Table Of ContentUiVliRSlTYAND BIOSTRATIGR.'VPH^' OF THE DIPROTODONTOIDEA OF
RTVERSLBIGH. NORTHWESTERN QUEENSLAND
KAREN BLACK
Black, K. !907;06:30: Divcrsilyandbiosiraligraphyu!theDiprotodonloideaolRiversleigh.
norihwtvsicmO'tccnslanci. Me(f\oinoftheQaeenslcmdMuseum 4H2): 187-192. Brisbane.
ISSN0079-8835.
The diversityunddistrihutionofRiversieigh's Dipruiodonloidcais 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'/cnandPropalorchesfvsT^wgtihruu^houiSysicniN A.BmidCexhibitiug;i tnorphocl-
ine. Stage-of-evoluliun biocoiTclation with oihcr Ausiralian and New Giunean Ternary
mummal I'aunas is inaccord with previouslye&iabllshed biosiiatigruphie hyputhcscs. How-
ever .some basal Riverslcigh deposits may predaic (he central Ao'^tralian Wipajiri and
EladunnaFnrmalions.Diproiodontoiddiversityis;high niSysieni Adeposus.low iiiSystem
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.
,
KarenBlacKSchoolofBiologicalScience. Universityc^NewSoHihWtt/es,NewSouih HWA'.v
2052,Australia:received4November 1996.
ThepaucityorradionictricanydatedAustralian Liihostraiigraphy and Systems terminology
Tcniaryfreshwater fossil deposits has resulted in follow Archerel al. ( 1989, I994al. Cusp nomen-
marsupial stagc-of-evolulion biochronology clature lolhms Archer (1984) and Rich el al.
being ihc most commonly employed method of (1978) Molar hontology ts (hat proposed hy
dating fossil assemblages (Wuodburne el al., LuckctH 1993k PicinolariiumbcrfollowsFlower
I'J85).FundametUal tothistnethodisanadequate (1867) Higher level systematic nomcnelaiurc
understanding ofphyiclic succession within the follows Aplin & Archer 1987).
(
ta\on (Megirian, 1994). Chronologic and phy-
letic succession within diproiodontoid lineages BIOSTR/XTIGRAPHY
arc well documented (e.g. Stirion et al., 1967;
Murray. 199Ub; Murray el al. 1993). Conse- Figure 1 lists the diprotodt)nlids and pal-
quently, the group has played a major role in orcheslids from different local faunas withm
previous biochronologic analyses (e.g. Stirton ct Riversieigh's siratigraphic units. These units are
al.. 1967: Woodbtirncei al.. 1985; Woodbumeel defined hy Archer et al (1989. 1994a): System
al., 1993; Archerctal., 1994; Archeretal., 1995) A, late Oligocene lo early Miocene: System B.
and hascontributed significantly totheconstruc- eariy to middle Miocene; System C. middle to
tion of the chronological framework of earlylaieMiocene;andPleistoceneassemblages.
Australia'sTertiary mammal faunas. Silvahistms wichaelhini is the most picsio-
Similarly^ because oflimited siratigraphicdata morphic zygomaiurinerecognised {with thepos-
and the lack of radiometric dates, much of the sible exceplion ofHaemeotheriinnyatkolai Rich
current chronologic sequence o\ Riversieigh's ci al-, 1978) and may be antecedent to (he entire
local faunas has been constructed on the basis of /ygomaturineradiation(Black&Archer, 1997a).
vertebrate biocc^rrclalion (Archer el al.. 1989), FourspeciesofNeohelosarerecognised(P. Mur-
Recent reappraisal of diprotodontoid material ray pcrs. comm). Neohelos sp. nov. I, a stnall
from Rivcrsleigh indicates at least 9 genera and picsiomorphic form; N. firarensis a medium-
K species. This paperoullines the diversity and sized moderately derived form from the
I
distribution of diprotodontoids throughout the Kuljamarpu Local Fauna, South Australia;
Riversleigh Systems and discusses their bearing Neohelossp.nov.2.alargederived formfromthe
on current biostratigraphic understanding, Bullock Creek Local Fauna. NT. anieccdcnt lo
tiioc(irrelatu:m of Riversieigh's Oligocene- Kolopsis torus (Woodburne, 1967) o\' the laic
Miocene deposits with other Australian fossil MioceneAlcotitaLocalFauna.NT;andNeohelos
assemblages is alsodiscussed, sp- nt)v. 3, a highly derived form structurally
—
188 MEMOIRS OFTHEQUEENSLAND MUSEUM
SYSTEV A SYSTEM B SYSTEM C PL
L MID H LOW MID HIGH C+ El
LOCALFAUNAS D B H RB BS AJ IVP AL WH WW CS NG DT MM Ni TR G DI JJS HH JJ 90 CJ DS C NE TE
S. michaelbirti 1
S.johnnilandi 2
Silvabestiussp. 1 10
Neohetossp. nov. 1 1 1 1 1
N. tirarensis 1 1 1 I 1 1 1 1
Neohelossp. nov. 2 h— I 4
Neohelossp. nov. 3 2
Ni. lavarackorum 1 1 1 24 1 2
Nifiibadonsp. 1
Zygomaturine
9
gen. nov.
