Tarsiers, adapids and the integrity of Strepsirhini

Jeffrey

Takers, adapids and the integrity of Strepsirhini

H. Schwartz

Departmmt oj Anthropologv. 1 ‘niuersit_~oJ’Pittsburgh, Pitt.rbur,~h. f’,4 /5X0. l’..C..4.

.L\lthough strepsirhinc primate can hr dcscrihed by their narial conligyration, this and most utht.1 drlinahle fcaturcs are prohahly primitivr rctvntions; only thr dcvclopment ofa groomin!: claw of the second pedal digit and uf a twthcomb j the lattrr of which has been lost in Daubentonirc) cmrr,qc as potential apomrlrphirs III’ the group. Within this assemhlagr tunurids. Lepilemur. the indriids. ancl Daubentorriu can he argued tr, constitutr a monophylctlc group whosr rrtationships cladistically arr in the qucnce listrd: Lrmuridar* .md 1ndriid.x can themselves hr detineatrd as mwwphylrtic ,groups. ‘l‘hr rrmainingstrcpsirhine primates-the cheiroqalcids, galagids. and Iurisid-also appear to constitute a dctinahlr ctadc, with the fijrmrr qrwp rrprecnting the sistrr taxon ofthr latter two families: cath family C;LII bc united on thv hasis of distinct synapomorphics. Although thrre art’ fcatuws --up&A! of thr car rqion-which prrsrnc thrmsclvvs as putcntiatl\ wllrctivc 01’ thr Gstcr relationship of ‘Tarsu\ + A\tlthrupoidca, othrr charS~cters. including the possession of the grouminq cI;tw. arr suggrstivc ofnn altcrnativc scheme,: 7hrrzu.s may he tht sister ofthc extant lorisiform q,aup. thrrch> rccclnstitutinq. albeit in a novel form. thr pr imatc suhordcr Prosimii It also appears that fossil “tarsioids” may in fat I tw ~nm~closcl~ rrlat~d to the extant lorisiforms than tu ‘Ihrtiu\. .\ ru unsidcration of thr v-~~.tltctl fkil Icmurs. thy adapids, kads to the oulclusion that :ldnpl\-lilr pr irn,itch XC’ ‘1 clad? apart from Pe!>,-udu!. .\rjtharrtu<. .Smildrc te\ ~ntt their most irnmrdiatc rrlatives. and Ina, thcmselvcs constitutr .I cl,~(Ic that ib rclatrd as the primitive sistrr to all other “prosimianb” b\ 1 iT111(. 111‘the, rt~~\~&,pm~~rlr of IhP so-called frr? lnrl-dhullar t! rnpat~ic I iy

Ian Tattersall

Rrcrivrd 30 Sovemlxr 1986 Rrvision rcrcived 2J June 1987 and acqtcd 25 ,Junr 1987 Publication datr Kuvcmhcr 1987

Introduction The

decade

Primates

of the

into

Strepsirhini course,

1970s witnessed

Prosimii

and

and

Haplorhini

in conxnsus

alzay

Anthropoidea

and

the

(c.g.,

1975: Szalay.

was the reinterpretation

same

period,

hlalagasy concept

doubt

began

the Eocene

Luckett,

towards

of the relationships forms

adapids

(e.g., Tattersall were

in some

\iew.

and groups

recent

to survey

re&ws

the morphological

of primates. as Aiello

and tabulations

Lllith regard

cvidencc

(1986) and MacPhec

of features

in apparent

Relationships

support

to this shift, During

of Strepsirhini

ancestral

of into

as a group

of the into the

to thr

(e.g.. Schwartz & ‘Tattersall, the evidence for these points of

WC have referred

of‘this

order

1974). Most recently,

fGr the relationships

Br Carrmill

of the tarsier.

division

\vay hroadly

to Haplorhini,

the subdivision

1973). Crucial

& Schwartz,

extant strcpsirhine primates has been subject to question 1983). It may thus be appropriate at this juncture to review these

from

division

of the enigmatic

to be cast on the neat geographical

and non-Malagasy that

a shift

among

and within

principally

( 1986). which provide

to such

summaries

clad?.

among the strepsirhines

The first question

to be addressed in this context ix whether Strepsirhini (i.e., “lemurs” + “lorises”) does indeed constitute a coherent clads., and if so, what arc the relationships within the clade? Based on thr analysis of sc\xxral studies investigating various different aspects

of morphology

0017 L’184/87/010023

(Bugge,

+ 18 $03.OCl/~l

1974; J ou ff ray P/ a/..

