Postcranial Remains of the Extinct Monkeys of the Greater Antilles, with Evidence for Semiterrestriality in Paralouatta1

2006·4.6 MB·English

by  R. D. E. Macphee

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3516, 65 pp., 37 figures, 16 tables May 17, 2006
Postcranial Remains of the Extinct Monkeys of the
Greater Antilles, with Evidence for Semiterrestriality
1
in Paralouatta
R.D.E. MACPHEE2 AND JEFF MELDRUM3
CONTENTS
CONTENTS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
IInnttrroodduuccttiioonn .................................................................................................................. 33
BBooddyy SSiizzeeiinnXXeennoottrriicchhiinnii ............................................................................................ 33
MMaatteerriiaallaannddMMeetthhooddss.................................................................................................. 66
AAbbbbrreevviiaattiioonnss.............................................................................................................. 77
AAnnaattoommiiccaall.............................................................................................................. 77
MMeeaassuurreemmeennttss.......................................................................................................... 77
OOtthheerr...................................................................................................................... 77
JJaammaaiiccaannFFoossssiillssaannddLLooccaalliittiieess........................................................................................ 88
CCuubbaannFFoossssiillssaannddLLooccaalliittiieess.......................................................................................... 1111
HHiissppaanniioollaannFFoossssiillss aannddLLooccaalliittiieess.................................................................................. 1144
PPoossttccrraanniiaallEElleemmeennttssAAttttrriibbuutteeddttooXXeennootthhrriixx,,PPaarraalloouuaattttaa,,aannddAAnnttiillllootthhrriixx ...................... 1144
HHuummeerrii .................................................................................................................... 1155
XXeennootthhrriixxAAMMNNHHMM226688000022,,226688000055,,aanndd226688000088 .................................................... 1155
PPaarraalloouuaattttaaMMNNHHNNCCuu7766..11001100––7766..11001133,,7766..11002244,,aanndd7766..11003355.................................. 1188
FFuunnccttiioonnaallCCoonnssiiddeerraattiioonnss...................................................................................... 2211
RReellaattiivveeMMeeddiiaallEEppiiccoonnddyylleeSSiizzeeaannddOOrriieennttaattiioonn.................................................. 2211
TTrroocchhlleeaarrOOrrggaanniizzaattiioonn ...................................................................................... 2244
UUllnnaa aannddRRaaddiiuuss........................................................................................................ 2255
XXeennootthhrriixxAAMMNNHHMM226688001100.................................................................................... 2255
1Contribution11totheseries‘‘OriginoftheAntilleanLandMammalFauna.’’
2DivisionofVertebrateZoology(Mammalogy),AmericanMuseumofNaturalHistory([email protected]).
3DepartmentofBiologicalSciences,IdahoStateUniversity,Pocatello,ID83209([email protected]).
CopyrightEAmericanMuseumofNaturalHistory2006 ISSN0003-0082
2 AMERICAN MUSEUMNOVITATES NO. 3516
PPaarraalloouuaattttaa MMNNHHNNCCuu 7766..11001144,, 7766..11001166,, aanndd 7766..11001177 ...... ........ ........ ........ ........ ........ 2277
FFuunnccttiioonnaall CCoonnssiiddeerraattiioonnss...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 2299
OOlleeccrraannoonn SSuurrffaaccee ooff TTrroocchhlleeaarr NNoottcchh ........ ........ ........ ........ ........ ........ ........ ........ 2299
OOss CCooxxaaee.... ........ ...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 3300
XXeennootthhrriixx AAMMNNHHMM 226688000099.... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 3300
