Palorchestes selestiae, a new species of palorchestid marsupial from the Early Pliocene Bluff Downs Local Fauna, northeastern Queensland

7 Pages·1995·2.4 MB·English

PALORCHESTESSELESTIAE, A NEW SPECIES OFPALORCHESTID MARSUPIAL
FROMTHE EARLYPLIOCENEBLUFFDOWNS LOCALFAUNA, NORTHEASTERN
QUEENSLAND
BRIANMACKNESS
Mackness,B.S. 1995 1201.Palorchestesselestiae,anewspeciesofpalorchestidmarsupial
fromtheearlyPlioceneBluffDownsLocalFauna,northeasternQueensland.Memoirsofthe
QueenslandMuseum38(2): 603-609. ISSN0079-8835
Palorchestesselestiaesp.nov.fromtheearlyPlioceneBluffDowtuLocalFaunaisdescribed
onthebastsofan isolatedLMKItislargerthanP.patneiand P.parvusbutsmallerthan P.
azaelinbothlengthand widthoflophs. Itisalsodiffercntiable fromall otherpalorchestids
onthe basisofthecombination oflinksbetween,orstemming fromlophs. It is thesecond
undoubted Pliocene species and shares features ofboth P. parvus and the Pleistocene P.
azael.Withitsrelativelyderivedanteriormorphologyandplesiomorphicposteriormorphol-
ogy, P, selestiae challenges current notions about the phyiogenetic relationships of
diprotodontid marsupials which suggest that smalt size and simple structure are probably
pleisomorphicfeatures. Palorchestes,palorchestids,BluffDownsLocalFauna,Pliocene,
Diprotodontia,Marsupialia.
Brian Maekncss, SchoolofBiologicalSciences, UniversityofNewSouth Wales, P.O. Box
I. KensingtonNSW2033, Australia; IAugust J995.
PalorchesteswasdescribedbyOwen(1873)on salt. Archer, inArcherScWade(1976)noted,but
the basis of an anterior portion of a cranium didnotnamethreeotherdiprotodontoidsfromthe
includingtherostrum.Theholotypewasobtained Bluff Downs Local Fauna, a species of Eu-
by Dr Ludwig Becker and described by Owen ryzygoma, a species ofZygomaturus and an un-
(1874)asP.azaelfromanunspecifieddepositin identified *notothere\
Victoria interpreted by Mahoney & Ride (1975)
as the River Tambo in Gippsland. Owen Terminology
(1873:387)consideredPa/a/rrteifestobe".. .the Dental homology of cheekteeth follows
largestform ofkangaroohithertofound". Luckett (1993) such that the first adult molari-
Owen regarded Palorchestes to be a form tooth is Mi and the deciduous molariform
macropodid,aview followedbySimpson(1945) tooth isdp*. Terminology ofcrown morphology
who placed the genus within the subfamily (Fig. 1)followsthatusedbyArcher(1976)except
Macropodinae. Raven & Gregory (1946), how- that the metaconule is used instead of hypocone
ever,placeditinthesubfamilySthenurinaewhile following Tedford & Woodburne (1987). Tbe-
Tate (1948) placed it in anew subfamilyofkan- gotic terminology used follows Every (1972).
garoos, the Palorchestinae. Woods (1958) was Abbreviations for specimen numbers: QMF,
thefirsttorecognisethediprotodontidratherthan Queensland Museum fossil collection; UCMP,
macropodid affinities ofPalorchestes. Archer& UniversityofCalifornia,Berkeley;P,SouthAus-
Bartholomai(1978)followedthisassignmentbut tralian Museum.
elevatedTate's Palorchestinae tofamily level.
SYSTEMATICS
Palorchestes selestiae sp. nov. is described
LM
here on the basis of an isolated ! from the
fluviatile and lacustrine deposits of the Al- OrderDIPROTODONTIAOwen, 1866
lingham Formation, northwest ofCharters Tow- SuborderVOMBATIFORMES
ers, northeastern Queensland. This formation Woodburne, 1984
VOMBATOMORPH1A
containsadiverseassemblageknownastheBluff Infraorder
&
DownsLocal Fauna, and hasbeen the subjectof Aplin Archer, 1987
many studies, most recently including those of Family PALORCHESTIDAE(Tate, 1948)
&
Vickers-Rich (1991), Mackness et aL (1993), Archer Bartholomai (1978)
Boles & Mackness (1994) and Mackness (in PalorchestesOwen, 1873
press). Archer& Wade (1976) assigned anearly
Plioceneagetotheassemblageonthebasisofthe TypeSpecies
mtepreted age ofthe overlying Aliensleigh Ba- PalorchestesazaelOwen 1873.