1984; Luckett, 0

1974; Cartmill, 1987 hcadcmic

Prrss

197.5; Lirnit~~d

24

.I. H.

Ford,

1980;

Tattersall, have

Hofer,

in previous

the

group,

hypothesis The

and

most

obvious

feature,

as the basis

which

are

anterior

teeth

the presence However,

a broader

a broad

comparison

and thus of no utility

the group

include

developed

occipital

Table 1

the presence extension

Distribution

lemurs

of a prrpollex, of the

cerebral

of various morphologies

hnur’s

l&i.

Daub.

purpose.

loriscs

Our

reveals

laterally

lemurs

these

median

an elongated

fourth

hemispheres,

pedal

among extant strepsirhines

Galagids

with

to bc primitive rctcntions

digit,

and

also shart

promincnccs.

primitive

short

Cheiro’s

the loss

latrrally,

nasal

characters

Similar

is the by this

comprcssrd

and loriscs

slit or creased

this clade.

Indriids

united

Unquestionably,

growing, Thr

as a whole

Of the taxa

and unfused

mammals

in recognizing

&

1) that

of relationships

for that

a toothcomb.

i.e. the nostrils

intcrnarium,

among

and

of the foot.

of this genus.

of the nares,

hypotheses

Schwartz (Table

1.

the digit

1984;

and states

again

of continuously

an autapomorphy

of a rhinarium,

retentions,

uniting

acquisition

configuration

Stephan,

once

form, Daubentonia, possess the

represent

the “strepsirhine”

hcrc

in Figure

claw on the second

and

1986;

for establishing

employed

is represented

all but one extant toothcomb

& Cartmill,

a table of some 45 characters

synapomorphy

of a grooming

AND 1. TATTERSALL.

studies

of relationships

presence of the

MacPhee

1985), we have compiled

served

within

1980;

SCHWARTZ

broad

ii)r

moderately olfactory

and Tarsius

Lorisids

Tartiur

Grooming claw: 2nd pedal digit 3rd pedal digit

X

X

X

0

II

0

0

Toothcomb

X

X

x

X

X

X

X

Ectotympanic: aphaneric phanrric fused/unfused tubular Position

X

unfus

carotid

Promontory

f.

artery

Stapedial

X

arter)

unfus

postlat.

‘lat.

‘lat.

‘lat.

sm.

sm.

sm.

sm.

k.

vrst.

k.

lg.

unfus

i-us. postmed.

sm.

vest.0

sm. \‘i%t.0

Ascending pharyngeal artery (pharyngeocarotid anastomosis)

0

0

0

X

Foramen lacerum exposed

0

0

0

X

0

Middle meningeal a. fed via ophthalmic a.

0

0

0

X

0

Anterior cavity

0

0

0

0

accessory

Tympanic

cavity

Annular

bridge

Mastoid

cavit)

Epitympanic complex

lg. X

sm.

1% X sm.

lg. X Sill.

1% X k

recess: 0

0

X

INTEGRITY

OF

i t2.1

STREPSIRHINI

Table 1 rontimed Lemur’s

I‘@.

Daub.

Indrilds

(:hciro’s

sub= srp.

shal./ flat

Galagids

Lorisids

sub=

sub=

Xl’.

SC,‘.

Aal./ flat

shal./ flat

hr.

mod.

mod.-mk.

0

rnlld.

mod:mk.

7hrTlur I;rt.>>mcd

undist. 0

(Hqmi X (Hapa.)

strrp.

strep.

hr.

hr. unfus. sh.hr. pow sh.

sh.hr. unfus. mod. lrrg atr.

ctr.

St,

\tr.

0 (Hnpa. I

0

0

II

arc. Ing meldrd tall

0

rnrldcd tall

melded tall

d&y.

ana.

an-.

