Platyrrhine evolution in the West Indies

Platyrrhine

evolution

in the West Indies

There are now at least thrre, and prohahl) five to sewn. distinct u~d~mit platyrrhine species known from the Grratu Antilles. All arc 01‘ I.atr Pleistocene or Holoccnr age. These include at lest threr fromJamaica, the distinctive .t’mothrix mqrqori and tv,o others knowm onl! from proxim.tl femora. All three are approsimatcly the same size. One of thrsc is Ao presrnt on Hispaniola. Dental remains from sr\wal sites on Hispaniola all hrmmsi,~ (kchich is liktly not .SuNimin). prohahl) helong to “Saimiri” Postcranial remains lir~m Hispaniola may not belong to the same m~mkc~>, their phenetic (hut perhaps not phylogenetic) afinitirs arc to callitrirhids. Cuba also had two endrmic platyrrhinrs. lrss changed from their mainland relatives, a hov,lcr monke! and a spider monkey (:lMrs rmlhrr~p~~,umorl,h/~r, I. One or mow of thrsc linrages proha& rearhvd the islands prior to thl. Pleistocene. and almost certainly through dispcrwl (!?a raliinq) end n,o vicariancc or hurn~n trx~uport.

Introduction The

fossil record

e.g., Fleagle, 1985,

et al.,

includes

the remains

the radiation recent

1983,

chapter

For almost

70 years

(Ameghino,

~Zntilles in some

There

remote

Greater

clear

Antillean

to a vicariant

evidence

islands

usually

questions

about

adaptive

niches

of endemic

(Hispaniola, origin.

not directly the

evolution

previously

New

Jamaica, This

record

associated of body unknown

Overview

travelling

from the Greater

monkey

and Cuba), includes

with

one another.

size

and

although

and relationships

or if

Central 1982).

from

of the

three

questions fossils

in fact,

remain

and

important

suggest

have

of New World

as

postcranial

raise

on islands,

may,

thr

there,

& LYoods,

cranial,

These

competition and

species

dental,

reached

from either

2; see MacPhee

M:orld

Pleistocene

in isolation

by canoe

to

the most

1).

platyrrhines

evolution

(Figure

for primates,

bearing on our understanding of the origins the South American mainland.

monkey

and

related

ways,

(Figure

)-ears

1985).

that of the Late

as to whether

independent

argued

In many

enigmatic

of a platyrrhine

doubt

million

ut al.,

are more clearly

are others.

is the most

by Amerindians

researchers

or dispersal

remains,

remained

& Rosenbrrger,

MacFadden

of the platyrrhines,

past and underwent

as most

than

the first report

to the islands

America,

is now

following

1990;

(see.

1986. 1988;

the last 25 or more

Some of these

lineages

history

in the last decade

1985; Setoguchi

spans

1985,

the Caribbean,

1910), there

they were introduced or South

platyrrhine

from

record

set of monkeys.

of the evolutionary monkeys

considerably

et (II., 1987; Ford & Morgan,

1982; Setoguchi,

This

MacFadden.

of a diverse

of the modern

Holocene

Antilles

& Woods,

1984;

has grown

1983; Fleagle

et al., 1988).

1987; Setoguchi

(Marshall

monkeys

& Bown,

et al., 1987; MacPhee

Kay,

and

of platyrrhine

1990; Fleagle

new

important

monkeys

from

by island and site

Hispaniola C’ueva de Berna: Hispaniola. place.

“Saimiri” bernensis. Three primate specimens have been reported from The first (CENDIA 1) is a fragment ofa right maxilla with P”, M’. and M” in

plus the alveoli

for the anterior

two premolars

and part

of the canine

alveolus

(see

al&&tine

SOUTH

Figure I. &lap of

Figure

3). This

Dominican

specimen

Republic.

specimen

of the trigon

and the small

the Hispaniolan size.

in extant

specimen

Several

workers

have

Suimiri, and that it may be a cebine Woods,

1982; Rosenberger,

(UF

28038)

with

de Berna between

at the eastern the molar

dated

at 3860 +/-

with a separate

suggested

than

it should

phyletically

more

135 yrBP

its Saimiri-like

any known

(Rimoli,

of

1977).

(especially

traits.

However,

Saimiri, close to Cebus in

not be considered closely

of this

as the holotype

and offset hypocone

to M’ are among larger

end of the

structure

related

congeneric

with

to Cebus (MacPhee

&

1978, 1979).

Caverne Sawo: mandibularfragment. fragment

primates from the Crtater Antilles.

of Suimiri, it was designated

on the molars, is considerably

Gueva

similarity

members

size of M2 relative

ofknown fossil

from

on the great

bernensis; it has been

ofsaimiri

The dominance

body

was recovered

Based

and that found

a new species on M’),

the Clarihhean, showing locations

AMERICA

MacPhee

an extremely

& Woods worn

(1982) described

Mt in place

a right

and the alveoli

mandibular for Py and I’.&

(Figure 4). This specimen was recovered from Caverne Sawo on the southern peninsula of Haiti, the opposite end of Hispaniola from Cueva de Berna. It is dated by “‘C to 9550 +/150 yrBP,

more than twice the age of the type specimen

noted

the close

lower

molar

basis oftheir ancestral analysis, particular

similarity

in size to “S.” bernensis and,

and low crown similar

conclusion

relief,

suggested

that

of “S.” bemtmsis. MacPhee based

its closest

for “S.” bernensis), although

on the general affmities

noting

& Woods

shape

are with

of the

Cebus (the

that these may represent

platyrrhine conditions. The occlusal morphology is too worn to allow a detailed but they note that there are some significant differences from modern Cebu.r, in the marked distolingual projection of the first molar in the fossil.