PPaarraalloouuaattttaa MMNNHHNNCCuu 7766..11003366 ...... ........ ........ ........ ........ ........ ........ ........ ........ ........ 3322
FFuunnccttiioonnaall CCoonnssiiddeerraattiioonnss...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 3322
AAcceettaabbuullaarr WWaallllss........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 3322
FFeemmuurr ........ ........ ...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 3377
XXeennootthhrriixx AAMMNNHHMM 226688000033.... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 3377
PPaarraalloouuaattttaa MMNNHHNNCCuu 7766..11001188aanndd7766..11001199 .... ........ ........ ........ ........ ........ ........ ........ 3388
FFuunnccttiioonnaall CCoonnssiiddeerraattiioonnss...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 3399
HHuummeerruuss//FFeemmuurrPProropporotriotinoanlaitlyity................................................................................... 3399
DDiissttaall EEppiipphhyyssiissAAnntteerrooppoosstteerriioorr CCoomm-pression and Condylar Proportionality. . . . . 39
?prTeisbsiiaon. a.n.d. .C.o.n.d.yl.a.r.P.r.op.o.r.ti.o.n.a.lit.y... ........ ........ ........ ........ ........ ........ ........ ........ ........ 3490
TibXiaen.o.th.r.ix. .A.M. .N.H.M. . .25.9.9.0.3.... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 4400
XPaenraoltohuraixttaAMMNNHHMNC2u597960.130.3.4. ...... ........ ........ ........ ........ ........ ........ ........ ........ ........ 4400
PAanrtaillloouthartitxaUMSNNHMNMCu257466.18023.4....... ........ ........ ........ ........ ........ ........ ........ ........ ........ 4403
AFuntniclltoiothnraixl CUoSnNsiMdeMrati2o5n4s6.82... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 4434
FuFnicbtuiolanralACpopnresisdsieornati.o.ns....... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 4444
CheirFidibiaul.ar. .A.p.p.re.s.si.o.n..... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 4445
ChPeiarirdailaou.a.tt.a.M. .N.H.N. .C.u.7.6..1.0.3.7.a.n.d. .76...3.05.9. .(T.a.lu.s.). ........ ........ ........ ........ ........ ........ 4456
PPaarraalloouuaattttaaMMNNHHNNCCuu7766..11002307(aCnadlc7a6n.e3u0s5)9a(nTdaMlusN)H.N. .C.u.7.6..1.0.2.1. . . . . . . . . . . . . . 46
Par(aMloiudadtlteaCMuNneHifNorCmu)7.6..1.0.2.0.(.C.a.lc.a.n.e.u.s).a.n.d..M.N..H.N..C.u.7.6...10.2.1..(M..id.d.l.e.C.u.n.e.if.o.r.m.) 5500
PPaarraalloouuaattttaa MMNNHHNNCCuu 7766..11002222,, 7766..11002233,, aanndd 7766..11002255––7766..11002288 ((MMeettaappooddiiaallss)) .. .. ........ 5511
PPaarraalloouuaattttaa MMNNHHNNCCuu 7766..11002299––7766..11003333((PPhhaallaannggeess)) ........ ........ ........ ........ ........ ........ 555121
FFuunnccttiioonnaall CCoonnssiiddeerraattiioonnss...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 5555
DDiiggiittaall RRaayy LLeennggtthh .... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 5555
DDiissccuussssiioonn...... ........ ...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 5577
CCaalllliicceebbiinnaaee aanndd XXeennoottrriicchhiinniiaassPPhhyylleettiicc AArrrraayyss.. ........ ........ ........ ........ ........ ........ ........ 5577
SSiizzee VVaarriiaattiioonn iinnPPaarraalloouuaattttaa...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 5577
LLooccoommoottoorr AAddaappttaattiioonnss ooffXXeennootthhrriixx.. ........ ........ ........ ........ ........ ........ ........ ........ ........ 5588
SSeemmiitteerrrreessttrriiaalliittyy iinnPPaarraalloouuaattttaa.. ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 5588
ExcElxucdlaubdlaebPleaPttaerttnesrn.s.:.?. ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 5599
ElbEolwbo.w.:.?........ ...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 6600
KnKeenee.:.?.. ........ ...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 6600
ProPxriomxaiml TalarTsaurssu.s:.?....... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 6600
DigDitisgi.ts.:?. ........ ...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 666010
AAcckknnoowwlleeddggmmeennttss .. ...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 6611