X

X

X

Ku1 ICInhhrrr IatiorlJ: Lcrnur’s = Lcmurids; Lpi. = I.epilemur; i)mb. = Ikmbentoma: Cheiro’s = Cheirogaleids. I. = l~tan~~.n; X = prracnt; 0 = not present: pass. = possihlv; - = !l,lt applicahlr; fus. = fused: unfus. = unfuwd: shal. sm. = small: lg. = = hh.dlow: pock. = pockrt; lat. = lateral: med. = mrdial: c~rtt = central; pust = posterior: larqv: wst. = \.cstIgial; hr. = hruad; mod. = modcratc: mk. = markrd: dist. = distrndrd; undist. = undistrndcd: WC. 1 occ-asional: strep. = strepsirhincs: sh. = short; Ing = long: nar. = narrow’; str. = straight: arc. = arcuatc: o~rnp = comprcswd; ant. = antPrior: scp. = separate: rnd = round: ang. = an,gular: cs. = rxtra; taxa in hracktw = C’S,cptir,ns: Halnpa. = Hapdtmut: G.cms<. = Gala,qo uaurmudatuc; C’h:hpiro = Cheirqnleus.

26

J.

1. Hypothesis

Figure

peduncle,

poorly

disruption

developed

during

the upper

the base exposure features

unite

Saban

pharyngeal” “anterior

as

the

(better carotid”)

homology united

that

the trigonid which

known,

on the lower

hypoflexid

tends

A clade

Lemuriformes, forms

synapomorphies broadening and

bc

molars

artery

the

lcmuriform

and

Cartmill

(1975)

Lemur identified

by the

of the

by its earlier be noted,

“ascending

appellation

Such an anastomosing

In turn,

the lorisids

to

vessel

that this claimed and galagids

ring fused to the bullar

of tall protoconids

of

does not proceed

and we doubt

a tympanic

a

These

homologue

it should

and

in the adult; artery.

and cheirogaleids. in genus

that rings

anastomosis;

ophthalmic

identified,

vessel,

established.

and metaconids

arc edge,

and a deep

the protoconid. Lcmuridac,

by Lepilemur), Daubentoniidae that include compressed lowering of the upper

distension posteriorly anterior elongation.

galagids,

that includes

to isolate containing

by the

that

of a narrow

stapedial

as it does in the latter.

as securely

a paracristid

a pharyngeocarotid

may

this small

bases;

fenestrae

at least on Ml; lower

the possession

that the vessel

tubaire”,

anastomosis,

that

stage:

of an anterior

palatine

cristae,

ofthe

primarily

in any non-cheirogaleid

can be regarded

basin;

and more appropriately

of lorisiforms;

and the development

the living

admits

i.e. the lorisids,

by a suite of synapomorphies notch

thr

of a choriovitelline

anterior

at their

reduction/loss

is supplied

“rameau

has not been identified

the

by folding),

by the development

artery;

of prehypoconc

(1986) suggested

form a pharyngeocarotid

is completed

placenta

that are melded

lacerum,

artery

& Cartmill

(1963)

below)

the development

the lorisiforms,

and MacPhee

(which

is characterized

(see

truncating

of the foramen meningeal

primate.

nasal fossa, and, in placcntation,

amniogenesis

facet of the calcancus;

middle

the major groups of extant strepsirhine

of an epitheliochorial

pharyngeal

cusps,

trochlear

I. TATTERSAI.1

fovea, a capacious

during

and metaconids

of these

elongate

among

a subgroup

incisors;

protoconids

AND

free yolk sac.

clade,

or ascending

separate molar

of a large

this wider

carotid

retinal

layer

the development

and the presence Within

SCHWARTZ

of relationships

of Rauber’s

presence

H.