B.angulaiu/n 1 1 2 4 1 3
Bemaiheriumsp. 1
Ngapakaldiasp. I
Prporiticulus I 1 2 1 1 1 1 I
Pr. novaciiia
cephalus 1 I
Palorchesiesanulus 1
P. Qzael 1
Diprotodonopiaiuiu 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.ThedistributionofdiprotodontoidtaxathroughtheRiversieighsequence.Numbersindicatetheminimum
numberofindividuals foreach species found in a given local fauna. PLEI=Pleistocene. System B heading is
subdivided into: L=Low, MID=Middle and H=HighLocal FaunaAbbreviations: AL, Alsite; 90, Alan's Ledge
1990; BO, Burnt Offering; BR, Bone Reef; CO, Cleft ofAges (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'sCarousel;JJ, Jaw
Junction; JJS, Jim's Jaw Site; MM, Mike's Menagerie; NG, Neville's Garden; RT, Ringtail Site; SB, Sticky
BeakSite; TE, Terrace Site: WH, White Hunter; VIP. VIPSite; WW, Wayne's Wok.
transitional between Neohelos sp. nov. 2 and Murray pers. comm.) with reduction or loss of
Kolopsisyperns(Murrayetal, 1993)fromthe late premolar cusps a relatively common phenome-
MioceneOngevaLocalFauna,WaiteFormation. non. Comparable cranial material for Ni. lav-
Bematherium sp. is known from a single maxil- arackorum has yet to be processed. Propalor
lary fragment which is plesiomorphic relative to chestes ponticulus (Murray, 1990b) is plesio-
B. angiilum. A new unnamed species of morphic relative to P. novaculacephalus. Pro-
Ngapakaldia is more derived than the central palorchestes novaculacephalus occurs in the
AustralianN. tedfordibutplesiomorphic relative Bullock Creek Local Fauna; Murray (1990b) in-
to N. bonythoni. Fragmentary material of dicated thatthis material is morederived than the
Zygomaturine gen. nov. has an uncertain phylo- Riversieigh specimens. Palorchestes anulus is
genetic position. Nimbadon sp. is similar to N. intermediate between Pr. novaculacephalus and
lavarackoruminsize,molarmorphologyandcra- Palorchestes painei from the late Miocene Al-
nial profile; however, differences in upper pre- coota Local Fauna (Black, 1997). Palorchestes
molar morphology may imply a more azaeland Diprotodonoptatum the mostderived
,
plesiomorphic position within the genus. How- membersofPalorchestidaeandDiprotodontidae.
ever, I doubt whether this single skull is indica- respectively,areknown fromTerraceSite, which
tive of a new species. Extreme intraspecific has been radiocarbon dated at approximately
variation in P-^ morphology is common among 23,900-F/- 4,100-2,700 years BP (Davis &
Tertiary zygomaturines (e.g. Neohelos spp.; P. Archer, 1997).
DIVERSITYOFDIPROTODONTIDS AND PALORCHESTIDS 189
DISCUSSION al, 1993). Murray (pers. comm.) suggests thai if
this material is found to represent tirarensisthen
AGREEMENT WITH PROPOSED STRATIG- someofRiversleigh'solderdeposits maypredate
RAPHY. The dislribution of diprolodonle^ids mammal-bearing Etadunna Formation.
throughout the Riversleigh systems is generally Myers & Archer (1997) correlated White
consistent with Archer et al/s (1989, 1994a, Hunter Site (System A) and MammaUm Hill
1995) proposed stratigraphic framework. The (Zone D. Etadunnal Fomiation). the laller 24.7-
most plesiomorphic forms are restricted to the 25 MY BP (Woodburne et al.. 1993).