RReeffeerreenncceess...... ........ ...... ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ ........ 6611
Ap?p??e?n?d??ixAP1:PSEpNeDcimIXen1s of Extant Primates Utilized for Morphological Comparisons and
MSpeeacsiumreenmseonftEsx.t.an.t.P.r.im.a.t.es.U.t.il.iz.ed. .fo.r.M.o.r.p.ho.lo.g.ic.a.l.C.om. .pa.r.is.on.s. . . . . . . . . . . . . . . . . . 65
andMeasurements
2006 MACPHEEAND MELDRUM: POSTCRANIALREMAINS OFANTILLEAN MONKEYS 3
ABSTRACT
This paper describes postcranial remains pertaining to the endemic xenotrichin callicebines of
theGreaterAntilles,allofwhichareextinct:Xenothrixmcgregori(Jamaica),Antillothrixbernensis
(Hispaniola), and Paralouatta varonai and P. marianae (Cuba). These monkeys differed
considerably in body size and inferred locomotor behavior. Xenothrix and Antillothrix are
estimated to have weighed 2–5kg, which is well within the middle range of body sizes found in
extantSouthAmericanmonkeys,butParalouatta(,9–10kg)wouldhavebeennearlyaslargeas
the largest living platyrrhines. In line with previous studies, we interpret Xenothrix mcgregori as
a rather short-limbed, slow-moving arboreal quadruped possessing some unusual features not
otherwise seen in platyrrhines (e.g., adductor process or ‘‘fourth trochanter’’ of the femur). Its
closest locomotor analog among living primates remains uncertain. Paralouatta varonai also
exhibitsfeaturesnotseeninotherplatyrrhines,butinthiscasethereareintriguingresemblancesto
certainOldWorldmonkeys(e.g.,retroflexedmedialepicondyleandnarrowtrochleaonhumerus,
stabilization features of talocrural joint, short digital rays), especially so-called semiterrestrial
cercopithecines whose locomotor repertoire includes a significant amount of movement on the
ground (e.g., Cercopithecus lhoesti). At the same time, the Cuban monkey conspicuously lacks
most features uniquely connected with suspensory activities, otherwise seen in all living
platyrrhines of large body size. The locomotor and postural repertoire of Antillothrix is
unresolved, as the only element currently available for analysis is a distal tibia. The tibia of the
Hispaniolanmonkeyisnotveryinformativefromafunctionalstandpoint,althoughitexhibitsless
emphasisontalocruralstabilizationthandoestheequivalentelementinParalouatta(e.g., sizeof
medialmalleolus).Thediversepostcranialspecializationsexhibitedbyxenotrichinsareconsistent
withtheirlong isolation(at leastsinceOligocene) on landmasses in the CaribbeanSea.
INTRODUCTION and elements already known from published
materialaregivencursorytreatment.
The amount of fossil evidence pertaining to
In previous work, MacPhee and Fleagle
Xenotrichini, a clade of extinct callicebine
(1991) showed that theknown postcranium of
monkeys endemic to the Greater Antilles, has
Xenothrix was distinctively different from the
grown substantially over the past quarter
postcrania of other New World monkeys in
century (MacPhee and Woods, 1982;
a number of ways. Paralouatta is, if anything,
MacPhee, 1984, 1996; Ford, 1986a, 1986b,
even more distinctive—but in ways that con-
1990;FordandMorgan,1986;MacPheeetal.,
verge to some extent on living Old World
1989, 1995, 2003; MacPhee and Fleagle, 1991;
monkeys that spend significant amounts of
Jaimez Salgado et al., 1992; MacPhee and
time on the ground. To what degree
Iturralde-Vinent, 1995, 2000; Horovitz, 1997;
Paralouattamayhaveactuallyresembledthese
Horovitz and MacPhee, 1999; MacPhee and
cercopithecids in locomotor and postural
Horovitz, 2002, 2004). With the exception of
behaviors is a complicated question that will
specimens of the Hispaniolan species
be taken up in the Discussion. However, the
Antillothrix bernensis (5 ‘‘Saimiri’’ bernensis),
fact that at least one lineage of platyrrhines
under study by others, the only xenotrichin
might have experimented with semiterrestrial-
fossils not yet adequately described consist of
ityasearlyastheEarlyMioceneisonlyslightly
postcranial elements of Xenothrix mcgregori
less surprising than the fact that the lineage in
(Jamaica) and Paralouatta varonai (Cuba)
questionevolvedinanislandcontext.