CARIRBEAN

‘39

PL.ATYRRHINF:S

Fignn- ?. Proposed altrmatiw modes of colonization of primates of the Grratcr :\ntilles, includirlg importation hy humans. travel over a land connection (either a land hridqc or an early vicariant cvrnt). rafting cm mats of\qctation, and hurricane dispersal (hi,qhl> unlikeI\ to hc succrssf~d for an) thing otht.~ than a VPYJ vnall animal). :Zs discussed in tat, dihprrwl h\ raking appears the most rt~as~mal~l~~ ckplanation at prrwnt for prim;ttcs.

.Smnanu Bq: right

tibia

Republic

tibia. The (Figure

human

contact

inferred

body

Hispaniola, closest

published

size

places

it close are

(Ford,

a particularly

the trochlear facet which angled posteromedially

from Hispaniola the Samana

age but

with

members and

deep

of the

is a distal

Bay region

heds

that of the

(Ford, much

and pitheciines); to more-or-less

facet

of a

any

evidence

01‘

198&z). Althou,gh

dental

specimens

smaller-bodied

from

callitrichids.

the tibia shares

on the distal

its

1978). its

198&z; Rosenberger,

1988). In particular, fibular

fragment

of the Dominican

predate

1952; Ford,

for both

Cebus monkeys

198&z; see also Ford, long

from

& Roopman.

to estimates

the size of extant

found to a lesser degree in the atelines trochlear facet which extend inferiorly

platyrrhine its lateral

f rom

1929; Williams

affinities

toSaguinus

the callitrichids:

specimen

254682)

5); it is of unknown (Miller,

or about

phenetic

especially

third

(USNM-Mammals

end

(both

with also

anterior and posterior edges ofthc equal degrees; a posterior border of

is fairly sharp; and a median ridge of the trochlear (the last two features developing independently

facet which is in selreral

lineages). The tibia also shares with Saguinus a centrally placed fibular surface, an anteroposteriorly narrowed medial malleolus (also fijund

facet on to some

240

_A.-----~-/ Figure

degree

3. Type specimen

its presence

noteworthy

the tibia

of platyrrhines

atelines

(although

World

(CENDIA

in all other

callitrichids).

posterior

These

sharing

size (&bus

to it in body

primitive,

general

features

features

groove

while

(as

few, are

that characterize

or the pitheciines)

platyrrhine

view.

malleolar

similarities,

in that the tibia lacks any of the derived closer

I). Ckclusal

or of the larger

found

in most

New

monkeys).

Original

study

callitrichid would

of this tibia

lineage,

isolated

be a secondary

for gigantism

However,

an alternative

several

Hispaniola.

mammals

following (see reviews

as discussed

below,

it may

represent

in Ford,

a descendent

If so, the relatively

a well-documented

recent

Most

In 1986, Charles

Woods

new

from

specimens

of these

localities

trend

larger in island

in Hispaniola

body

of a size

mammals

1980a, 1986a). This remains

discoveries

dentition (lacking

and co-workers Haitian

are

fragment (UF 28038) was found. Dating now being studied. The primate material and lower left femur

that

Antilles.

a viable

lend credence

to

explanation.

new material.

collected

led me to suggest in the Greater

development,

in small

hypothesis.

Haiti:

brrnenm ii-om Hispaniola

and C’ebuella), and the lack of a horizontal

in Cullithrix

in Suguinus, versus particularly

of “Saimiri”

/-----

_----

near

sites Caverne

at the Florida on

the

Sawo,

where

has not yet been completed, includes an almost complete

from one locality, as well as a few isolated only the distal epiphysis), and the distal

State

southern

Museum

peninsula the

of

mandibular

and this material is and unworn upper

teeth, much of an immature portion of a humerus from

nearby sites. Preliminary work suggests that the upper dentition is extremely similar in both size and structure to that of the type specimen of “S.” bernensis from the other end of the island. described

The lower lower

R/I, of the new specimen molar

from

the

area

is also the same size and shape (MacPhee

& Woods,

1982),

as the previously but

the

occlusal

CARIBBE,

.‘,

,.:., .::: _.

il ::

.‘,.

.:

:

“.

I).

..”

;

:.,

‘.

“,:,

,,:,‘;

:

,,

..T,

‘. ..:, .,

.t :.

.. :

:: . ‘.’ ,-..li f .., .;: , *..

