1 . reduced. 2. greatly reduced. 3. absent. 

C9. M} entuconid size. 0. large. 1. reduced, 2. absent. 

* Three derived states recorded for this character tsce text). 



vanccd camasiialisation tor derived dasyurids 
and ihylacinids. noting (hat the shift to a predom- 
inance of longitiidiiial vcrlical shejir in marsupial 
carnivores diminishes the requirement fora brace 
against transverse forces (i.e., the likely role for 
the hypoconulid notch). 

Archer (J 982b) regarded a hypoconulid notch 
in the anterior cingulum as a possible dasyurid 
synapomorphy. But Wroe ( 1 997) concludes that 
it may have been in the common ancestor of 
Dasyuromorphia and was almost certainly in the 
a)rnmon ancestor of Dasyuridae-Thylacinidae. 
For specialised dasyurids iSarcopfulus and 
CUiucodon) and \\\y\-iic\f\\i\Sr{ThyUiciHt4s, Wuhul- 
(H'ituis) loss of the hypoconulid notch correlates 
with advanced carnossialisation. 

Mclaconid . A well-developed mctaconid on all 
lower molars occurs in all putative Dasyuru- 
murphian outgroups. The same is so for A< 
tiraren.sis, K. yvoorihuntei und W. tasseJIi. All 
ihylacinids show marked reduction or loss of the 
meiaeonid on all lower molars. Among 
Dasyuridae this feature is variable. Plesio- 
morphic ia^\a for each subtamiiy show no reduc- 
tion of the metaconids. However. specialJNed 
Dasyurinae and Sminthopsinac show derived 
characicr-stale.s. Derived dasyurincs sht)W a re- 
duced M| mctaconid, but less reduction lor M3-4 
Ttietaconids iDasyccnus, Dttsyuwuies, Dasy- 
urus. Sa/cophilus). In Plnfii}^ali' meiacDiiid dim- 
inuiion is les.s iidvanced on Mi and more uniform 
through M2-4. This phenomenon shown for 
t^lontgole is also common to ihylacinids which 
relain nielaconids (except Badjcinus tiimbulU 
which shows the typical dasyurinc condition). 
Mu\l^riphi4s orhi4S .shows uniform reduction of 
Ml 4 mclaconids, ihe chanicter-.slate conimc^n lo 
Pkmigale among dasyurid.s and Muribacinus, 
Nimbacinus and N^atnalacuiHs among thy- 
liicinids, 



Localised meraconid reduction (i.e., a clear dif- 
ferential shown between Mi and M^-j meiacoiiid 
reduction) as shown by some dasyurines, 
sminthopsincs and Badjcinus, is probably related 
to brachycephalisation, shortening ol the tooth 
row on the anteroposterior axis and concomitant 
prem(jIarisaiion of Mi (Archer, 1976). 
Generalised inetaconid reduction (Mi-4) corre- 
laies with increased size oi the proiocunid and 
primacy of the paracrisiid and postmctacristae in 
vcnical .shear. For carnivorous dasyurids and rhy- 
iacinids these derived features ai'e associated with 
alignment of the venical shearing crests with the 
long ixm of the looth row. 

Enloconid. Large entoconids arc plesiomorphic 
fordasyuromorphian outgroups. A rirarcnsis, AT. 
woodhurneL basal dasyurines and sminthopsincs, 
and phascogalines. Entoconids are reduced t«- 
absent in some Sminthopsis and Antevhiitomya 
among smmthopsines and Paratitechimis, 
PscudantedUmts, Dasyuroides, Dasyrrrcus, 
f}<isvkuluUi, most Dasyunts (excepting D. 
hallticauis), Surcoplultt.s and Glaucodon. All thy - 
lacinids show .some eiUoconid reduction, with the 
least reduction \r\MHribuiinus:xni^ the greatest by 
Thylacinus. Mayi^riphus orbus shows mi)deratc 
reduction ft>r this feature Archer ( 1 98 i ) and San- 
son ( 1 985) note that no clear form-function rela- 
lion.ship explains the distribution of entuconid 
reduciiun and loss among dasyurids. bul note that 
this reduction is greatest for arid-adapted species 
(some Sminthopsis. Antechinomys. Lkisyuroides, 
Dasycerctis) However, considerable reduction 
or loss is shown for some species found in less 
extreme environments and for large dasyurid and 
thylacinid carnivores this phenomenon is likely 
associated with carnivory. More form-function 
data is required here. 



MfAYIGRfPUVSORBm, EKTGMATIC DASYL»ROMORPHlAU 



44 *» 



TABLE 3 Characler/taxonmatr2?L 



Mayigriphus orbui 


2 1 1 a2 


00) 1 


Alphadon marshi 


DOOOO 


0000 


Marmosa sp 





0000 


Duieiphis marsupujHs 


10000 


0000 


Drtvmciops austrnlts 


00100 


1000 


Yarala burchfieldi 


3 


2 00 


Ankotarinja iiraransU 


1 00 


000 


Keeuna woodburnti 


???a 1 


0700 


Wak/imalhij tasselli 


07200 


1700 


Murihacinus gadiyuli 


I 10ft2 


0010 


Hadjcinus lurnhulU 


no«2 


01 10 


Nimharinus dicksoni 


?or?2 


70 17 


Nganialar.inus timmulvaneyi 


'/Old 


1020 


Wabuiacinus ridet 


077c2 


172 2 


Thyiacinus machu^ssi 


OOU? 