collected in recent years by American
Museum teams and collaborators (tables 1,
2). Here we present brief descriptions of this BODY SIZEIN XENOTRICHINI
material and evaluate its significance for un-
derstanding the functional morphology of this Body size or mass (BM) is an important
vanished group of island primates. As this is variable when considering functional attri-
meant to be an interpretative paper, the best butes of extinct species, especially when such
preservedandmostinstructiveelementsreceive species have few or no modern analogs for
the bulk of attention; fragmentary specimens theirinferredlocomotoryorpositionalbehav-
3
4 AMERICAN MUSEUMNOVITATES NO. 3516
TABLE1 TABLE2
Postcranial Specimens: Xenothrixmcgregoria Postcranial Specimens: Paralouattavaronaia
AMNHM Element Localityb Description MNHNCub Elementc Description
259900 femur(R) LongMile completeexcepthead 76.1010 humerus(R) completeexcepthead,
259901 humerus(R) LongMile proximalend capitulum
259902 tibia(R) LongMile proximalend 76.1011 humerus(L) diaphysis
259903 tibia(L) LongMile proximalend 76.1012 humerus(R) headonly,damaged
259904 oscoxae(L) LongMile acetabulum,partsof 76.1013 humerus(R) distalarticularend
ilium,ischium 76.1014 ulna(R) tricepsprocessonly
259905 humerus(R) LongMile shaftonly 76.1015 radius(L) proximalend
259906 humerus(L) LongMile shaftanddistalend 76.1016 ulna(L) proximalend,damaged
259907 ulna(R) LongMile proximalend 76.1017 ulna(L) proximalend,damaged
259908 sacrum LongMile partialsacrum 76.1018 femur(L) proximalend,severely
268002 humerus(R) Drum proximalend damaged
268003 femur(R) Somerville distalend 76.1019 femur(L) distalepiphysis
268005 humerus(L) New(?) proximalend 76.1020 calcaneus(L) talarfacetareaonly,
268008 humerus(R) Drum completeexcept damaged
head 76.1021 middle complete
268009 oscoxae(R) Skeleton acetabulum,partsof cuneiform(R)
ilium,ischium, 76.1022 MC1(R) complete
pubis 76.1023 MC2(R) complete
268010 ulna(R) Somerville proximalend 76.1024 humerus(L) distalarticularend,
aFororiginaldescriptionsofcraniodentalmaterialnot partialtrochleaonly
76.1025 MT1(L) complete
discussed here, see: Williams and Koopman (1952), 76.1026 MT1(R) proximalend
holotype mandible; Rosenberger (1977), redescription of 76.1027 MT3(L) complete
holotype; MacPhee and Horovitz (2004), skull and 76.1028 MT3(R) complete
76.1029 proximalphalanx, complete,?pes
additionalmandibularremains.
DR1(R)
bLongMileCave,collectorH.E.Anthony,1920;New 76.1030a–c 3proximal acomplete,bdistalend,
Cave,collectorsK.KoopmanandM.Hecht,ca.1950;all phalanges, cproximalend
othersitesworkedbyAMNH/CMCexpeditionsin1990s. DR2–5
76.1031a–l 12middle mostspecimens
ExceptforAMNHM268003,whichwasfoundinsurface
phalanges, complete
debris,allspecimenswererecoveredincourseofexcava- DR2–5
tions. 76.1032a–d 4distalphalanges complete,withexpanded
tips
76.1033 1distalphalanx, completebutwater-
DR2–5 rolled,unexpanded
76.1034 tibia(R) shaftanddistalend
76.1035 humerus(L) shaftanddistalend
ior within their phylogenetic group (Martin,
76.1036 oscoxae(R) acetabulararea,partof
1990). To provide a consistent basis for iliacblade
estimating BMs among xenotrichins, we se- 76.1037 talus(R) body,damaged
lected Conroy’s (1987) ordinary least squares aFor additional descriptions of material, see: Rivero
approach, which relates log of average m1 and Arredondo (1991); Horovitz (1997), Horovitz and
e
area(independentvariable)toBM(dependent MacPhee (1999), and MacPhee and Iturralde-Vinent
variable).Weacknowledgethat,asapredictive (1995).
technique,anyregressionapproachhaslimita- bCurrentMNHNCucatalognumbersdifferfromthose
previouslyinuse.Itisespeciallyimportanttonotethatthe
tions, especially when (as here) the target taxa
(a)P.varonaiskull(holotype),previouslypublishedunder
are not actually part of the reference sample
the MNHNCu numbers V90-25 and V194, is now
(Jungers,1990).Evaluationsofresultsintable3
cataloged as 76.2565; (b) P. marianae talus (holotype),
areprovidedinfollowingparagraphs. previously P3059, is now 76.3059; (c) P. varonai partial
ForAntillothrix,theleastknownofthethree talus,previouslyV205andlaterP2565,isnow76.1037;(c)
knownxenotrichins,BMhastobebasedforthe P. varonai partial jaw V195 is now 76.1213; (d) and
presentonasinglejawfragmentcontainingm1 mandibularmolarspecimensV123,138,144,145arenow
(MacPhee and Woods, 1982). The tooth is numberedseriallyfrom76.1214through76.1217.
cAllspecimenslistedinthistablearefromCuevaAlta,
worn, but its sidewalls are intact and thus
excepthumerusMNHNCu76.1035andMT3MNHNCu
appropriate for measurement. In the case of
1027 which are from Cueva del Mono Fos´ıl. Material is
Xenothrix, the m1s of AMNHM 268001 and
currently housed in MNHNCu and was collected in
268004, two recently described mandibles collaboration with members of Grupo Espeleologico
(MacPhee and Horovitz, 2004), are severely ‘‘PedroA.Borra´s’’ofLaHabana,Cuba.