.:

A

C

0 Ii::::

4

I cm

morphology latter.

cannot

Pending

be closely

further

study

remains

from Hispaniola

monkey

to continental

new material although

there

not

several

retentions,

have

recent

be closely

or the other

If only

a single

reconsideration

taxon

here

Preliminary to both

Rosenberger

and

Cebus are

the hypothesis

two extant

,111 111~

M any

clmt~rl

The affinities

While

1989).

of wear

all of the primate

species.

sense)

Saimiri

supported

stage

of tlrih

analysis

of thr

Sain~iri and

( 1979, 198 I sister-taxa Saimiri and

that may

and

within

of the similarities

platyrrhines

C‘P/~I~.\,

O’P/UY of “,Y.”

be shared

primiti1.c

features.

affinities

on the femur

is represented

of the

will be analysis

comments

attributes.

19866; Kay,

of these

that

platyrrhine general

that

have

of monkey

of the

importance

of unique

derived

it appears

are not as yet clear.

suggested (Ford,

of the advanced

a single

(in the most

studies

related

not shared

material,

lineages

similarities

others

bernensis to one

Some

platyrrhine

are also a number

and

Platyrrhini, may

of the new

because

may represent

indicates

elsewhere)

compared

dentally

similar-sized

of the structure

will be made

across

tibia1

Hispaniola,

fragment

of the new postcrania

below,

after a discussion

then

a

is necessary. from

01

the island.

of the material

from

J amaica. J amaica Long Mile C’uz~e:Xenothrix the smaller the

Greater

northwestern

recognized known

area

a human cases,

specimen,

extinct

Antillean

platyrrhine 1982; Morgan

third

molar

placing body

of the

(Williams

(AMNH

fauna

first

recovered

6). Although

W’illiams

(Rosenberger,

1986). The

It is slightly slightly

canines,

smaller smaller

than,

endemic

8r ILlorgan, fragment

well

and, in

data

for the

that this is indeed 1986; MacPhee

preserves

the first

it is clear

that

and incisors;

Cebus (Rosenberger,

than

to be

AYenothri.y is

in 1920 from strata is no absolute

1986u: Ford

from from

primate

inference

mandibular

for the premolars,

developed.

there

remains

1952).

with other

and Koopman’s

1977; Ford,

& Woods,

Antillean

& Koopman,

148198)

(Figure

all confirm

it close to, but probably

The

molars

are bunodont

a larger

premolar

alveoli

and

incisors

the

dentition

which

(Rosenberger, Hispaniolan

with

hypoconulid. increase

large

cusps,

The molar

teeth

in size posteriorly,

somewhat

procumbent.

has, to a large

on the deep

canine alveolus, He has recently new

source

primate

is Long Mile Cave,

the Hispaniolan

1977),

dental

the thus

remains

in

size.

include

Based

the

have yielded

The first of these

in the cave, and it is associated

and the alveoli

never

localities

mgregori

fragment

workers

& Woods,

two left molars

island,

deposit

an endemic

Three ofJamaica.

Xenothrix

midden

recent

island

of the

as endemic:

from a mandibular

below some

mcgregori.

Antillean

and expanded

part,

on the wide

and the alveoli This

rendered

gonial

especially

are large relative mix

region,

suggest

of traits

its affinities expanded

with

trigonids,

and the)

to the size of the jaw, the canines

gives

Senothris

any living

premolars,

The

were small

forms

a unique obscure.

and relatively

small

Rosenberger (1977) suggested affinities with the titi monkey. Cullicebus. expanded this to include ilotus as well, which two he regards as sister-taaa 1990).

However,

material

and

small females

canines

occur

of many

in other

sexually

platyrrhines,

dimorphic

including

platyrrhine

the

species.

And expanded genial regions and larger premolars have developed several times in primates (see Szalay & Delson, 1979; Fleagle, 1988), generally in large-bodied forms with more frugivorous/folivorous diets. Their presence in a form with an otherwise highl\~ individualized, complex molar structure may not be a good indicator of phylogenetic

CARIBBEAN

affinities.

and these

similarities

convergence. The lack of a third

molar

of the teeth

with

exception

the

possible

of the symphysis

aspects strongly

suggests

callitrichid independently sequence

and tibia

of development

bodied,

insectivorous

from

and subsequently

almost

have

certainly (Figure

7).

development mandible, expected

so that to

platyrrhines,

undergone

radical

have

teeth

which

is precisely

newly

noting became

in association dental

would

reconsider

the idca

that

that the theory dental structure

gonial

region

in

lineages

in hod) the

re-

of the

dentition. would

1

not bc

of frugivorous

for Xenothrix.

that the structure

a callitrichid

include

descendent

in the

its diet would

in its modified

other

if a small-

isolated

the change

callitrichid

found

features

that

could

of the

be obscured

those

the situation

with

structure

expansion

lost

an alternativt

body size increase,

of the gonial

incisor teeth, and the probable presence of a hypocone remain any callitrichid afinities. \Yhile new material from Hispaniola gigantism to a &bus-sized isolated on small islands,

suggested

(Suguinus),

in

the

was

ofcallitrichid-like

1986a)

significant

unlike

molar

of Senothria,

frugivorous

like

molars,

third

morphology

for example,

precisely

et al. ( 1990) argue

I (Ford,

changes

heritage

an enlarged,

the

of some

the wear pattern

as a tamarin

changes and,

that

with the recognition

underwent

its callitrichid

that

Rosenberger

such

Complementary

of a hypocone

also suggested

Later,

interpretation

on the upper

argued

Hispaniola,

but little elst and tamarins,

To the contrary,

was present

(1977)

of the unusual callitrichid,

Greatcar Antilles size

hypocone

Rosenberger

(1952)

alveoli.