1
604 MEMOIRS OFTHEQUEENSLANDMUSEUM
The anterior cingulum fails to extend around
the base of the protocone. A slight anterobuccal
cingulum is continuous with the anterior cingu-
lum. Otherwise there are no lingual or buccal
cingula.Anteriorcingulumhighresultingindeep
fossae where the forelinks join the cingulum.
Posterior cingulum closer to the metaloph than
the anterior cingulum is to the protoloph. As a
result, the posterior cingular valleys are not as
deepastheanteriorones. There isaconspicuous
interproximalwearfacetontheanterobuccalcor-
ner ofthe tooth, presumably caused by abrasion
againstP^.
There are a number of blades on the crown.
M Where the enamel has been breached, both sets
FIsaeGrle.esa1tl.isaDoei.parRgeersgaeimnotnsesdhouuwssieindnggfottrehermmehiaonslouolrtoeygpmyeeon(ftQsaMnFin12Ta41b5ol5fe)P..1 ocfanlebaedinsgeeanndontratihleinpgrboltaodleosp(hseannsduEmveetrayl,op1h9.72A)
Arrowindicatesanteriorofmolar.Abbreviations:ab.f numberofsecondarybladesarealsopresent.One
anterobuccalfossa;a.c. anteriorcingulum; l.f. lingual blade (the apical margin ofthe lingual forelink)
forelink; m. metaloph; me. metacone; mcl. leads from the protocone to the anterior cingu-
metaconule; ml. midlirtk; pa. paracone.;p.c.posterior lum. Another blade (the apical margin of the
cingulum;pl.f.postcrolingnal fo«a;pr. protocone. buccal forelink) connects the protoloph, approx-
imately one third of the way between the
Palorchestesselestiarn, sp. paracone and the protocone, to the anterior cin-
(Fig. 2) gulum. A well-developed primary midlinkjoins
the protoloph to the metaloph.The posteriorend
Materialexamined of this midlink contacts the trailing edge of the
QBHuloualfrofrtyDyo(pw1e9n:"s4QS3tM"aSFt,i1o21n,445n5°o5r3,t6h'eEaa)ns,teAirlsnollQtauntegeehdnasmLlaFMnodr1.m,aMtiaoinn, TMAB1LinEm1m,.ComparativemeasurementsofPalorchestes
AW PW
Specimen No. L
Etymology
P.selestiae Holotype 22.6 16.6 16.9
ForSelesti SmithofBluffDowns Station.
P.pan'us QMF784(cast) 20.7 15.7 15.4
QMF12476 15.4 15.3
Diagnosis
QMF2963 19.3 14.9 14,4
Small LM* with lingual forehnk terminated in
the cingular valley and not in contact with the QMF3719(cast) 19.3 15 ri 14.:
cingulum; second incipient buttress on theback QMi 2967 19.4 15.5 15.6
ofthemetaloph; posterolingualfossawelldevel- QMF2965 20.9 14.5
oped; posterior cingulum rises to an apex mid- QMF789 19.5
length;incipientbuttressingcmtheposteriorwall P. cfpQrvus P24097(R) 18.8 I3.fi 12.5
ofmetaloph; onemidlink and twoincipient mid- P24097(L) v, 13.9 13.6
1
links;doubleforelinks. P.azael QMF772(cast) 26.6 21.9 21.5
QMF3837 25.8 20.7 19.7
Description Ml370 28.3 21.8 21.4
The holotypeconsistsofacomplete, relatively
P3137) 28.3 22.6 21.9
unworn LM*, lacking any remnant of the roots.
The metaconule is the tallest cusp with the pro- P31372 26. 22.9 21.9
tocone, paracone and metacone subequal in P. painei UCMP70553(R) 16.5* 13.6* 13.8
height. Protocone linked to the paracone by a UCMP70553(X) 16.8 14.4 1-3,7
well-defined sinuous protoloph. Metaconule UCMP7055O(R) 16.7 13.9* 13.7
linkedtothemetaconebyawell-definedcrescen- UCMP6652KL> 17.8 14.0* , ; 2
tic metaloph whose ends are posteriorly de- =approximation
flected.
NEWSPECIESOFPALORCHESTiDMARSUPIAL
(SOS
TABLE2: Summaryoflinks inPatorchestesM1 PHYLOGEtfETIC AFFTN1TES
()Incipient link; -Link/sworn.*cast.
The early Miocene Nagapkaidia tedfordi and
Specimen Ml Forelinks Midlink Hindlinks Pitihanria dailyi (Stirton, 1967) from the Tirari
&
Desert of South Australia have previously been
xelestiae Hoiotype 2 HI) (2)
sp.tmv. regarded as thepleisomorphic sistergroup (sub-
P.parvus QMF784* 2(1) i 2 familiaeIncertae)oftheapparentlymorederived
QMP124T6 2 2 Palorchestinae based on synapomorphies in the
qmr:^ auditory region and neurocranium (Stirton eta).,
2(1) 2 2 1967). Ithasbeen furthersuggestedby Stirton et
QMF3719* 2 2 2 al.(1967:154)thatPitikantia"...ismuchnearer,
OMF2967 2 2 2 if w\t in, an ancestral position to Palortkestes*'.