2006 MACPHEEAND MELDRUM: POSTCRANIALREMAINS OFANTILLEAN MONKEYS 5
damaged.Measurementsarethereforebasedon talar length, which suggests in turn that one
the type, AMNHM 148198, whose m1 is should be conservative in accepting a very
fortunately well preserved. For Paralouatta, large BM estimate for this monkey.
measurements are based on a sample of five For Antillothrix, the calculated estimates of
teeth, four of which represent isolated finds 4.5–5.0 kg also seem large, but it is hard to
(MNHNCu76.1214–1217) and are identifiable have a good sense of the size of this primate
only as ‘‘m1 or 2’’ (Horovitz and MacPhee, because the only other published craniodental
1999: their table 1). The fifth m1 is present in fossilofthisspeciesisthetypemaxilla.R´ımoli
a beautifully preserved partial jaw, MNHNCu (1977), the type’s discoverer, thought that
76.1213(HorovitzandMacPhee,1999),andits it represented a squirrel monkey (hence
locusisthereforecertain.Thefactthatmostof ‘‘Saimiri’’ bernensis), but allowed that its
the teeth in the sample were isolated finds is owner would have been closer to a living
important; first and second lower molars are capuchin in BM. Ford (1986a) suggested
homomorphic in Paralouatta, and in practice that Chiropotes was an equally good size
they cannot be metrically or morphologically proxy, but had in mind a body weight closer
discriminated. Thus, even though m2s are to 3 kg.
doubtless present in our sample, in principle The estimates for Xenothrix, 5.5–6.5 kg,
theyshouldnotaffectBMestimationsbasedon also seem large and moreover are in conflict
themeasurementsemployedinConroy’s(1987) with the 2–4-kg range predicted by MacPhee
regressionexpressions. andFleagle(1991)usingRuff’s(1987)femoral
We utilized Conroy’s expressions for ‘‘all cortical cross-section method on AMNHM
primates,’’ ‘‘anthropoidgrade,’’and‘‘monkey 259900.(Unfortunately,thismethodcouldnot
grade’’ (table 3). Although on grounds of beusedasanindependentcheckondentition-
narrow allometry the ‘‘monkey grade’’ esti- based BM estimates for Paralouatta because
matesmightbepreferred,allthreeexpressions there are no suitable femoral specimens in the
produced rather similar results. In the case of current hypodigm.) An obvious possibility is
Paralouatta, estimates are in the range of that Conroy’s (1987) method yields over-
9–10 kg, with MNHNCu 76.1213 producing estimates not only when an unknown is in
the largest result. BMs in this range are theupperrangeoftoothsizesforitsgroup(as
large for platyrrhines; the implication is he acknowledged), but also when the molar
that Paralouatta groups with the largest row is reduced—the theory being, in the case
living taxa, such as Brachyteles arachnoides of Xenothrix at least, that m1 is larger than
andlargespeciesofAtelesandAlouatta,allof would be ‘‘typically’’ expected. An analogous
which meet or exceed 10 kg in BM (Fleagle, issue may affect estimates for ‘‘big-toothed’’
1999:table5.4).However,aswenotelater(see species like Cebus apella, whose ‘‘monkey
Cheiridia), compared to other platyrrhines, grade’’ predicted range of 4.0–5.5 kg
Paralouatta has a relatively large m1 for its (Conroy, 1987: his table 1) lies well beyond
TABLE3
BodySize Estimates: Antillothrix,Xenothrix andParalouattaa
Bodysizeestimates(g):
All Anthropoid Monkey
Taxon Mandibularm1specimens BL3MD Log m1area primates ‘‘grade’’ ‘‘grade’’
e
Antillothrixbernensis UF28038 4.935.5 3.2940 4520.9 5174.8 5176.6
Xenothrixmcgregori AMNHM148198 5.136.1 3.4372 5840.4 6476.8 6482.9
Paralouattavaronai MNHNCu76.1213–1217 5.737.0b 3.6863 9104.2 9581.7 9551.3
Paralouattavaronai MNHNCu76.1213only 5.637.4 3.7242 9740.6 10,168.7 10,133.0
aBasedonordinaryleastsquaresregressionexpressionspublishedbyConroy(1987).
bMeanvaluesforthesefivespecimens.