functional

to callitrichids,

any of the marmosets

& Koopman’s

of the incisor

two lineages.

large

similarity

resembles

of Williams

and angle

in these

in the relatively

phenetic

or mandible

that a fairly large

pattern,

Callicebus and .Yenothrix may represent

between

is an obvious

in the morphology

“-13

PI.ATYRRHINES

lineage

reached

that

region,

the size of the post-

strong arguments against is currently causing me to

island

and

there

underwent

form, I feel the argument is still logically sound for platyrrhines given our knowledge of patterns of change in primates, and

provides a possible on *Jamaica and

is

scenario worthy

for of

the origin of the unique .Yenothri.u further consideration. LJnder this

Early tamarinlmarmoset (approx. 500 gm)

_
‘1

._@>L,,;,;~8 ,f ‘I i r’ --. GiY

Ancestral Platyrrhine (approx. 1000 gm)

Island tamarin descendant, result of gigantlsm (approx. 2500 gm)

scenario,

those

convergences Several

few

postcrania

Xenothrix mandible

structure

appears retained

which

Senothrix described

Mile Cave. belonged

in the proximal

with

Callicebus

would

represent

niches.

as coming

MacPhee

Some of these postcrania to have

shares

filling similar dietary

were originally in Long

study of this material. particular

traits

in platyrrhines

from the same strata

and Fleagle

are currently

may indeed be primate.

to a very unusual

primate,

portion (for those areas preserved),

as the

completing

A partial femur in

with a conservative but a highly derived

distal epiphysis. For example, its short, fairly robust shaft ends in a distal epiphysis most reminiscent of the distal femora of sloths. If this femur belonged to a monkey of the same species as the Xenothrix mcgregori mandible, was a very distinctive primate indeed. Two other

caves on Jamaica

and they are very similar in body size, .Yenothri.x

have yielded

primate

remains,

in both cases proximal

femoral fragments close in size to the remora of extant Cebus and Chiropotes. \Vith at least three specimens of primate postcrania known from Jamaica and three from Hispaniola, it is remarkably fortuitous for purposes of comparison that four of the six are femoral fragments preserving some or most of the proximal epiphysis, including all three 01‘ the

CARIBBEAN

,Jamaican

specimens.

nothing

This

of the structure

comparisons

narrow

record

of other

is disappointing, portions

in base and functional

studies

The first of these

583.50) from Sheep

Pen Cave,

located

Study

from this site has just

of the site is difficult,

but the remains

older

tllan 50,000 yrRP.

ifnot

older

Ii-cm

the Ciaribbean,

(Goodfriend,

100,00(1-200,000 Miocene

of the islands.

The specimen

small,

It is distinctly

Both

its affinities

within

right

of the island. femur

:30.000 \.rBP

(Figure

The third

9) and

than

& Morgan,

the limits

theoldest

known

.Jarnaican

some

mammalian

in particular

to

the probable

of ‘“C: dating,

a combinatiori

primate

the functional

femur

8). thus

100,000 ).rBP old mammalian

site dated

predates

( LIF

(Figure

1988). Dating

hair human

due to its especially

trochanter,

fragment

Mile Cave

may bc at least

1988). and it clearly

and

making

femoral

another

1989) and

lesser

little

fossils between

preserved

in

occupation wide femoral

not l‘ound in

;uI\.

from Long Mile C:avck.

significance

of its structure

arc

1990).

The Coca

(Goodfriend

unlike

platyrrhines

(see Ford,

~AKOReu (,‘azre:pro~imnlf~mur. interior

(Poinar,

displaced

platyrrhine. at present

older

from

is very unusual,

prosimallv

other unclear

et hl.,

(MacPhee

(Ford

that the femur

1986). It may then be among

from Hispaniola

right

2.5 km from I,ong

are clearly

we know primates.

verv difficult.

been completed

to material

in that

of Antillean

is a proximal

only about

suggests

in addition

yrRP

amber

neck and

Evidence

however,

of the skeleton

S’/zeep Pen CUZV:pro.~inxzlJknrur. of the femur

245

PI.ATI’KRIiINES

primate

Ree specimen

was

recovered

& Mitterer,

1987).

locality

oneJamaica

(UF 40097) from

deposits

is C:oco Ree (Love, in the

is also a fragment dated

to between

ofa proximal 30.000

and

246

5.