older System A local faunas, and more derived INgapakaldiasp. nov. in SystemAextendsthe
taxa become moreabundant in Systems B and C. generic distribution from South Australia. This
A similar pattern oi' distribution is evident species appears to be more derived than N.
within lineages. There is a gradual evolution in tedfordi from the Ngapakaldi and Ngama Local
cranial and dental morphology accompanied by Faunas (Etadunna Fmn) and the Tarkarooloo
an increase in body size within Neohelos and Local Fauna (Namba Fmn), yet plesiomorphic
PropalorchesTes. respectively, through the relative to N. honythoni from the Ngapakaldi
Riversleigh sequence (Murray, 1990b; pcrs. Local Fauna. Ngapakaldia also occurs in faunal
comm.). Although some temporal overlap is evi- zones C. D and E ofthe Etadunna Fmn. As with
dent, generally Neohelos sp. nov. 1 is the domi- Neohelos, species level identification is neces-
nant form in System A, N. tirarensis is most sary before Ngapakatdia will be o^ use in
abundant in System B.Neohelos sp. nov. 2 spans biocorrelation
low to high System C and Neohelos sp. nov. 3 is Propalorchesies novaculacephalus and
unique to high System C Jaw Junction Site. Neohelos sp. nov. 2 in low to mid System C
A similar succession is exhibited by pal- assemblages confirms previous hvpotheses
orchestids. Plesiomoq")hic Propalorchesiespon- (Archeretal., 1989; 1994a; 1995) that'theyareof
ticulus is relatively common in System A and B a similar age to the Bullock Creek Local Fauna
Sites, and is succeeded by the more derived Pr. (Fig. 2). Correlation is further supported by the
novaeuiacephalus in mid-high System C Sites. shared Nimbacinns dicksoni (Muirhead &.
Archer, 1990), Wakaleo vanderleiiri (Munay &
INTRACONTINENTALCOMPARISONS.The Megirian, 1990) and Balharoo sp. (Flanncry et
Riversleighassemblagescontainthemostdiverse al., 1983).
array ofdiproiodontids and palorchestids ofany Neohelos sp. nov. 3, the largest and most de-
single region on the continent. Of the 9 genera, rived species oi' Neohelos, occurs in the Jaw
Neohelos, NimbadofU Ngapakatdia, Propalor- Junction assemblage suggesting that this high
chesiesandPalorchesfesarealsoknown fromthe System C deposit is younger than the Bullock
Oligocene-Miocene in central and northern Aus- Creek Local Fauna. Neohelos sp. nov. 3 exhibits
tralia. The biochronological potential ofsome of an upper third premolar morphology that antici-
these genera is, however, limited, because of pates the condition ftiund in the more derived
uncertainty about interspecific afi'inilies. zygomalurineKolopsis, which firstappears in the
Neohelos, PropalorchesiesandPalorchesfes. are Alcoota and Ongeva Local Faunas of the Waile
biochronologically most significant. Fmn, Northern Territory. Ancestor-desccndenl
Neohelos tirarensis from System B is close to relationships are well supported for the
the type material from the Kutjamarpu Local Neohelos!KolopsislZygomatunis clade (Stirton
Fauna suggesting age equivalence; this is sup- etal., 1967; Murray etaL, 1993). The presenceof
ported by other shared taxa such as Wakiewxikie Neohelos material, structurally transitional be-
/^/H'iYva(Godthelpclal.. 1989). Pa//V/ra(Archer, tween Bullock Creek's N. sp. nov. 2 and (he
1994), V/akaleo oldfieldi (A. Gillespie pers. WaiteFormation's Kolopsis furthersuggests this
comm.), Namilamadeta: Namharoo (Archer ct correlation making some of Riversleigh's high
al., 1989) and Litokoala (Black & Archer. System C deposits, such as Jaw Junction Site,
1997b). youngerthan the Bullock Creek Local Fauna but
Neohelos sp. nov. I in Systems A-C is more older than late Miocene deposits ol' the Waile
plesiomorphic than N, tirarensis suggesting thai Formation.