6 AMERICAN MUSEUMNOVITATES NO. 3516
its empirical average of 2.5–3.6 kg (cf. Silva individuals. For this study, the necessary
and Downing, 1995; Fleagle, 1999). compromise was to use the best of what there
Insummary,wesuspectthataverageBMin is, even if the specimens in question lacked
Paralouatta could have been as much as 9 kg, provenance data. Insome cases, species inthe
butmaywellhavebeensomewhatless.Thisis same genus have been combined when there
still, of course, very large for a platyrrhine was not enough material to make up an
monkey. With regard to Xenothrix, it seems adequate sample.
quite unlikely that the Jamaican monkey Obviously, providing equal coverage in the
couldhaveexceeded5 kginviewofitsgeneral figures for so wide a range of taxa is not
similarity in long bone lengths to monkeys possible, nor even particularly useful unless
such as Cebus. The conclusion is the same for noted resemblances have some significant
Antillothrix, whose dental measurements are correlation with function. In this paper we
slightly smaller. We continue to prefer the 2– concentrate on illustrating major limb bones
4 kg range originally predicted by MacPhee (especiallythehumerus,ulna,andtibia),using
and Fleagle (1991) for Xenothrix, which for comparative purposes a set of New and
means that this monkey would have been OldWorldmonkeys whicharepredominantly
approximately one-half to one-third the quadrupedal in their locomotor behavior
size of Paralouatta as currently estimated. (whether in the trees or on the ground). BMs
Interestingly, at 0.9–1.3 kg (Silva and forthecomparativesetareasfollows(dataof
Downing,1995;Fleagle,1999),theonlyextant Fleagle [1999], male/female BMs in kilo-
callicebine, Callicebus, is but one-tenth the grams): (1) tufted capuchin, Cebus apella
estimated size of the Cuban monkey. (2.5/3.7 kg), a medium-sized arboreal quadru-
ped;(2)redhowlermonkey,Alouattaseniculus
(5.2/6.7 kg) a large-bodied arboreal quadru-
MATERIALAND METHODS
ped with some arm swinging capability; (3)
The comparative set of taxa employed by gray-cheeked mangabey, Lophocebus albigena
MacPhee and Horovitz (2004) for evaluating (6.0/8.3 kg), a predominantly arboreal quad-
thecraniodentalmorphologyofXenothrixwas ruped in the size range of Alouatta seniculus;
essentially limited to extant pitheciids and (4) patas monkey, Erythrocebus patas (5.7/
Aotus. Adequate appreciation of the unusual 10.6 kg),anagileground-foragingspecies,and
postcranial morphology of Antillean primates (5)gelada,Theropithecusgelada(11.7/19.0 kg),
requires wider comparisons, and the compar- a very large Old World monkey whose
ative set has been broadened accordingly (see locomotor activities are conducted almost
appendix 1), with special attention given to exclusively on the ground. The last two
species of the following genera (platyrrhines: species are markedly sexually dimorphic.
Callicebus, Chiropotes, Ateles, Brachyteles, Other elements are much more poorly repre-
Alouatta, and Cebus; catarrhines: Trachy- sented in the xenotrichin collection and are
pithecus, Erythrocebus, Lophocebus, and accordingly given less attention in this paper.
Theropithecus). Incidental observations on In general, the usual run of biomechanically
other anthropoids are also provided where important ratios emphasized in functional
appropriate. We have accepted species desig- morphology (e.g., Anapol et al., 2005) could
nations as they appear on specimen labels, not be taken on highly incomplete fossil
although it is clear that, given the current material, although we have tried to fashion
drive to relimit numerous species of primate a few others that seem to have some in-
taxa (e.g., Groves, 2005), the AMNHM terpretative significance. Appropriate com-
collection stands in need of major revision. parative illustrations of conditions in other
Although the primate collections of the primates for some elements (femur, os coxae)
AMNHM are very extensive, for not a few can be found in the paper by MacPhee and
species there is little postcranial material; Fleagle (1991).