OMF2965 2 j Archer(1984),however,observedthattherewas
P.azael QMF772* 2 i(2) littleevidence toregard thetwogroupsasmono-
QMF3837 2 phyletic, wilh species of Ngapakaldia and
1
P.painei UCMP70553 i 1 J PtirtaicLtaendiinaalsaaclkibnogntehse sopfecipaalliozrecdhemsotlianress,anedvreen-
UCMF*552l
J 1 1 thoughtheirbasicranial morphologywassimilar.
posteriormetalophblade.Anotherlessdeveloped Murray (19S6) described the middle Miocene
midlink occurs on the lingual side ofthe tooth. PropalorchestesnovacutacephalusfromtheBul-
This smaller link is steeply v-shaped rather than lockCreek LocalFaunaand, based on itscranial
shallowly v-shaped as is the primary midlink. base, glenoid fossa morphology and auditory re-
There is a very deep fossa between these two gion, placed it within the subfamily Pal-
midlinks. orchestinae. Subsequently, Murray (1990)
Thereisalsoaswollenbuttressontheposterior describedthedentitionofP.Tiovaculacephalusas
flank of the protoloph, buccal to the primary well as a new taxon. F, ponticulus from D-site
midlink. Another steeply inclined buttress de- (System A), Riversleigh, Queensland. The latter
scendsposteriorlytotheposteriorcingulumfrom was alsoplaced withinthe Palorchestinae.
methneettatcibpolnoafudlteehettomheatttheaccpooonsntunelereicoatrnsdcistnuhgpeuplouarmpt.sexAansoimfnaclitlpheier- cohrTacrhhaeecsttieMdrsi1s,(eWdthobeoydmesol,asbt1o9rd5ai8ta;egdnWofosoortdei-bc,uhrtionnoedt,-ha1fn9od6r7m)pi.adli--l
blade-like structure is present on the posterior links. These features are lacking in species of
wall ofthe metaloph, buccal to the primary pos- Ngapakaldia and Pitikantiabutpresentorincip-
teriorblade. There is some slight crenulation of ient in Propalorchesfes novaculacephalus* con-
the enamel at thebase ofthe lophs. sidered to be the sister group of Palorchestet
Murray(1986, 1990).Patorchestespaineihasthe
Remarks simplestsystem Oflinkswithonefore-,onehind,
P. selestiaeis approximately3/4the sizeofP. and one midlink which is thecondition alsoseen
azael Owen and is 1/5 Larger than P. parvus De in P. novacutacephalus. All otherspeciesofPal-
Vis.WhilstcloserinsizetoP.parrns* P.selestiue orchesteshsvctwolorchnksasasynapomorphy.
IS morphologically closer to P. azael. P. azael
differs in that the lingual forelink links the pro-
tocone to the anterior cingulum, and lacks the
second incipient buttress on the back of the
metaloph. P. selestiae has a better developed
posierolingual fossa and the apex midlength on
the cingulum is absent in P. azael. In P. parvus
the anterobuccal fossa is less developed, both
forelinksjointheanteriorcingulum,therearctwo
well-defined hindlinks which join directly with
the posterior cingulum, and two midlinks are
present. P.panelWoodburne has a definite mid-
link,absentinP.selestiae,anditlacksthedouble
forelink and the two incipient buttresses on the FIG. 2. Palorcheues seiestiae hoiotype QMF12455.
posteriorwall ofthemetaloph. Actualsize.Occlusalviewstereopair.Anterioratinp.
606 MEMOIRSOFTHEQUEENSLAND MUSEUM
verse correlates of age. By
23 - /** those measures, P. painei,
A \ as the smallest species,
\ ought to be the oldest fol-
22 ~ lowed, in decreasing age
*
1 A : and increasing size by P.
pan>us, P. selestiae and P.
21 i azael.However,P.selestiae
A t
I i comes from ademonstrably
r early Pliocene site, whereas
20 thesmallerP.parvuscomes
IJ* from early to middle
Pliocene sites which does
19
not support the
i biostratigraphic argument
ofStirtone/a/.(1967).
Several authors (Archer,
1976b; Murray, 1990) have
17 proposed that thebilophod-
onl uppermolars ofspecies
of Palorchestes are deriv-
16 ablefromsubselenodontan-
imals such as wynyardiids.