1731 


Thylacinus cynocephalus 


002c2 


1032 


Smimhopsis leucopus 


102b0 


0002 


PlamgaU macidaia 


2l3b0 


0012 


PlanigaU gilest 


3l3b0 


0012 


Planigale tenurostrix 


2I3bO 


0012 


Murexia tongicaiuiata 


OIUI 


0000 


Phascogale tapolafla 


lOIal 


0000 


NeophascogalE lorentni 


lOIttl 


0000 


Myoirxis meias 


12I11I 


0000 


Paronltchinus apicalis 


223cl 


0102 


Dasyurus hallacatus 


322»1 


n 10 


Daysurus tnaculatus 


322c2 


2121 


SarcophiliLs harrisii 


323c2 


2121 



DISCUSSION 

BIOSTRATIGRAPHY AND ECOLOGY. To 
date M. orbas is restricted to early late Niiocene 
Archer ct al. (1995) Encwe Site at Riverdcigh 
Encore has produced a fauna ihai includes several 
unique laxa, including a large dasyuromorphian 
of uncertain affinity (unpubL data), a giani 
Ekaltadeia (Wroe, 1996a), a derived koala 
(Black, pers. comm.), a palorchestid structurally 
intermediate between species from Riverslcigh 
Syslejn C and the late Miocene Palorchesies 
oainei of Alcoota (Black, 1997) and a Warenja- 
like wombat CAxcher el al., 1995). The rootless 
teeth of this wombat (unknow n for other species 
ai Riversleigh) and the relatively low abundance 
of the frog Lechriodus inter gervis, common in 
other Miocene Riversleigh deposits (Godihclp. 
pers. comm.) indicate ihat climatic conditions 
may have been drier for the depositional episode 



during which Enoorc was produced, Tenuiiivc 
support for a relatively laie age for Encore, site. Is 
also forwarded by Wroc (1997a). If M. orhu,^ is -^ 
dasyurid then the derived dentition (relative lo 
other Miocene dasyurids) might also suggest a 
laic age for Encore site. As noted above for small 
dasyurid*;, circumiitantial evidence correlates cn- 
loconid reduction with adaptation to rejativcly 
dry environn^ents. 

Mayigriphus orbus is the smallest dasyiim- 
morphian from the Oligocene and MioccrJC of 
Riversleigh and is comparable to Planmile. 
maculatus in size. Only one other marsupial io- 
seciivore has been identified that might have 
competed closely with M. orbus, the diminutive 
bandicoot Yarala burchfieldi (Muirhead, 1995). 
As with modem Flanigaie (Denny, 1 9S2) the diet 
of M. orbus probably included invenebr^tc^^ 
frogs, small lizards and/or small mammals. 

PH^XOGBNY. Mayigriphus orbus shows a 
unique mosaic of features among dasyuro- 
morphlans. Two features of A/ orbus may imli- 
cate a relationship with Planigale (the greatly 
reduced Mi paraconid concurrent with a mocicr- 
ately reduced Mi meiaconid, and rclauvcly uni- 
form diminution of the Mm meiaconids). 
Although a comparable degree of Mi paraconid 
reduction is al.^ common to many derived 
dasyurines (e.g., Pseuiianiechintis), *un these laxa 
diminution of itte Mi mctaconad is far mnrc ad- 
vanced and a clear differcniial is produced be- 
tween thai shown by Mi and M2-4- Additional 
apomorphics shared by M. orbus and Planigale 
(e.g., reduction of M4 talonid, entoconid and Pi ). 
arc also found in other specialised dasyurid la;ta. 
On the basis of cyiochrome-b data, Painter ei al. 
(1995) estimate the oldest branchings withm 
Planigale ai 11-15 raya, thus the possibility that 
A/, orbus represents an early branch of this radi- 
ation can not be discotmted. However. M. orbus 
shows at least 2 derived features not in Planisiale 
(wide angle formed between the paracri.iid, 
metacristid, a buccal shift in the point of termi- 
nation of the cristid obliqua). The oldest material 
clearly attributable to Plamgole is Pliocene 
(Archer^ 1 982a). Based on available daia, a sister 
laxa association for A/, orbus with Planigale is 
considered equivoc-al. 