FORI)

Irl.

c,

Figure 9. Proximal posterior; c, lateral

This femur

right femoral fragment from Cow Ret Cake. Jamaica views (from Ford & hIorgan, 1986).

is very distinctive

1988, in preparation). smaller,

dwarfed

from the other

Its closest

callitrichids

phenetic

from the mainland

specimens

this view. One is the presence

the femoral

neck,

ofilotus,

found

primarily

Aotus, and Suimiri among size than small

the Coca

represents

within

While

state ofthis

Brudyteles

ofsome

ofthe

and slightly

& Morgan,

1986). Several surface

reports

of

Cullicebus,

the medium-sized

1987), all somewhat (1988)

much

less so to

ridge on the posterior and

Hershkovitz

may indicate

& Morgan

(1986)

is a narrow, the shape

examined

increased that

smaller

finding

in hod)

this ridge in a

from that seen in UF 40097. There

consistent trochanter.

in most

in the British

by Hershkovitz.

leaping

oval area of insertion

1988), it is fairly

present (Davis,

this is not a strong

of this area varies

vestige

of a third

feature

for this feature,

in the two skeletons

this feature feature

of a distinct

(see also Davis,

Recently,

(set Ford & Morgan.

of Cebus updla specimens. He reports a less well-developed rid,q:e in sonic or slight ridge in this region also equivalent to either the “low mound” for some pitheciines and other platyrrhines by Ford (198Ob, 198661, which a less derived

diversity

Ford

America,

t).

percentage

pitheciines, described

lacking

of South

in the callitrichids

platyrrhines

Ree femur.

femora

is to the remora

Callicebus, and Saimiri (Ford

some individual features

support

two Jamaican

resemblance

(UF 43097). a. anterior;

genera

Although

1987), Hershkovitz indicator

in some examined.

Hershkovitz

as present in UF 40097, this is due to dillerrnces ridge is present, but not a distinct, well-defined

but

suggested

that

(1988) concurs

with

of locomotor tendon

The (1988)

third

pattern.

on the lesser

platyrrhines

is also some

specimen,

WThile it has been

of the iliopsoas

somewhat

Museum

A second trochanter.

(see also Hershkovitz. feature

describes

is the loss

the third

of an)’

trochanter

in describing and coding of this feature. I\ third trochanter such as occurs in man).

specimens of ,4otus, most strepsirhines, and Eocene primates. While none of these features are unique to callitrichids, the callitrirhids arc the onl) other platyrrhine group in which all occur regularly. This is particularly interesting given

CARIBBEAN

the similar

finding

of thts traits

listed

closely

resembles

singlt.

specimen.

identical

for the Hispaniolan above

PI.ATTRRFIINES

tibia.

in a very small

Although

Th e newly

in size and

recovered

comparable

species,

not affiliated

similar-sized

postcranial

f‘rtnul, from Jamaica Rcc femur must

and

developing

tocotnotor

lineage,

from that island

Bay tibia

convergence, behaviors

mentioned Ree femur.

and ofthe

to callitrichids

similar

which

possibly

phenetic

more

the fhssil

together

in an!’ appears

Pending

detailed

expressed

conspecific

to a oft ht.

affinities

Coca

similarities

so than

due to a monkey

to those

each

above

the phylogenetic

The general

presumably

has found

represent the same species. from Hispaniola proves to belong

with any callitrichid

the Samana

femur

to the Coca

need to be re-evaluated.

then represrnt

of Ce6uJ specimens,

Hispaniolan

in structure

material

(1988)

rare in Cebus and do not occur

stud\-. it appears likely that these two specimens As discussed above, if all of the dental material single

Hershkovitz

percentage

in size, most are estremely

2-17

Ret

of the C:WX)

to other

platyrrhines

of the general

size of (,‘rb~

in some callitrichids

(SW Ford,

t !W) Cahu Cuezz de Boca de1 Purial: (MAhl-ULH U nivcrsidad

de La Habana)

Alontaneia Purial,

anthropomorpha

Provincia

teeth

the

Cuba.

except

substantially Ate1e.s species of the other scbparated

recovered

from

all subsequent

have

1986a,

Based

to those

assumed

that

brought

MacPhee

del

remains,

were very similar

of .4teLes, probably

Ford,

de Boca

human

little or no fossilization.

researchers

species

e.g.,

Cueva

with prehistoric

that the teeth

de 1a

of the

over

& Woods,

b!.

1982:

1952). of

an

& \‘arona

(1983) suggested

endemic

Cuban

indicated

specimen

that

consists

for the right

in the following

features:

bones

of

of this

spider

tason

mandibular

left M:+. Arredondo

ofextant

and molars;

from it b!. a tnarked

other

that the teeth are specificall)

species

of 16 isolated

1, and

to the teeth

premolars

were

of teeth

de Biologia

( 19 IO) as a new genus.

by Ameghino

to have undergone

(see,

A collection

de la Facultad

association

of an extant

Arredondo

similar

10). They

almost

mainland

‘l’hcy

‘l‘hc t!.pe

dentition

the

remains

anthropomorphu.c).

reported

in direct

appearing

the remains

howe\;cr,

distinct,

Montane

(I 916), who suggested

& Koopman,

Recently,

Figure

Ecuador,

from

anthropomorphus.