Megaladapidae and Lc,muridae,

(as represented is united

among

by a set ofdental

upper molar cusps bearing distinct crests, a molar protocone in the sequence Ml-“, some

of the metaconid in the lower molars, and anterior premolars with Generally, the presence of a large stapedial branch and a small

promotory artery ha‘s been regarded as a primitive is possible, however, especially in view of the

character presence

retained by these primates; it in Plesiadapis of the re\rerse

INTEGRITY

condition,

that it is the latter

character exists

would

about

character

represent

Within

condylc,

distended

para-

distension

of the

indriids

a rounded,

expanded

de-emphasis and

is, of course, globular

gonial

with broadened

a lower

and

condyles

posterior

Two primary

points

anthropoids

of view are current

of the lemurs

in the suborder

however,

to consider

alternative

sister of Strepsirhini first of these of supposed

shared

robust

alternative While more

possibilities

theory (Figure

Prosimii

recent

substantial

Figure

of relatedness

is deeply

excavated

shares a deep

plane,

of the tarsier. Prosimii,

Ml-2

with

the

mandible quadratr regions.

teeth

can, however,

this enigmatic of the

1986). It may be worthwhile, Tursius is the

two: either

one of these groups.

set of assumptions

anterior

Either

or it is the sister

are essentially

and the ensemble

of enlarged

on Ml-“. posterior

and mesostylar

by Aiello,

which

a complex

trigons

the

of the

of Tarsius

or it nests within

(see below),

possession

uniting

To sustain

with the plesiadapiforms (see review

be constructed

the

Tarsius with a host on the

by Schwartz,

by considering

1984). X the other

2). as conventionally

arguments considerations.

region-primarily

prefcrrd

requires

plus

aspect

this form

in the coronal

in the suborder

possibilities,

that

but

and metaconule

(see review

+ Anthropoidea,

fossil relatives

basis ofthe more

Haplorhini

this entire

marked

splanchnocranium,

on the affinities

+ lorises

broad

Mr with

premolar

rounded para-

to leave Daubentonia

with

on Ml-“, elongate

with deepened

and enlarged

in this

uncertainty

on the posterior

autapomorphic,

The relationships

form is the sister

we prefer

is united

articulation

on Ml-2,

highly

cranium

region

hypocone,

(Megaladapidae)

regions

condition

suficient

of relationship.

of the postprotocristae

metastylar

However,

at present

hypotheses

of a secondary

metaconid,

Daubentonia

that

constructing

Lepilen~~

and

distally. with

when

by the development

mandibular

In that case, the lemuriform for the group.

of this morphocline

aside

Lemuriformes,

indriids

is derived.

a synapomorphy

the polarity

complex

which

27

OF STREPSIRHINI

constituted

for the existence Initial

the aboral

‘2. Two possible hypothrsrs scheme is shown left.

features

rounding

was essentially

of a haplorhinc brought

to bear focussed

of the narcs

of the relationships

based

between

on notions

ofgrade.

clade have been founded (Pocock,

To&r

on aspects

1918)-to

and other

which

primate

on more

ofthe

narial

were

groups.

Our

later

28

J. H. SCHWARTZ

added

features

(Hill,

1955).

aspects

such as fusion Most

capsule

and

platyrrhines Further,

and placentation

a fused upper

can occur,

namely

lip, he points

the coalescence

regard

1986), have

to the configuration

in Tarsius as well as in some

reminiscent

(1955)

of the orbit,

& Cartmill,

With

out that

creases Hill’s

rhinarium

and attendant

1974)) as well as fcaturcs

pointed

confirms

naked

placentation

1986; MacPhee

ofHaplorhini.

developed

(1980)

in possessing

this condition

Aiello,

has recently

variably

Hofer

(Luckett,

(c.g.,

of a moist,

of hemochorial

for the reality

(1980)

one finds

anthropoids

region

support

Hofer

while

lip and the absence

the development

auditory

been cited as further of the nostrils,

of the upper

recently,

of amniogenesis

nasal

AND I. TATTERSALL

of a strepsirhinc

conclusion out that there

condition.