Even at the family level, the phylogenetic po- 
sition of Af. orbus is considered uncertain, be- 
cause it has a number of features that mighi be 
mteTpreted as synapomorphies for either de- 
rived dasyurid or Uiylacinid clades. but no un- 
equivocal synapomorphies (wjthin Dasyurn 



446 



MEMOIRS OF THE QUEENSLAND MUSEUM 



morphia) for either family. Two synapomorphies 
for the Dasyuridae are in the lower dentition of 
M. orbits: reduction of Pi (Tate, 1947; Archer, 
1982b; Marshall, 1990) and the hypoconulid 
notch in the lower molars (Archer, 1982b). Status 
of both as shared-derived features for Dasyuridae 
is questioned by Wroe (1996b, 1997b). Reduc- 
tion of P3 is certainly common within Dasyuridae 
which culminates in the loss of this tooth among 
specialised taxa. However, reduction or loss of P3 
may have occurred independently at least 3 times 
in the Dasyuridae (Archer, 1981). A further argu- 
ment against the phylogenetic value of this char- 
acter at the family level is the P3 smaller than P2 
in 2 thylacinids from Riversleigh (Muirhead & 
Wroe, in press; Wroe, 1996b, 1997b). The status 
of the hypoconulid notch as a shared derived 
feature for dasyurids has been undermined by the 
discovery of plcsiomorphic thylacinids with a 
well-delined hypoconulid notch in the lower mo- 
lars (Wroe, 1997b; Muirhead & Wroe, in press). 
Marked reduction of the Mi paraconid in M. 
orbiis is common to specialised dasyurids but not 
thylacinids, excepting Badjcinus turnbulli 
(Muirhead & Wroe, in press). None of these 
features represent unequivocal synapomorphies 
for Dasyuridae and each have been independently 
derived within specialised dasyurid lineages. At 
least 3 features in M. orbus suggest a possible 
alliance with thylacinids. Firstly, the lack of a 
clear differential between metaconid reduction 
on Ml and M2-4 in M. orbus is known only for 
Planigale among dasyurids but common to 
plcsiomorphic thylacinids. In all dasyurids ex- 
cept Planigale, Mi metaconid reduction clearly 
exceeds that of M2-4. Although reduction of the 
M2-4 melaconids is less pronounced in A/, orbus 
than in known thylacinids, excepting 
Muribacinus, it is greater than for most dasyurids 
except D. maculatus, Sarcophilus and Planigale. 
Secondly, the wide angle formed between the 
paracristid and metacristid in M. orbus is found 
in basal thylacinids, but only D. maculatus, 
Sarcophilus and Glaucodon among dasyurids. 
Thirdly, among specialised dasyurids, reduction 
of the Ml -4 talonids and melaconids is commonly 
associated with a buccal shift in the point of 
termination of the cristid obliqua. In both M. 
orbus and plcsiomorphic thylacinids this is not 
the case, with the cristid obliqua terminating in a 
relatively lingual position. Ultimately, this may 
be related to Ride's (1964) observation of a dif- 
ference between Thylacinus and specialised 
dasyurids in the composition of the principal pos- 
terior shearing crest. Ride pointed out that, in 



Thylacinus, the posterior shearing crest runs from 
the protoconid directly to the hypoconid, while in 
derived dasyurids (especially Sarcophilus) the 
posterior shearing crest connects the protoconid 
and metaconid. 

CONCLUSIONS 

KM. orbus is a dasyurid it represents the most 
denved member of the family known from pre- 
Pliocene times, with the possible exception of the 
Miocene Dasylurinja kokuminola (Archer, 
1982a) from Lake Yanda in central Australia. A 
special relationship between these two taxa can 
not be discounted, with both showing 
specialisations associated with camassialisation 
(the much larger size of D. kokuminola precludes 
the possibility that the 2 taxa are conspecific). D. 
kokuminola is known only from a single upper 
molar and direct comparisons with Af. or^w^ can- 
not be made. Among known dasyurids M. orbus 
shares the greatest number of derived features 
with Planigale: two character-states (disparate 
reduction of the Mi paraconid and metaconid and 
uniform reduction of the M1.4 metaconids) sug- 
gest the possibility of a special relationship for 
the 2 taxa. However, uniform diminution of the 
Ml -4 metaconids is also shown by some basal - 
thylacinids. 

Mayigriphus orbus shows no unequivocal syn- 
apomorphies for either dasyurid or thylacinid 
clades. For Dasyuridae, unique derived-features 
(within Dasyuromorphia) are only found in the 
basicranium (Wroe, 1996b, 1997b). Unique de- 
rived features (within Dasyuromorphia) uniting 
Thylacinidae have been identified only in the 
upper dendlion (Wroe, 1996b). Neither region is 
known for M. orbus. Confident phylogenetic as- 
signment for M. orbus has been further tempered 
by the identification of possible thylacinid 
apomorphies in this taxon, which must be consid- 
ered in the following context: investigation of 
Oligocene and Miocene material from 
Riversleigh is revealing a complex dasyuro- 
morphian phylogeny dominated by a diverse thy- 
lacinid clade, showing greatly expanded 
intrafamilial variation (Muirhead, 1992, 1997; 
Muirhead & Archer, 1990; Muirhead & Wroe, in 
press; Wroe, 1996b, 1997b) a close relationship 
between Dasyuridae and Thylacinidae has been 
established by molecular studies (Lowenstein et 
al, 1981; Sarich et al., 1982; Thomas, 1989; 
Krajewski et al., 1992), and a relatively recent 
genesis for Dasyuridae has been suggested 
(Archer, 1982a; Krajewski, 1992; Wroe, 1996b, 