someoi'Rivcrsleigh's stratigraphic units predate ThisisalsotrueofRiversleigh'sEncoreassem-
theKutjamarpuLocal Fauna.Neohelos from fau- blage which, on the basis ofits derived and often
nal /ones D-E of the Etadunna Formation have unique fauna, is believed tobeearly lateMiocene
not been specifically identified (Woodburnc et in age (Archer et al., 1995). This is based on the
190 MEMOIRS OFTHEQUEENSLAND MUSEUM
LATE
MIOCENE ALCOOTA
ENCORE
10.4
MY
MID
MIOCENE BULLOCK SYSTEM
C^
CREEK
16.3
MY
FARiA
vuo< em:
SYSTEM B
KUTJAMARPU
23.3
MY
NGAMA TARKAROOLOO SYSTEM A
LAIE
NGAPAK.ALDI
Ol IGOCENE
ERICMAS
DiTJIMANKA
PINPA
SOUTH AUSTRALIA NORTHERN RIVERSLEIGH
TERRITORY 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
diprotodonloidslage-of-cvolulioncomparisons.
presence of several taxa which exhibit adapta- cies. Conversely, upper System C deposits ex-
tions to the onsetol drierclimatesduring the late hibit a decline in diprotodontoid diversity. Ifwe
Miocene as well as taxa whose lineages extend considerthesmall numberoftaxabeinganalysed
into Pliocene and Pleistocene times. These in- suchchangesindiprotodontoiddiversity maynot
clude: Palorchestes aniilus which is structurally besignificant. However, asimilardecline in both
antecedent to P. painei from the late Miocene family-andgeneric-level diversity isevident in a
Alcoota Local Fauna (Black, 1997); a small number of other marsupial groups during the
Phascolarctos\ahypselodontvombalidrelatedto middle to late Miocene. This decline may be
the late Cainozoic Warendja wakefielcli (Archer related to the late Miocene onset of 'icehouse'
et al., 1995); a thylacoieonid intermediate be- climatic conditions resulting in the regional col-
tween Wakaleovanderleuriandthe lateMiocene lapse ofrainforestandsubsequent spreadofopen
Wakaleo alcootense (A. Gillespie pers. comm.); forest and woodland/savanna (Archer et al.,
a giant rat kangaroo, Ekaltadetajamiemulvanei 1994b; 1995).
(Wroe, 1996), which is possibly annecetant to Replacementofrainforesthabitatbymoreopen
Pleistocene Pmpleopus; and Mayigriphus forest may have benefited select diprotodontoids
orbusCWroe. 1997), a dasyuromorphian with which is reflected in an increase in species abun-
some featurescorrelatedwithdrierenvironments dance for some System C Local Faunas. Rela-
in moderndasyurids. tively high 'minimum number of individuals'
estimates for species o[Nimbadon and Neohelos
DIVERSITY AND FAUNAL CHANGE. Sys- sp. nov. 1 have been recorded from AL90 and
tem A sites contain the highest number of con- Cleft of Ages Sites respectively. This may sug-
temporaneous diprotodonloids with 5 species at gest that individuals ofthese species were roam-
SiteDand4atStickyBeakSite(Figs 1-2).Adrop ing in mobs, a feature characteristic of slow
in diversity is evident in System B withonly 1 or moving medium- tolarge-sizedherbivoresinrel-
2 species per site. Diversity increases slightly in atively open environments.
low to middle System C assemblageswithgener- It is also feasible that high abundance in the
ally 1-2 contemporaneous diprotodontoid spe- above System C assemblages, and conversly low
DIVERSITY OFDIPROTODONTIDS AND PALORCHESTIDS I9J
abundancein SystemA-B assemblages, mayrep- and Territories; the Queensland National Parks
resent sampling orlaphonomic biases. Likewise, and Wildlife Service; the Commonwealth World
the absence ofdiprotodonlines in System B and Heritage Unit; ICI Australia Ply Lid; ihe Austra-
C may be an artefact of incorrect taxonomic lian Geographic Society; the Queensland Mu-
assignment. Even so, several zygomaturine spe- seum; the Australian Museum; ihe Royal
cies recovered from System B and System C Zoological Society of New South Wales; Ihe
assemblages appearto have developed a number Linnean Society of New South Wales; Century
ofdiprotodontine- like featuresoftheirdentition. Zinc Ply Ltd; the Riversleigh Society Inc.; and
These include the reduction ofthe paraslyle and private supporters including Elaine Clark. Mar-
loss of the hypocone on P-' and a reduced garet Beavis, Martin Dickson, Sue & Jim Lav-
paracristid on Mi. A similar phenomenon has arack and Sue & Don Scotl-Orr. Vital assistance
occurredintheAlcooiaLocal Faunawheredental in the field has come from many hundreds of
and cranial morphology ofthe zygomaturineAl- volunteers as well as staffand postgraduate stu-
kwertathehumwebbi is convergent on that ofthe dents of the University of New South Wales.