worse, even for some well represented species, MacPhee and Fleagle’s (1991) study should
so much postcranial material has been ‘‘lost’’ also be consulted for additional comparative
overtheyearsthatitisnowhardtofindintact observations and illustrations, as well as
2006 MACPHEEAND MELDRUM: POSTCRANIALREMAINS OFANTILLEAN MONKEYS 7
measurements and ratios not defined here. MC metacarpal (with or without digital
Anatomical names mostly conform to the ray specified)
most recent Terminologia anatomica (Feder- ML mediolateral(ly)
ative Committee of Anatomical Terminology, mltr medial lip of trochlea(humerus)
1998), except in instances where homologs in mm medial malleolus
msr medial supracondylar ridge
Homosapiensarenotobviousoranotherterm
MT metatarsal (with or without digital
is to be preferred.
ray specified)
mw medial ‘‘wing’’ of the olecranon
ABBREVIATIONS surface of the trochlear notch (ulna)
nv navicular
Anatomical
ol olecranon
ac anteriorcrest (tibia) olf olecranonfossa
ap adductorprocess (femur) OSTN olecranonsurface of trochlear notch
apl abductor pollicis longus m. (inser- PP proximal phalanx
tion) ps patellar surface
BL buccolingual pt peroneal trochlea(talus)
BM bodysize rf radial fossa
br brachialism. (insertion) rn radial notch
brf brachialisflange sc supinator crest
bt bicipitaltuberosity scf semicondylar facet (tibia)
CC craniocaudal srp superior ramus(pubis)
cf coronoidfossa st sustentaculum tali
cp coronoidprocess tcj talocrural joint surface
cpt capitulum tm teres major m. (insertion)
ctf cotylarfossa (talus) tn trochlear notch
df dorsalepitrochlearfossa tp triceps process (ulnar)
DP distalphalanx tro trochlea
dpe deltopectoraleminence
DR digitalray (unspecified)
Measurements
DR1 digitalray ofthumb orbig toe
DR2–5 otherdigitalrays (manusorpes)
AdjMDW adjusted maximum distal width (of
DV dorsoventral
longbone)
ef entepicondylar foramen
APW anteroposterior
ff fabellarfacet (femur)
DR2L total length of second digital ray
fdp flexor digitorum profundus m. (ori-
(including MC2)
gin)
DVW dorsoventral width
ftp fossafortibialisposteriorm.(origin)
MaxL maximum length(oflimb bone)
gt greatertubercle (humerus)
MLW mediolateral width (oflimb bone)
gtr greatertrochanter
MC2L maximum length of second metacar-
ib interosseousborder
pal
if iliacfossa
MEPD posteriordeviationofmedialepicon-
ig intertubercular groove(sulcus)
dyle(of humerus)
ip iliacplanum
TPL total phalangeal length (i.e., finger/
is ischialspine
toe length)
it ischialtuberosity
lc lateralcondyle (femur)
le lateralepicondyle Other
lsr lateralsupracondylarridge
lt lessertubercle (humerus) asl above sea level
ltc lateralcuneiform bp before present (i.e., before radiocar-
ltr lessertrochanter bon datum, AD 1500)
lw lateral ‘‘wing’’ of the olecranon sur- L leftside
faceof the trochlear notch (ulna) PCA principal componentsanalysis
m.,mm. muscle, muscles R right side
m1,2 mandibularmolar +locus rev. sides photographically reversed (in
me medial epicondyle figures)
8 AMERICAN MUSEUMNOVITATES NO. 3516
UF FloridaMuseumofNaturalHistory, sites are concentrated on the western end of
Gainesville, FL Portland Ridge, a prominent elevated lime-
MNHNCu MuseoNacionaldeHistoriaNatural, stone area in the southernmost part of
LaHabana, Cuba Jamaica, near Jackson’s Bay (17u459N,
AMNHM AmericanMuseumofNaturalHistory
77u159W) in southern Clarendon Parish. New
(Mammalogy)
Cave is known only from a manuscript
USNMM United States National Museum of
notation and has not yet been relocated. The
Natural History (Division of Mam-
only site on the north side of the island that
mals)
has yielded Xenothrix remains is the type
locality, Long Mile Cave. Descriptions of
JAMAICAN FOSSILS AND LOCALITIES
these caves and their speleological, paleonto-
Although it is easy to appreciate the logical, and paleoenvironmental significance
difficulty of assigning isolated bones to in- can be found in papers by MacPhee (1984),
dividual species of primates in contexts in Fincham (1997), McFarlane et al. (2002),
which there are several possible choices, MacPhee and Flemming (2003), MacPhee
assignment can be problematic even when and Horovitz (2004), and references cited
there is only a single conceivable choice. The therein.