AM}767f//'//t/y a/?fl6'5. ENIGMATIC DASYIROMORPHIAN 



447 



1 997b), Given this emerging climate olcomplcx- 
ity for dasyuromorphian phylogeny, Ihe curituis 
nwx ortlerivcd iealures in M. orhus (olherwisc 
considered diagnostic of either specialised 
dasyurid or ihylacinid clades) makes reliable 
placement whhin cither family impossible, par- 
liciilarly as Ihis decision must currently be based 
&;}lcly on elements of the lowci dcniition. Prob- 
lems associivled with determining the phyloge- 
nctic position of M. rtrhus highlight an 
unexpected phylogeneiic scenario. Despite an 
abundance of dasyuromorphian material from 
Oligoccne-Miijcene deposits of Riverslcigh no 
Ujxun has yet been described which can be un- 
equivocally associated with elements of the now 
ubiquitous dasyurid radiation. 

ACKNOWLEDGEMENTS 

M. Archer. H. Godthelp, and J. Muirhead pro 
vided invaluable assistance through their con- 
SJruciive enticism and comment, J. Muirhead 
also kindly took the photographs presented. Vital 
i^uppori for this research has been given by the 
Australian Resc:irch Council: the National Estate 
Grants Scheme (Queensland); the Department of 
Environment, Sports atid Territories; Ihe Queens- 
land National Parks rmd Wildlife Service; the 
Comnumwcallh World Heritage Unit fCan- 
hcrra); ihe University of New Svuuh Wales; ICI 
Australia; the Australian Geographic Society; the 
Queensland Museum; the Australian Mtisetun. 
Century Zinc; Mt Isa Mines; Surrey Bcatly & 
Sons; the Riverslcigh Society . ; the Royal Zoolog- 
ical Society of New South Wales; the Linnean 
Society of New South Wales, and many private 
supporters. Skilled preparation ol most of the 
Riversleigh material has heen carried out by 
Anna Gillespie, 

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Dasyuridae. Journal of Mammalian Evolution 4: 
19-52. 



STRATIGRAPHY AND PHYLOGENV OF THE GIANT KXTINCT RAT KANGAROOS 
(PROPLEOPINAE. HYPSIPRYMNODONTIDAE. MARSUPIALIA) 

S. WROE 

Wroe, S. \991 06 30: Stratigraphy and phylogcny of the giaiu cxuaci Rai-kangaroos 
(Propleopinac, Hypiiprymnodoniiduc. Marsiipialia) Memoln of The QuraishnJ Mtiwum, 
41(2): 449-456. Brisbime. ISSN 0079-8835- 

The Giant Rat-kangarooK vvca* placed in the I'mplcopinae by Arclict &. Flanncry ( 1 985) iind 
in ihc Hypsiprymnodontidae by Ride { 1993). Cladislic analysis ul tkctltatUla material from 
Rivcrslcigh, northwestern Queensland (Wroe, l9%)sugge,stcdthaiamiddleiolate Miocene 
dichotomy in Ekahadeia nia>' have ptoduced two lineages of Pho-Picistoccne Fropleopus, 
indicating polyphyly for Pwpieopns and paraphyly for EkalmJeia. Melrical data for pro- 
pleopincs and siraligraphic infomiatiiin support \Vroc".s (1996) cladislic analysis of pro- 
plciipines. □ Propleopinac, ilyp.Hiptymnodontidae, Riversletgh, Ekaltadeta. cladhtics. 

S. Wrot. School ofBhb^icul Sctvnvv, Umyently of New South V^/cdva. NS.W, 20fi2 
Auxiralia: received 4 November 1996. 



Giant Rat-kangaroos (Hyp-siprymnodontidac: 
Propleopinac) may be the picsiontorphic sister 
group of potoroids (Flanncry, 1987). Archer & 
Flanncry (1 985) considered Ekaltadeta i!na,iP\g. 
I) the .sister group to Propleopus De Vis, 1888 
with M oscillans De Vis. 1888 (Fig. 2) the more 
piesiomorphic and P. chillagoemis Archer et a1. 
(1978) (Fig. 2) ihc inore upomorphic within Pro- 
pleopus (Fig. 3). Pri>plc(>pinc species described 
since 1 985 are Ekaltadeta janiiewulvaneyi Wroe 
1996, (Fig, 4) and Jackmahoiieya toxonieusis 
Ride. 1993. Wroe (1996) suggested another pos- 
sible phylogeny for the Propleopinac with £\ ima 
and P. chillagoensis forming the sister group to 
another cladc containing a new species, £. 
Jamiemtilvaneyi, as the sister laxon to P. 
wellin[^tonensis and P. oscillans.iF'ig. 5). As an 
adjunct to the ciadistic analysis (Wroe, 1996), 
metric and .stratigraphic data for propleopines are 
used 10 clarify intrasubfainilial relationshtps. 