I I Murray (1990:49) suggests
15 b ". .The possibility of
; .
paraphyletic bilophodonty
could not be ruled out, par-
14
ticularly with respect to the
diprotodontids". In this
13 view those animals such as
the late Miocene P. painei
II whichhadtheleastdevelop-
12 1 I i i i 1 1 i ment ofselenodontfeatures
16 1B 20 TOOT2H2 LENGT2H4 26 26 30 would be regarded as the
most derived. It would then
ANTERIORWIDTH P.paOfnel P.selOestiae P.paarvus P.azAael follow that,barringconver-
POSTERIORWIDTH • A gence, the more elaborate
the cresting, the more
p^lesiomorphic the animal.
FIG.3.Bivariateplotsoftooihlengthagainstanteriorwidthf)andtoothlength seledont groundplan is
againstposteriorwidth M1 ofPalorchestesspecies. still evident in P. selestiae,
even though ithas been ob-
Palorchestesselestiae and P. azael share acces- scured to some degree by
sory midlinks and the reduction ofhindlinks as the developmentoflophs.
synapomorphies while P. parvus has two
hindlinks and two midlinks as autapomorphies. PALAEOECOLOGY
The developmentofaccessory hindlinks and the
Ungualforelink notextending tothe anteriorcin- Palorchestidsare uncommon elements in most
gulum in P. selestiae are considered au- Australian marsupial faunas although they have
tapomorphies forthis taxon. a long Tertiary and Quaternary history (Murray,
Palorchestes selestiae indicates need for cau- 1991).Theoldestknownrepresentativesarespe-
tion in determining phylogenetic relationships cies of Ngapakaldia and Pitikantidia from the
within diprotodontoid lineages based on size Oligo-Miocene deposits ofcentral Australia and
alone. Stirton et al. (1967) suggested that molar Riversleigh in northwestern Queensland. At the
sizeandcomplexity in diprotodontoids were in- younger end of their time range, some may be
NEW SPECIES OFPALORCHESTID MARSUPIAL 60?
P.Mfest/a* F.parvus equipped&with large laterally compressed claws.
Flannery Archer(1985) haveused postcramal
material and teeth to attemptareconstruction of
twospecies. Their analyses suggested the possi-
bility that speciesofPalorchestesweremedium-
sizedfolivoresthatmayhaveusedtheirpowerful
clawsandarmstoripbarkofftreesforfoodorlo
uproot shrubs with tuberousroots.
ACKNOWLEDGEMENTS
TWOFORELINK&
I thank Michael Archerand Suzanne Hand for
helpful comments on the manuscript. Sue
Creagh, PeterMurray, NevillePledgeand Ralph
Molnar for access to other palorchestid speci-
TOiFEMVIEDLUONPKMSENT mens. Jack, Rhonda, Bram, Troy and Selesti
SmithofBluffDownsStationcontinuetoprovide
tremendous support for this ongoing research. 1
thank Henk Godthelp, Darren Sprott and Frank
Coffafortheirassistance.CollectionoftheBluff
Downsmaterial wassupportedinpartbyanARC
Program Grant to M. Archer; a grant from the
Department of Arts, Sport, the Environment,
M Tourism and Territories to M. Archer, S. Hand
FIG.4. Cladogramoftaxabasedon 1 morphology. andH.Godthelp;agrantfromtheNationalEstate
Program Grants Scheme to M. Archer and A.
only 30000 yBP(Murray, 1991) and it has been
suggested that Palorchestes was contemporane-
ous with humans. Murray & Chaloupka (1984)
have interpreted an Aboriginal rock painting in
the Arnhem Land escarpment as a possible ren-
dition ofa species ofPalorchestesalthough this
&
hasbeenquestioned(Lewis, 1986; Chaloupka
Murray, 1986; Mackness, 1992).
The reduction and retraction of the nasals,
along with a conspicuous fossae for large
nasomaxillolabialis muscles and a narrow, pro-
tracted rostrum have been used as a basis for
reconstructingpalorchestids with atrunkand the
basts of their popular description as *marsupial P.parvus
tapirs* (Bartholomai, 1978; Flannery, 1983;
& P Aml*xUiM
Flannery Archer, 1985; Murray, 1991). The h-
long,narrowdeeplygroovedsymphysissuggests
that palorchestids had long protrusible giraffe-
liketongues (Murray, 1991).Thespatulatelower
incisorsarealmostparallel tostronglydeveloped
diastemal crests and interpreted by Murray
(1991) as probably being used to strip leaves or
crop clumped vegetation. Little has been pub-
lished about the postcranials of palorchestids.
Woods (1958) cast doubt on the association of
postcranialswithskullremainssuggestedbysev-
eral authors (Owen, 1876; Gregory, 1902;
Fletcher, 1945). Archer (1984), however, has FIG. 5. DistributionofspeciesofPalorchestes,
noted that Palorchestes has powerful forclimbs
.
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