gradual accretion of the hypodigm of A nearly complete humerus (AMNHM
Xenothrix mcgregori is a case in point. The 268008) of Xenothrix was recovered from
describers of the Jamaican monkey, Williams a position just above layers dated to 11,220
and Koopman (1952), reported that they 6 100 bp and 10,250 6 80 bp in the Brown
found several unusual mammalian postcra- Dust Passage of Drum Cave (fig. 2A;
nials in H.E. Anthony’s 1919/1920 Jamaican McFarlane et al., 2002). This establishes that
faunal collections, but decided that no con- the Jamaican monkey lived in the vicinity of
clusive allocation of these elements was then Jackson’s Bay—now one of the driest regions
possible. After examining this same material, on the island—at least as early as the close of
MacPhee and Fleagle (1991) concluded other- the Pleistocene. Dates reported by MacPhee
wise: Some of the unusual postcranials could, (1984) on nonprimate bone scrap from levels
they thought, be assigned to Xenothrix on in Long Mile Cave that produced monkey
general morphological grounds, even though remains indicate that Xenothrix was present
thebonesinquestiondidnotgreatlyresemble onthenorthcoastlateintheHolocene.These
those of any other platyrrhine or even any are the only radiometric dates available, and
other anthropoid. Other elements had to be are likely to remain so for the foreseeable
excluded from consideration because they future. Because of the scantiness and fragility
were either too incomplete or too ambiguous of the new Xenothrix material, it was decided
morphologically to assign to any particular to treat all specimens with a consolidant
mammalian group. Since then, renewed col- (Butvar).Thetypejawhadearlierbeentreated
lecting has produced both new cranioden- with a varnish or resin of some sort, pre-
tal material of Xenothrix (MacPhee and sumably by Anthony. Thus all specimens of
Horovitz, 2004) as well as the postcranial Xenothrix in the current hypodigm are con-
remains described in this paper. Although taminated with recent carbon, and they will
these specimens are unassociated and dam- therefore never produce reliable 14C dates.
aged to a greater or lesser extent, they have This is of little paleontological consequence
helpedtoclearupsomemysteriesaswellasto because it is beyond doubt that this monkey
reinforce the likelihood that the majority of survived very late, perhaps even into early
allocations made by MacPhee and Fleagle European colonial times (MacPhee, 1984,
(1991) were correct. 1996). Although it would be of interest to
To date, six localities (figs. 1, 2; table 1) ascertain more precisely when its extinction
have yielded remains attributable to Xeno- took place (cf. Steadman et al., 2005), this
thrix:LongMile,Skeleton,Somerville,Drum, kind of inquiry can only be adequately
Lloyd’s, and ‘‘New’’ Caves. With the excep- conducted through the discovery of many
tion of the first and (possibly) the last, these more specimens than exist at present.
2006 MACPHEEAND MELDRUM: POSTCRANIALREMAINS OFANTILLEAN MONKEYS 9
Fig. 1. Jamaican localities: Locator maps for sites mentioned in text. The greatest concentration of
xenotrichinsitesisinsouthernClarendonParish,nearJackson’sBay(A,B).Cavesinthisareathatyielded
postcranial remains discussed in this paper are in capital italics (C); for other localities, see MacPhee and
Horovitz(2004).Thelocationof‘‘NewCave’’isnotknownbutisbelievedtolieonPortlandRidge;Sheep
PenandCocoReearenotdefinitexenotrichinsites(seetext).MapBbasedonSheetH,SeriesE721(D.O.S.
410),U.K. Directorateof Overseas Surveys forthe JamaicaGovernment; mapC afterFincham (1997).
10 AMERICAN MUSEUMNOVITATES NO. 3516
Fig. 2. Jamaican localities: A, Drum Cave, excavation in Brown Dust Passage near entrance #3. B,
SkeletonCave,showingrearofMapRoomafterexcavationofmostoftheupperpartofthefill(sediment
stain line can be seen to left of individual’s head). Drum Cave has yielded two humeri of Xenothrix
(AMNHM268002,268009)as well as the holotypejawof the ?heptaxodontidrodent Xaymacafulvopulvis
(MacPheeandFlemming,2003).ExcavationsatSkeletonCaveproducedanoscoxae(AMNHM268008)as
wellasthetwonewjawsofXenothrixdescribedbyMacPheeandHorovitz(2004),oneofwhichwasfound
whereindividualispointinginB.ThepaleoecologyofthislocalityisdiscussedbyMcFarlaneetal.(2002).