Dental homology for premolars follows Flower 
1.1867) and Luckett (1993.) for moiars. Higher 
level systematics of kangaroos follows Flanncry 
( 1987) and Ride (1993). Specimens arc housed in 
the Queensland Museum (QMF). Other prefixes 
include; UCM (University of Calilbrnia Mu- 
seum), NMV (Museum of Victoria). 

METHODS 

Specimcnsof £A:n//£/£/t'/a from Riverslcigh rep- 
resent 30 individuals from several stratigraphic 
levels. The relative paucity of specimens and 
chronological data precludes a siriclly stjato- 
phenetic approach (yt'«.N7/Gingerich. 1976, 1979; 
Bown & Rose, 1987) to proplcopine phylogeny. 



However, a more general consideration of stratig- 
raphy in phylogenettc analysis may he appiopri 
ate in association with cladislic Irealmenl where 
specimens are siratigraphically disjunct or 
sparsely distributed (Gingerich. 1990). 

Sites with Ekaltadeta are late Oligocene to 
early late Miocene (Archer et a!., 1989, 1994. 
1995). A number of characters were analysed to 
assess the development of time-dependent 
changes. Specitnens were ranked to indicate rel- 
ative age (Appendix 1). Stratigraphic levels ate 
from Archer cl al. (1989, 1995): level I=lale 
Oligocene early Miocene; level 2=early 
Miocene: level 3=lalc early Miocene; level 
4=mid Miocene: level 5=latc mid Miocene; level 
6=early late Miocene; level 7=PIioccnc: level 
8=Pleistocene. 

I included all propleopines possible, although 
Pliocene and Pleistocene Jackinahoncya and 
Propleopus are known frorti inaierial often llnv 
ned to portions of upper andyor lower dcnlilions. 
Most Propleopus are from the Pleistocene, al- 
though material has been recorded from early 
Pliocene local faunas (Archer & Flanncry. 1985). 
Propleopus clullai;ocnsis was described as 
Pleistocene (Archer et al. 1985), but could be 
older, possibly late Miocene or early Pliocene 
(Archer pcrs. comm.). Jackiuahomya roxonien- 
sis is Pliticenc (Ride. 1993). 

Differences in molar gradient were used by 
Archer & Flannery (1985) and Wroe (19961 to 
di.slinguish proplcopine species. Molar gradient 
retlecis bulb the surface area and length t»f the 
molar lotith row. In proplet»pines a high molar 
gnidienteorrclalcs with areduclitm in hoih molar 
surlacc area and the \cT\gih of ihe lomh row. 



450 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Reducing the distance between 
condyle and sectorial loolh 
ma\imi/.e.s leverage applicable 
10 ihe tooth (Young el al., 
1989). Through shortening the 
molar row. leverage on the 
large shearing PVa of pro- 
plcopincs is increased. This ef- 
fect is achieved ai the cost of 
molar length. 

Relative P^ size and molar 
gradient for upper and lower 
dentitions has been quantified. 
Distinct reduction in tooth size 
posteriorly occurs in upper and 
lower dentitions of E, ima. In 
the upper dentition this steep 
gradient begins with a reduced 
posterior width (pw ) relative to 
the anterior width (aw) of M- 
which then ramifies through 
M^"^. In E. ima M"^ pw is <l/2 
M- aw. The upper dentition of 
P. cliiliagoensis is similar to 
that of E. ima. Lower dentition 
is not known for P. chUlagoen- 
sis. For P, oscillans M-^"* are 
missing but M-^ pw is only 
slightly less than M- aw sug- 
gesting a less extreme gradient 
This supposition is strongly 
supported by the lower denti- 
tion in which molar gradient 
contrasts strongly with E. ima, 
Mi-4 looth widths decrease 
steadily anteroposienorly in E. 
ima hut are reversed in P. os- 
cillans where loolh width in- 
creases posteriorly for M|.3» 
with only a slight decrease in M4. 

Several methods to quantify molar gradient 
have been considered. Accurate determination of 
individual molar surface areas and comparisons 
between leeth would be useful but would require 
2 or more teeth/ specimen, greatly limiting data 
sets, particularly for upper dentitions. Molar gra- 
dient might also be estimated geometrically by 
determining the angle al which a line drawn bucc- 
ally or lingually through the faces of the crown 
intersects the mid-line of the dentary or skull. 

In this study the clear initiation of a marked 
molar gradient at M- in the upper dentitions o\E. 
ima and P. cliiliagoensis permitted estimation of 
the gradient from a single molar by comparing aw 
to pw, In lower deniilions the gradient is less 




CS3."^'^:tV 



FIG. 1. Ekalladeta ima. x 2. A, occlusal view of QMF12436 luppers). B, 
buccal view of QMFr2435, left dentary containing Ii , alveolus for I:, P2-^, 
M1-4. C. occlusal view of QMFl 2435. 



distinct and 2 molars were required to demonstr- 
ate a gradient. Measurements were made using a 
Wild MMS 235 Digital Length-Measuring Set 
attached to a Wild M5A Stereotnicroscope. Ab- 
breviations are: I=lenglh, w=width. aw=anierior 
widlh, pw=poslerior width, dd=depih of deniary, 
G-value=ratioofanteriorlo posterior tooth width. 