Antropologico

Spiritus

condition,

are

Amerindians

(see

of Miller

.Ite/es ,/iucicep.r from

LVilliams

(=Montaneia)

was originally

de Sancti

and are in excellent on the analysis (Cuban

Ateles

1376, in the Museo

\‘arona

monkey

“escotadura”

of M1_g is elevated

(ectoflexid)

buccally;

argue

that,

on

louc~~

whilta

from all knolqrr

exceeding above there

(.i/r/u.c fi)und

entire

differs

P, is very wide buccolingually, the trigonid

been

teeth-an

and

Atelex, the Cuban

monkey

have

the width

the talonid

and

is a well dcvelopcd

on hII_,; hl,i has sis cusps arranged around a central fovea; Ci-hf.c Icngth I ikt 38.5 1111n) is greater than that of other .4teleJ. ‘I’hc r\idence presented by Arredondo and \.arona strongly indicates that the teeth do

tiypoconulid

rt’prest’nt

an

endemic

Cuban

species

of spider

monkey,

Pinar de/ Rio: howler monkey .skuU. A well-preserved

.-Iteles anthropomorphu.c.

skull (Figure

11) was recently

discovtarrd

in ;I C;IW on Cuba, at Pinar de1 Rio, west of the Cordillera de los Organos, along with a hutneral and a femoral fragment (River0 de la Callc, 1988). ‘l‘hese specimens are beirlg prepared and studied by C>scar Arredondo and Manuel Rivero de la Calle. ‘l‘hcir preliminary

report

A/ountta. hut much

indicates largrr

that the skull is extretnely than

an); known

species.

similar ‘l‘hr teeth

to the living howler are clearly

monkey,

.Ilouatta-like

and

s.

A

hl.

IOKI)

CARIBREAN

I

I

I

very

diKerent

from

anthropomorphus).

monkey.

the

In the

absence

occupation

of the island,

was viewed

with skepticism.

that a primate

presence

on thca Yucatan ol‘thc

to cross

in isolation Cuba

on Cuba

from

which

presence

specimen

(Ate/u\

similar quite

howler

that

means

monkey

and surrounded

one can only

developed

the known

of both

speculate:

range

rsist

and both

from

& Llioods,

unique

species

and .lteleJ

end of&ha,

different

by the narrow

both

to human

skull indicates

representatives

& ‘12’oods, 1982; Morgan

1986), they

prior

Roth .4iouatta

to but sli,qhtly likely

of howler represent

as an endemic

close to the western

(by what

1986; MacPhee

on Cuba

coincidental.

is quite

barrier

primates

Cuban

ofAte/es nnthropon1orphu.r

the newlJ.discovered

are very

& ‘I\:oods,

j

0

setting.

the water other

other

of platyrhine

It appears

(see Morgan

of the

I

preliminary

is not simply

today,

7

in an island

monkeys.

I9860; Ford & Mayan,

teeth

1

!

6

belirlre it is a new genus this specimen may views,

However,

on Cuba

American

managed

evidence

I

5

de la Calle

the interpretation

Peninsula

two specimens

Central

supports

I,

I

4

River0

size increase

of other

I

I

3

monkey

and

study

case of phyletic

I

I

2

spider

Arredondo

If further

another

I

I

I

cm

l’I.ATYRRHINkS

genera

set

1986)

of fauna

Ford, I’herc,

nati\.c

to

attributes.

Discussion Depending

on the relationship

to oncb another, endemic tnore

of the various

and on one’s interpretation

Antillean

(Figure

primate

12). Certainly,

species,

very probabl)-

a minimum

of three

ofapproximatel>,

the same body size: X’enothris

(:ave

Pen Cave.

and Sheep

Hispaniola,

possibly

specimen).

However,

“Suimiri”

conspecific

bemensis

with

the general

Hispaniolan

of the Cuban

and postcraniat

species

and the two round, may have of some

been

as many

occurred respectively, species

of the Hispaniolan

as seven

on.famaica,

ol all

in Coca Ret

the only primate

Jamaican

remains

there are at least thref

five, and possibly different

one of the three

similarities

dental specimens,

present

on

(the. COW Ret, postcrania

to

callitrichids and ofthe dental remains to both C’ebw (in some features) and Siz~nimiri(in other features) leaves c)pen the possibility that, despite the ,general similarity in body size Ihr all of the Hispaniolan

remains,

.Jamaica.

conclusion

A firmer

more

than one primate

for Hispaniola

must

species ab-ait

is represented, more

complete

as is the cast’ on stud!,

of the nt’~

250

Endemic Antillean Primate Species Possible

Most Likely Estimate

Xonothrfx Shnp

“glani

material.

Although

it has

(or human

researchers

strongly

suggests

The

various

specimens

groups.

While

the ancestry

obscure,

others

Callicebus, primate

possibly

lineage

Specifically

of other

Neogene the known

inhabitants relatives There

of the various are currently

radiation

ofplatyrrhines,

existence

ofNew

early

anthropoid

Although

evolution geology

current

Cuba

howler

monkey

Cuba

are

current

Pen

simply

work different

femur,

mainland

remains

indicates

recent

by Cuban

species.

to several

pattern

is open

that

completely possibly

more

Antilles,

than

to one

since

However,

there

lineages

1980; Schlcc,

(Fleagle,

the

evolved

earl)

diversified

is no evidence prior

that

b) the

to the earl!,

of New IVorld

monkeys

of the earl?,

1988, 1990), does allow for the

than this time, but there earlier

of South

and there

among

1990). l’he diversity

of platyrrhines

of the dating

not

are already

the exislencc

(see MacFadden, earlier

were

of

1988), along with

1987: Rieppel,

the primates

platyrrhine

in Argentina

(whether

The presence

(Poinar,

primates

indicating

slightly

islands

prior to the Late Pleistocene,

that

record.

Antillean

Antillean

to speculation.