diprolodontinc Pynunios alcootense (Murray, Skilled preparation of most of the Riversleigh
1990a). In general, zygomaturines display a material has been carried out by Anna Gillespie.
higherdiversity and abundance in Tertiary fossil
assemblages than diprotodonlines. This may re- LITERATURE CITED
fiect their greater ability to adapt to changing
environments than their diprolodontinc counter- APLIN, K. & ARCHER, M. 1987. Recem advances in
parts. It is further rellected in Rivcrslcigh's Sys- marsupial sysicmaiics with a new syncretic clas-
tem B and System C Local Faunas, where sificalion. Pp. xv-lxxii. In Archer. M. (cd.). Pos-
diversifying zygomaturines have subsequently sumsandopossums: studies inevolution. (Surrey
radiated into vacant niches occupied by Beatty &. Sons & Royal Zoological Society of
diprotodontines during the late Oligocene. New South Wales: Sydney).
Diprolodontoids are not known from ARCHER. M. 1984. The Australian marsupial radia-
Rivcrslcigh's Pliocene assemblages. They seem tion. Pp. 633-808. In Archer, M. & Claylon, G.
to have declined markedly in generic diversity (eds). Vertebrate zoogeography and evolution in
Australasia. (Hesperian Press: Perth).
throughout the Australian Pliocene. Significant 1994. The ringtail possums (Marsupiulia,
faunal turnover is characteristic of most marsu- Pseudochciridae) of Riversleigh. Riversleigh
pial families during the late Miocene and early Symposium Abstracts, p.24-25.
Pliocene (Archeret al., 1995). ARCHER, M.. GODTHELP, H., HAND. S.J. &
The drop in diversity ofdiprotodontid and pal- MEGIRIAN. D. 1989. Fossil mammals of
orchestid species continues into the Pleistocene, Riversleigh, northwestern Queensland: prelitni-
withonlyonememberofeach familyrepresented riiiry overview ofbiosiratigraphy, correlation and
(Fig. 1). Both species are common throughout environmental change. The Australian Zoologist
aAruestirhaelilaa'ssl oQfuattheeirrnarresypefcotssiivledleipnoesaigtess,. Tyehtisbodteh- ARCH2E5R:3.5-M6.9..HAND,S.J. & GODTHELP. H. 1994a.
cline in diprotodonloiddiversity may be aconse- 199R4ibv.ersPlaetitgehr.ns2riindtehde.ihiRseteodryBooofkAsu:stSryadlnieay')s.mam-
quence ofunsuccessful competition with rapidly mals and inferences about palacohabitals. Pp.
diversifying mesic and xeric macropodoids 80-103. InHill, R.S. (ed.). Historyofthe Austra-
(Archerel al.. 1994). lian vegetation: Cretaceous lo Recent. (Cam-
bridge University Press: Cambridge).
ACKNOWLEDGEMENTS
1995. Tertiary environmental and biotic change in
Australia. Pp.77-90. InVrba.E.S..Denton.G.H..
I wish to thank Mike Archer and Peter Murray Partridge. T.C. & BurcUc, L.H. (eds), Palaeocli-
whocritically read a draft ofthis manuscripi and maie and evolution with emphasis on human
Anna Gillespie. Henk Godthelp and Steve Wroe origins. (Yale University Press: New Haven).
BLACK, K. 1997. A new species of Palorchestidae
forihcirassistance with taxon correlations. Vilal
supportforresearchatRiversleighhascomefrom (MMiaorcseunpiealEinac)orferoLomcatlheFalautnea,miRdidvlereslteoigeha,rlnyorltahl-e
the Australian Research Grant Scheme (to M. western Queensland. MemoirsoftheQueensland
Archer); the National Estate Grants Scheme Museum41: 181-185.
(Queensland)(toM.ArcherandA.Bariholomai); BLACK, K. & ARCHER. M. 1997a. Silvahesiius gen.
the University of New South Wales; the Com- nov. a primitive zygomaturine (Marsupialia,
monwealth Department of Environment, Sports Diprotodonlidae) from Riversleigh, northwestern
.
192 MEMOIRS OFTHEQUEENSLAND MUSEUM
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41: 193-208. iwvacidacephahis Murray and P. ponticidiis sp.
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