RESULTS 



M' aw / M- pw VS STRATIGRAPHIC LEVEL. 
(Fig. 6). For upper dentitions the ratio M- aw: M- 
pw (G-value) was used as an arbitrary measure of 
molar gradient, with M- being common to the 
largest number of specimens. 



STRATIGRAPHY AND EKALTADETA 



451 



*v 







r^ 



FIG. 2. Propleopuschillagoensis, x 2. A, occlusal view 
of NMV P 1 59 1 7, right maxillary fragment (juvenile). 
containing unerupted P^, partial M^ M"^'-\ partial M*^ 
(cast of holotype). B, Propleopus oscillans, x 2, oc- 
clusal view of OMF6675, left maxillary fragment, 
containing P^, IVP"^. 

A trend is apparent in this scatter graph of 
G- value against stratigraphic level. P. chillago- 
ensis and P. oscillans represent 2 extremes with 
G-values of 1.23 and 1.06 respectively, with the 
lower number indicating a lesser molar gradient. 
Ekaltadeta ima from levels 3 and 4 has a limited 
range of G-values ( 1 .09- 1 . 1 5). 

The 2 Ekaltadeta from level 6 both fell outside 
the range of E. ima from older strata. E. 
jamiemulvaneyi (QMF24212; Cleft of Ages 4 
Site) had a low G-value of 1 .05, slightly less than 
that of P. oscillans. E. ima (QMF2421 1; Henk's 
Hollow Site) had a relatively high G-value of 1 . 1 9 
approaching that of P. chillagoensis. These re- 
sults indicate a divergence in the Ekaltadeta lin- 
eage with one population leading to P. oscillans 
and another leading to P. chillagoensis. 

Ml pw / M2 pw VS STRATIGRAPHIC LEVEL. 
(Fig. 7). The molar gradient of the dentary was 
estimated by dividing Mi pw by M2 pw (G- 
value). P. oscillans had the lowest G-value at 
0.93. The G-values for P. wellingtonensis and / 
toxoniensis were slightly higher at 0.96. At levels 
3 and 4 the G-values for Ekaltadeta were I.Ol- 
1.08. The G-value for E. jamiemulvaneyi, from 
level 6 (QMF24200, Encore Site) was 0.97. This 
placed Eyaw/emw/vfltt^y/ about halfway between 
the lowest G-value from levels 3 and 4 and P. 
oscillans. Again the highest degree of divergence 
among Ekaltadeta was for the E. jamiemulvaneyi 
from level 6, possibly indicating a trend toward 



K 










-8 








-S 



K 

s 




g 


"4* 

1 




£:■ 




^ 




Go 


1 


:3 
.1 


'§ 




1 


^ 


^ 


^ 


o: 


a: 



FIG. 3. Cladogram for the propleopines from Archer& 
Flannery ( 1 985 ). Character states at nodes: A=gain of 
an anterior cristid emanating from the melaconid of 
Ml, gain of derived Ii morphology; B=incorporation 
of the protolophid into the anterior iophid of M 1 loss 
of P2 with eruption of P3, dentary deeper anteriorly 
than posteriorly; C=reduction of melacone/en- 
toconid, P3 hypertrophy. 

the species with low molar gradients (/ toxonien- 
sis, P. oscillans and P. wellingtonensis). 

P3 w/Mipw VS STRATIGRAPHIC LEVEL. 
(Fig. 8). In P. oscillans P3 width was small com- 
pared to Ml posterior width (1.09). For J. tox- 
oniensis relative P3 width was greater (1.27), E. 
ima from levels 3 and 4 had ratios of P3 w / Mi 
pw of 1 .35-1.52. E. jamiemulvaneyi from Encore 



452 



MEMOIRS OF THE QUEENSLAND MUSEUM 



silc (QMF24200) again posi- 
tioned between E. ima from 
lower strata and J, toxoniensis 
I P. oscillans, with a ratio of 
L28 

DEPTH OF DENTARY 
AGAINST STRATI- 

GRAPHIC LEVEL. Depth of 
dentary against slraligraphic 
level (Fig. 9). Dentary depth 
was measured from the alveo- 
lar margin of Mi to the ventral 
margin of the dentary perpen- 
dicular to the m<.)lar row. Vari- 
ation in the depth of dentaries 
was small for E. ima from lev- 
els 3 and 4 (19, 3-21.1 mm). £. 
jamietnulvaneyi (QMF24200) 
was much larger with a dentary 
depth of 28.8mm approaching 
P- ascillans (32.6mm). J. tax- 
oniensis between E. ima and £. 
janiiemulvancyi/P. oscillans 
with a dentary depth of 
23.3mm, 







t * 



^^ 






« .1 



4 

7^^ 



1 



STATISTICAL ANALYSIS. 