Tarsius resembles

that

arc two ways in which

very early in development

of the maxillary

and median nasal processes, and the developmentally later gradual obliteration of the median rhinarial fissure. The possibility that syncheilism has been achieved by different means

in different

Hershkovitz various

primate

(1977)

platyrrhines.

as equids

and humans,

fused

of the nasal

tarsiers

the nasal

the absence septum tarsier’s

upper

these

a place

of the greatly

of the olfactory

recess.

naked

lips of these hairs;

mammals

in others,

known

bear

c.g. apes

in this particular

to state that on the cvidencc

from Anthropoidea”. cited

(1975)

as a synapomorphy

has noted,

in Tmsius this region

orbital

Similarly,

region,

which

the presence

01

reduction

of the

is diminished

in size

factor

also accounts

of an “apical”

of the 1972; C avc, 1973; Aiello, 1986) as indicative Clearly, however, the cxtremc extent of the septum in

has been cited (Cartmill, affinity with anthropoids.

Tarsius is due to the great expansion

on the development

of the orbits

He has argued

that

and morphology

this area

conditions

shown

expansion

of the alisphenoid

hypertrophied

(Starck,

1975); indeed,

enormous

eyeball

respects,

including,

(superiorly)

the

Cartmill’s wait

Cave

(1973) points

most

contributory

strikingly,

the extreme

reduction

(not scaled

still

point

in the tarsicr.

with

the

shallowly

the

structure

of the maxillary,

towards

structure

as a whole

synapomorphy

while expansion

at least, Cartmill,

would appear to lack this expansion: 1978, p. 102), WC do not feel that

Even

in most

frontal

and

orbital

primates,

p. 246).

However,

for size) of the area of the Tarsius medial

essentially Thus,

cups

the to the

accepts

other

the tarsier

between

if one

potential

uniting

the latter.

autapomorphic

1981,

much

primates.

he has pointed

as it is, merely

expansion

(Cartmill.

intermediate

Tarsiu., and

in fact a highly

components

for comparison

elements

in both

wall in various

Principally,

as expandt,d

1984)-is

orbital

a conformation

anthropoids.

contribution

(Castenholz,

arbitary

appropriate

and

of Tarsius-which,

zygomatic

of the medial

in Tarsius shows

by strepsirhincs

orbit

for

interorbital

to deficiency in the septum as a conscquencc of orbital size. In a series of valuable contributions, Cartmill (1975, 1978, 1981) has provided detail

in

and also lack a moist,

enough

but as Starck whereas

of

hairs,

such

are sinus

in size has been

expanded

sinus

mammals

is insufflcicntly

remote

by the findings with

out that other

prominences

the fused

reduction

Tarsius and anthropoids (Cave, 1973), nasal capsule in anthropoids is absolute, by the encroachment

nasal

unfortunately,

“occupy

capsule,

furrows,

has pointed

platyrrhines),

Tanks,

is suggested

median

(1980, p. 247) was confident

Hofer

region

As regards

median

anthropoids,

(e.g. various

they are not.

but overall

this condition

of vestigial

( 1977), moreover,

have

cases

displaying (1977)

As in Tarsius and

In some

respect,

Hofer

Szalay

and bovids

rhinarium. hairs.

groups

and

the rrlativc. proportions the autapomorphic nature ofthc

with anthropoids

alisphrnoid (although

see illustrations this fcaturc can

of the of this

must remain

a

some of the latter,

of A/ouatta and &bus in bc employed with much

INTEGRITY

confidence (1984)

in phylogenctic

implicit

that

effecti\+ unique. An aspect of the anatomy of a macula howe\.rr,

and

Massopust,

has recently morphology.

(1930) that

1970).

by Woollard

concluded

the retina On

(1926).

On

recent

found

the tars& presence

of further

by Woollard

rc,semblcd (e.g.,

Starck’s

of Tursius is

the supposed

the basis

reports

with

area

used to associate

including

csscntiall>.

of more

concur

interorbital

been

that the “macula”

of the tarsicr

the basis

we must

of the

1986) is retinal

as reported

Kollmer

of fixation,

Esscntiall!,.

development

of the eye that

(Aiello,

lutea

the

70 .__

STREI’SIRHINI

rrconstruction.

conclusion

with the anthropoids

OF

materials.

was an artifact

that of Lo& Castenholz,

(\?t’olin &

1965,

3984:

Rohen, 1966; Wolin & Massopust. 1970) it is arguable whether or not a fovea exists (or if it does, if it occurs consistently) in the pure rod retina of the tarsicr. Following a rev&v of the available visual

evidence,

including

morphology

MacPhcc

& Cartmill

structures tarsirr

and

annular

bridge,

accessor) include

cavity in this

maxillary

of‘thc

tympanic

cavity

of this region

suggest

that

additional

the

of chcirogaleids, altogether.

uniquely

auditory

region

is far from

Table 2

Distribution

arc:

ofthc

Ira\ing

tnqjor

& Cartmill an of

artrrb

is also characteristic

artery potential

In

only three

a small

may constitute

the promontory anthropoids:

dc,\.elopmcnt

is small

or lackill,<

s)~rlapOm~Jr~~hi~S

of‘ a cc~mples

and a postc~romedial

point of penetration

of the

ph\~lo~cnctic

si,qnal of characters

of the

s~1111.

the

clrar.

of various

morphologies

to

in f&t to be a combination

morc>o\ cr. thrrc

and

an

1,) the,

and

MacPhce

show,

cavity,

primarily

arteries

stapedial

of

of an anterior One might also

bridge

in which

data

artcrv.

promontor\ this appears

the

absence (as opposed

artery atmular

an

,itld anthropoids.

Hcnvcver,

lorisiforms

by the carotid

of‘

of‘ the

between

prrbullar

and galagids,

by tarsicrs

and unrcduced

a large mastoid

region

the

These

mitldl~~ mrningcal

of lorisids

and lorisids,

& cartmill’s

auditory

(‘aLit\.

the absc,ncc,

since sc‘\‘ere reduction

hct\\ccn

recess,

of four.

of‘

indicators.

consideration

synapomorphics

carotid

of the

shared

stapcdial

galagids

MacPhcc

rpit),mpanic

supply

ofthc. latter.

characters,

confident

tympanic

thcsc-. however,

uniquely

th(, vestigial

a d(>tailcd

artt.l’y, nnd the development c-ar. as a di\crticulunl of the auditor), tube.

middle

synapomorphy

two scparatc

felt most

arc also characteristic

features

prtGd(.d

that attributes

\-aluc as taxonomic

of‘rlo

Of SCVCII pc~tcntial

of the internal

Among

arc

recently

base.

of a small

category

artcry.

have

they

possession path

of the tarsier

cranial

the anthropoids,

transpromontorial)

shart,d

(1986)

of the primate

c1984) has concluded

his own, Castcnholz

and ph>Gology

among extant strepsirhines,

Tarsius and anthropoids

30

J. H. SCHWARTZ

AND I. TATTERSALL

Table 2 rontinued

Position

carotid

f.

Path of internal carotid artery Promontory

Lorisiforms

post-lat.

post-med. (cheiro’s)

transprom

artery

Stapedial

I~emuriforms

artery

transprom

sm.

SITl.4

lg.

\‘est.0

Ascending pharyngeal artery (pharyngeocarotid anastomosis) Middle meningeal primary supply Anterior cavity

0

X

Tmius

cent.

perhull.

Callitrichids

C&ids

Catarrhinrs

post-med

post-med

post-mrd.

perbull.

perbull.

pcrbull

lg.

lg.

lg.

lg.

vest.0

vest.0

vest.0

vest.0

0

0

0

max.

max.

max.

HOVIO (I’)

a.: stap.

ophthal.

max.

accessory

Foramen lacerum exposed

0

X

X

X

0

X

0

0

X

X

0

(Homo, f’m~o) Tympanic

cavity

lg.

sm. (cheiro’s)

sm.

Annular

bridge

X

0 (cheiro’s)

0

Mastoid

cavity

sm.

lg. (cheiro’s)

‘g.

Epitympanic complex

Stl,.

S,Tl.

sm.

0

0

k.

lg.

recess: 0

X

0

X

X

X

(cheiro’s) Orbital

frontation

hr.

mod.-mk.

mk.

Rostra1

elongation

0

mod.-mk.

mod.

0

0

unred.

unrrd.

crimp. red.

ahs. red.

abs. red.

ahs. red

sub= Sep.

sub= sep.

lat.>>med fus.

sub= SCp

sub= Sep.

sub= Sep.

shal./ flat

shal./ flat

deep pocket

sha1.l Hat

shal./ flat

shal./ flat

dist.

dist.

undist.

undist.