& Cannatella,

were present

for the origin

understanding

Hlspanlola

anihropomorphus

from Hispaniola

(Poinar

no fossils

monkeys

Jam&a

Atales

two endemic

the Greater

to assume

particularly

World

the

understood,

amber

Miocene

for any argument

our current

important,

and Holocene

to late Oligocene/early

given

while

of the Greater

callltrlchld”

islands. reached

reason

Hlspanlola

Saimiri, to A4teles, CO-Ilouatta,

‘rhis

primates

that mammals

Pleistocene

Miocene.

at present

to Cebus and/or

bemans/

specimens

as the Sheep

callitrichids.

to Miocene

no compelling Late

such

land vertebrates

indicates

mainland prior

amnities

in origin),

the remains is currently

represents

phylogenetic

Jam&a Jamaica

“giant callltrlchld”

species,

the material

the Caribbean

hairs in Eocene

Cuban

that

how

mammalian

the

exhibit

to the

and

or dispersal

that

of mainland

of some,

colonized when

1984), clearly

argued

have ties variously

and

vicariant

been

introductions)

mcgrogo,,

Pen femur

“.!wmbl”

colonizations

Estimate

than

American

is little or no basis

mid-late

Oligocene.

fossil localities

and of

comples

poorl)

and origins. of the

reconstructions

Caribbean

basin

all suggest

is extremely

isolation

ofthe

Greater

Antillean

and

islands

in

the Caribbean basin early in the Cenozoic (Burke et al., 1984; Dickenson 8r Cone),, 1980; Donnelly, 1988; Hedges, 1982). After this time, various of the islands were periodically at least

partially

Hispaniola

submerged,

appears

to have

with

areas

had

a hybrid

of high origin,

relief with

probably central

and

remaining

emergent.

northern

Hispaniola

connected early in the Cenozoic to eastern Cuba and possibly Puerto Rico, but southern Hispaniola connected at some early point with Jamaica and probably western Cuba. ‘l‘he southern Hispaniola/Jamaica/\Vestern Cuba ,group may have maintained a somewhat

CARIBBEAN

later mainland Eocrne

contact,

(Donnelly,

1988;

Rosen.

connected

to each

The

distribution poeciliid

of these

1988; Hedges,

t 985;

studied

but none

other

of several

groups

taxa

any direct

land

1980; Morgan

connections

mainland

was complete

bc made

for the presence

the Paleocene presence model

with

& N’oods.

in the Greater for various

members

present

in the Greater

Antilles

through

rafting

then and the present. in particular the Long

,Jamaica,

are highly

distinctive,

either

a long

colonization,

in particular

mainland

rclativcs.

,.\ntillean

isolation

The generally suggest

in an island

‘l’he fossil record particularly

shows

on Jamaica,

evidenced

by the Coca

Hispaniola. emergent

As there

their proximity

island

oforigin

material. rapid

Thus,

that

there

and

at least

reached separate

cave

specimens

or both. little

change

of the Cuban

hJn1

ties. This suggests or even

earlier

In contrast,

othcl

changed

and clear tendency

evolutionary

has been

Ree femur

limited

both

prirnate

movement

from ,Jamaica

biogeographic

between

and

and

from

thril

to endemism

commonly

primates

on

occurs

indicates

Hispaniola

and manner

diversification between

the very

geologic

and Jamaica

etal., 1983: Mattson,

(MacPhee

of this monkey

primates

a Pliocene

relative]),

probably

and on several Pen

after

in thcil

lineages

dispersal,

1981). The

on Jamaica,

appear

the nature

to explain primate

phylogenetic

Antilles

can

America) of a vicarian

the various

Sheep

from the

Mitt)

the\. arc

Pleistocene.

is strong

land connection

and

2lMl

a vicariant

argument

event

via overwater

low fauna1 diversitv

perhaps

Thus,

ultimate

once isolated

that

setting.

colonizers,

Mile

oftheir

in the Greater

evolution the Cuban

islands

fairI>, recent

despite

of isolation

or very rapid

specimens. the

period

regardless

reccantl)

taxa

(or Central

than mid- to late-Miocene

between primates.

the

islands

of the appropriateness

the islands

occasions Some

ncithrr

1986; MacFadclen.

no reasonable

1986~~; also Pregill,

not earlier

were

in the (Caribbean.

ofthe

America

fauna.

the

appear to originate after the breaking 01’

for a vicariant

regardless

in Ford,

probably

others

isolation

in South

Antillcan

colonized

among

& Morgan,

and at present to allow

islands,

bridge.

of the islands

(see Ford

primates

ofthe

islands

land

the primates, of the islands,

1980). Certainty,

less earlier,

(see discussion

Antilles

the mainland

Antillean

other

the &cater

isolation

and in particular later cotonizations

ofplatyrrhine

beyond

a number of insect 1988). snakes (Kluge, 1988), suggests

by the early Cenozoic. much

including

much

1984; Rauchenbrrgcr. the

by any emergent

predating

1986; Simpson,

or Eocene,

to this,

of animals,

However, most mammalian groups, l’rom South America and to represent

have persisted

1988; Mattson,

Subsequent

(Rauchenberger, 1988), and boid

(Liebherr, IVest Indian

would

1982; Liebherr,

nor to the mainland

fishes

onychophorans origin ti)r these

contacts

et al., 1982).