(Table I). Because all 
EkalUideUi material has come 
from a relatively small area 
(Rivcrsleigh), a regional popu- 
lation of potoroids was consid- 
ered an appropriate control. 
Sixteen specimens from the 
Australian museum of Potor- 
o u s tridac t y / u s collected 
around Hobarl were used, this 
being the largest potoroid spec- 
imen satnple available. Varia- 
lion in the G-values of 
Ekaltadda from levels 3 and 4 
approached that of P. 
rridacty'lus. When G -values 
from the 2 Ekalfadeta from 
level 6 were included the variation fell well out- 
side that of the local P. rnV/«rrv/av population. 

DISCUSSION 



Increases in premolar and molar shear within 
the Propleopinae appear to be mutually exclusive 
and their relative importance probably rellects 
dietary preference. A requirement for high pre- 
molar shear might be associated with camivory 






I' 



8 






tik 



Th 



ii¥ 



T 



FIG. 4. Ekaltadetajamiemtdvaneyi x 2. A, occlusal view orQMF24200, left 
dentary containing P^, M|-^. holoiype. B. buccal view of QMF242(X). C. 
lingual view of QMF2421X). D, occlusal view of QMF242 1 2. left maxillary 
fragment, containing P-, dP^, M'-, referred specimen. E, buccal view of 
QMF24212. F. lingual view of QMF24212. G. occlusal view of 
QMF20842. left P\ referred specimen. H. buccal view of QMF20842. I. 
lingua! view of QMF2()S49. 



(Abbie, 1939), while a more extensive molar 
array may indicate a more herbivorous diet 
(Wells ctal.. 1982). 

Species with a large molar surface area and low 
molar gradient (P. oscillans, P. wellingionensiSf 
J. toxoniensis) have relatively small premolars. 
Species with high molar gradients and reduced 
molar shear (£. ima, P. chillagoensis) are 
characterised by P3 hypertrophy. The extraordi- 
nary change in function for P2 shown by individ- 



STRATEGRAPHY AND EKALTADETA 



4?3 



8- 

i 



TABLE I Statistical summaries for M- aw / M- pw 
(G-\'aliic) for propleopines and a local P. tridacryhis 

population. 



FIG. 5. Minimal tree produced by Wagner analysis for 
the Propleopinae (from Wroe, ! 996). Character states 
at nodes; A = gain of an anterior cristid emanating 
from the mctaconid of Mi; basally broad conical 
upper molars; B = presence of lingual cingula on the 
upper molars; C = reduced molar gradient, reduced 
P3; D = mcorporaiion of Uie protolophid into the 
anterior lophid of Mj, a dentary deeper posteriorly 
than anteriorly. 

ual £". ima (Wroe & Archer, 1995) probably con- 
stitutes a response to the increased loading placed 
on P3. Regarding molar gradient and relative si/x- 
of the P3, EJamiemuJvcmeyi is intennediate, fall- 
ing between E. ima specimens from lower levels 
and P, oscilUms/P. wellingTonensis/J. toxomen- 
sis. Using the same criteria / roxoniensis lies 
between E. jamiewulvaneyi and P. osvillanslP. 
wellingtonensis. In terms of variation in P3 size 
and molar gradient P chillagoensis and P. os- 
cillans represent opposite extremes in propleop- 
inc evolution and it is suggested that P. oscUkms 





N 


SD 


cv 


SE 


Propleopines to level 8 


13 


0.05 


4.53 


0.01 


EkaluiJeta lo level 6 


11 


0.38 


3.35 


0.01 


EkulititJeto 10 level 4 


9 


0.02 


2.07 


7.71 E-3 


P. indacnhis 


16 


0.02 


1.70 


4.61 E-3 1 



was largely if not wholly herbivorous. Other de- 
rived features interpreted as adaptations lo 
herbivory for P. oscillans include a large dia- 
stema between Pi and li, and large spaluiale 
lower mcisors (Wroe, 1996). Regarding dcniary 
depth E. ima is the smallest propleopine with a 
general increase in depth for laxa at higher strati- 
graphic levels probably rctlecting a general in- 
crease in body size. 

Siraligraphic and metric analysis support the 
proposal of a laie Miocene dichotomy in 
Ekaltadeta producing 2 lineages of Propleopus^ 
and a reversal of previous assumptions on relative 
apomorphy within Propleopus, with P. osdUans 
considered Ihe most derived and P. chillagoensis 
the most plcsiomorphic (Wroe, 1996). However, 
broad trends suggested in ihis study are not inier- 
prelcd here chronoclines in the strait>phenelic 
sense (sensu Down & Rose, 1987), The scarcity 
of material and uncertain chronology o\ both the 
Oligo-Miocene Rivcrslcigh deposits and the Plio- 
Pleislocene local faunas from which most pro- 
pleopine specimens are known necessitates 
caution in the interpretation of results. A consid- 
erable temporal gap exists between estimated 
ages of the most recent Ekaltadeta specimens and 
all other propleopines. As noted by Ride ( 1993). 
the period separating the latest known incidence 
of Ekaltadeta frtmi Plio-Plcistoccnc Propleopus 
and Jackmahoneya may be sufficient to have 
permitted a secondary reversal of character stales 
within Propleopus lo produce P. cliillagoensis. 

Many questions remain concerning the age, 
stratigraphy and method of deposition of 
Riversleigh's Oligocene-Miocene limestone de- 
posits (Archer. 1994, 1995; Megirian, 1992, 
1994). If the phylogeny for propleopines sug- 
gested by Wroe (1996) reflects evolutionary 
events, then it provides tacit support for Archer 
ct al.'s (1989) proposed stratigraphy, with an 
agreement of hypothesised superposilional and 
phylogenetic patterns. 

The capacity of stratigraphic occurrence to ex- 
plicitly mirror phylogenies is questionable ifin- 
gclmann & Wiley, 1977). Although strong 



454 



MEMOIRS OF THE QUEENSLAND MUSEUM 



f ^ 



10 



/? oscUUms 
a 

QMF(W75 

E. jamiemulvaneyt 
n yMF2J2n 



F chilla^oeruiis 
a 

NMV P15917 



E. ima 

DQMF243I! 



D o CD £• inia iscc caption for m.i) 



T.O 



1,1 1.; 

M2 aw / M2 pw 



1.3 



P. osciUans 

QM PiiOZ 

a 


a 


P. weilingionensis 
ucMPJsni 




o 


X lo.wniensis 






D E. jainiemulvaneyi 

gM F243O0 






£. fmfl QMH12423 

a 


E, ima 


QM FU4i5 QMPZ4I98 QM R4I9« QM P:4197 
□ □ Q a 






DD 






E. ima UM FM!<)6 yM P:4195 






, ■ 1 



0.9 



1.0 
Ml pvv / M2 pw 



1.1 



FIG. 6. M" aw / M- pw vs relative straiigraphic level FIG. 7. Mi pw / M2 pw vs relative straligraphic level 
for propleopines. Ekaltadefa ima from level 3, left to for propleopines. 
right, QMF24207, 24204, 24205, 12436, 24203, 
24208, 24209, and 24206. 



lOi 



4- 



10 



P osciUans 

° OM"'o^ Itaxonimsis 

□ AR 175W 



E. Jamie mulvaneyi 
yM i^;42<Hi 



OMin2424 qmpi24:j E, ima 
□ □ 



QMFI2435 E.iim 

QM F242(H £. ima 



^^ 1.2 1.3 1-4 l.S 

P3\v/Ml pw 



1.6 



FIG. 8. P3 w / Ml pw vs relative straligraphic level for 
propleopines. 

congruence between cladistic and stratigraphic 
arrangements has been demonstrated for many 
venebrate taxa by Noreil & Novacek (1992a,b), 
the same authors advised that correlation between 
the 2 diminishes rapidly where cladislic or strali- 
graphic data is poorly resolved. Debate over con- 
formity of age and cladislic information 
commonly centres around the value of super- 
positional data as an adjunct to phylogenetic re- 
construction. Where cladislic analysis is sound it 



i 6 









F oxciUanx 


J. joxonietuis 


n QM FiiOl 


AK 17579 






E. jamiemulvaneyt 


D 


QM P24200 


a □ QM ii::424 yM fU4>j E. inui 


a QM 1=124-^5 E. ima 




oQM p\2?A?M E. ima 

, T 1 


— 1 ■ [ 



10 



20 30 

Deniary Depth (mm> 



40 



FIG. 9. Depth of dentary vs relative straligraphic level 
for propleopines. 

may be useful as a test of straligraphic interpreta- 
tions. 

The propleopine phylogeny of Wroe (1996) is 
based on analysis of an incomplete data matrix, 
with important characters unknown for several 
species. Consequently the cladistic data pre- 
sented cannot be viewed as a robust basis for 
testing superpositional pattern. However, the pro- 
ductivity of the Oligocene-Miocene deposits of 



STRATIGRAPHY AND FKALTADETA 



455 



Rivcrslcigh engenders reasonable cxpecUition lor 
the reliable resolution of phylogcnics. 

ACKNOWLEDGEMENTS 

M. Archer. H. Godihclp and W. D. L. Ride 
provided conslruclivc criticism and cominenl. 
Vilal support lor this research has been given by 
ihc Australian Research Council (toM. Archer), 
the National Estate Grants Scheme (Oaeenslantl) 
(grants lo M. Archer and A. Harlholnmai); the 
Department of Environment, Spoils and Territo- 
ries; the Queensland National Parks and Wildlife 
Service; the Commonvcealih World Heritage 
Unit (Canberra); the University of New South 
Wales; ICI Australia Ply Ltd; the Australian Cico- 
graphic Society, ihe Queensland Museum; the 
Austiiiliun Museuuj; Century Zinc Ply Ltd; Ml 
Isa Mtnes Piy Ltd; Surrey Bcatty & Soiis Pty Ltd; 
the Rivcrsleigh Society Inc.; the Royal Zooluj^i- 
C^\ Society of New South Wales; the Linncan 
Societ^y of New South Wales; and many private 
supporters. Skilled preparation of uiiist of the 
Rivcrsleigh material has been carried Out by 
Anna Gillespie 

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