Sykes

2.5 I

PI.ATYRRHINES

ofdispersal

within

islands. similar

evidence

new that

islands,

‘I’he 1attc.r is femur there

liom

was

no

in the mid- and late-Cenozoic,

1984; Morgan between

8r 12:oods,

the two islands

1986). thta is unclear

at prf’scn t. ,Jamaica primates families

is especially

interesting

in particular. ofany

ofthe

This

Greater

in the composition

island Antillean

has

the

islands

smallest

of its mammalian number

and a smaller

fauna

of non-bat

mammalian

and

iry

mammalian

fauna

than would

l)e predicted by its area considering both the extinct and extant taxa (Morgan Cyr\l’oods. 108ti t 11 has bren considered a depaupcrate backwater of Antillean evolution. with a largeI! trndemic fauna notably lacking in both sloths and insectivores, otherwise tile Greater Antilles (ibid.; MacPhee et al., 1983). H owe\.er, it has the greatest primate bcvn

species, conver,qin,q

at least three, on

the

including

absent

sloths)

two highI>, distinctive and

a possible

forms migrant.

(one which The

present number

in 01

may ha\x-

diversit\,

and

diversification

ofprimates

of competititon South

they

and Central

their

America,

phylogenetic

In addition, monkeys small diet

and/or

better

directly

to the lack ol‘other

platyrrhinrs

very

two at least becoming

the wide range 1988),

were about in which

tropical

into

di&rcnt

so highly

l~luna. In the absc~lc~,

from

those

modified

known

li.cnn

as to grcatlb- ol)scurc.

origins. despite

to I5 kg; see Fleagle, manner

may well relate

diversified

the same

estimates

size, falling

these co-existing yet

islands

understanding

of the primate

evolution

and

within

adaptation

monkeys (Ford

on Jamaica on this

Hispaniolan

around

avoided

12-3 kg. ‘l’h( competing

on

some diffrrences

in

& Morgan,

1986,

will give a much

fascinating

IX.> gm

(from and

likely it was through

the forest

radiation

Jamaican size range

distinctive

but most

levels

platyrrhines

all three+

in a very narrow

obviously

is as yet unclear,

in use of dimerent

of mammalian

in hod!, size in mainland

by current

island.

1988).

clcarrr

and

.Z

picture

on islands

in

general. In

conclusion,

indicates

the

without

far more varied

record

of fossil

that the primate

and complex

taxa are still known phylogenetic

current

a doubt

than would

from only single

affinites

and

known

groups,

in particular

features

suggesting

niche

a howler possible

monkey

(but

Gigantism

isolated

island

fauna,

may

in the Caribbean.

probably

occurring

presence

of primates

in an island

have

occurred

Overwater

several

separate

times,

the

while

others

Ree femur

and some of the

among

during

is currently

that their

ties to specific Cebu.5 or Sclimiri

times

sometime

and

ago. Most

derived

clear

a well-documented

or more

dispersal

in the Greater

(the Coca

Antilles larger

with

relationships

setting,

one

have

an .dteles sp. on Cuba,

and

(.Ebnothri.s), or callitrichids

Greater

was much

less than a decade

Others

not certain)

(“S.” bernensis), Callicebus in other

the

islands

and some are so highly

are obscure.

Hispaniolan

postcrania).

from

on these

have been predicted

specimens,

adaptive

exhibit

platyrrhincs

radiation

phenomenon the

or after

best

platvrrhinrs the Miocene.

explanation

li)r the

Antilles.

Acknowledgements This

work

would

preparation Morgan, John

not

of fossils, to whom

Fleagle,

Thorington,

indebted.

Kinzey,

Ross

Jr, as well as the comments I am grateful.

Kimberly

Martens, Vercillo,

particularly to forge

without the contributions, through acquisition and collaboration, of Dr Charles Woods and

discussion,

I am deeply

Warren

for which by John

be possible

thank and

Figures

Figure

Drs Fleagle

present

this

Alfred

of Dr Arredondo

I thank,

of Ross MacPhee, as well as Less

and Rosenberger

overview

in their

symposium

Primatological workshop, and

and

for financial

Southern

Congress.

Illinois

University

and

by Karen

Schmitt;

and Figure Davis

Figure

3 by

assistance.

the opportunity

on platyrrhine

it:.

reviewers,

8 was photographed

for technical both

with

Richard

and two anonymous

for providing

XIIth Congress of the International interchange during the associated

from long discussions

Rosenbergcr,

1, 2, and 7 were prepared

4 is courtesy

all of whom

It has also benefited MacPhee,

and Gar),

evolution

1

for me at the

Society and the forum for some liveI> I thank the L.S.B. I,cakey Foundation

support

which

allowed

me to attend

the

CARIBBEAN

PI.ATYRRHINES

25:i

and stratigraphy: I’lrleor,rrtrbrc~ta.

Ger1chrcmol0~~ .tlumo~rr

L~tmurdinczire.

of

thr

c0ntinrnt.d

m;lmmal-h~,,lrin#

Qu.~tvr-~~.rt~) 01 S~~rltll

.\IIII,~ 1, ,t.

76 pp.

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.Serie C, Sr.

18.

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