Nasal fossa Pterygoid medial Glenoid

plates plates fossa

Premaxillary margin Ethmoid

exposure

0 (indriids,

mod.-mk

mod.

mod. 0

X

X

X

X

X

0

X

X

X

X

Ha@.. I&, ) Palatine-frontal contact Yates Internarium Internarial

hair

Median nasal prominences

0 (Imur) strep.

strep.

strep.

strep.

hap.

br.

hr.

ex.br.

hr.

hr.

nar.

0

0

X

sinus

sinus

x (?)

rus.

fus./ mdngr.

fus./ mdngr.

unfus./ mdnfiss.

unfus./ mdn.fiss.-sulc

hap.

fus.

INTEGRITY

OF STREPSIRHINI

Lorisiforms

Tarriut

3 I

Table 2 continurd

Ixmuriforms

X

X

lost

to>t

fold.

fold.

Callitrirhids

C:ehids

C:ararrhinca

X

tc,\r cav.+l;Itrl.

paraemhryo. orthomrs.

paraembryo. orthomes.

pararmtx\ mesomcl.

difl’,epi.chorio. le;.yik.sc.sm Al.

difT.epi.chorio.

hrm.sm.\lk.

0

embryo. orthomrs. hem.sm.ytk sc.mrs.hdv

sh.hr.

tng.nar.

p0I-

welt-de\,.

41.

sh.

sh.

shhr. unfus. “rod.

t”R

I”g

“wd.

&r to nbhrerkztion~: cheiro’s = CJheiroqaleids; f. = f&-amen: X = prrscnt; 0 = “ot present; ~- = not applicable: l’us. = fused; unfus. = unfused; post = posterior; ant. = antc’rwr: tat. = lateral. med. = medial: crnr. = wnrral; rransprom. = tra”spromontorial; perhull; = pcrbullar; sm; = small: Is. = large: vest. = vestigial: sup. = stapediat: ophthat. = ophthalmic: max. = maxillary; hr. = broad; mod. = moderate; mk. = marked: unrrd. = unwducrd; comp.rrd. = reduced via compression; abs.rrd. = absolutely rrducrd; shal. = shallow: disr. = distrndcd; undisr. = undistended; OK. = occasionally: strep. = atrrpsirhine; hap. = haplorhine: mdn. = mvdlan: tiss. = lissurr; sutc = sulcus; gr. = groove; implant. = implantation; pararmhryo. = paraembryonic; embryo. = cmbry~x~ic: orrhomes. = orrhomcsometriat; mesomet. = mesomrtrial: diKepi. = diffuse cpithchochorial: chorio. = choriovitelline placenta; ylk.sc. = yolk sac; all. = allantoic vcsictc; hem. = discoidal hemorhorial; mes.bdy = mcsodtvmal hody stalk: sh. = short; 1°K = long; “ar. = “arrow: wrll-dev. = well-dewloped; str. = straight: arc = = arcuate; camp. = compressed: ant. = anterior: scp. = separarr; rnd = round: a”%. = angular; cx. = extra’ uxa in hrackcrs = exceptions; Hapa. = Hnpalemur; L.epl. = Lepilemur: Ikzuh. = Dnubentunia: .\f.mur. = .\licrorebur murinrtr: (;.rh~. = Goinqo dtmidoC: (~.~a.~~.= fkhqo rmtrwwdcztur: CW~~II’LI = rww~pitht~rids.

32

J. H. SCHWARTZ

I,uckctt

(e.g.,

1974, 1976) has argued

AND I. TATTERSAI.1

rcpcatcdly

that fixture

of‘amnion

fitrntatiott

placentation convincingly unite Tarsius and Anthropoidca. Among thr fcatutw thr de\.clopment of a mesodermal hod>, stalk. a rudimctttary allantoic waiclv. yolk sac, the prcsencc stage,

of a primordial

and the development

anthropoids,

as in tltr implantation

endometrium,

and

the shift

attributed

to the tarsicr’s

there

other

arc

hemochorial “anthropoid” (Luckett,

1974). Moreo\~cr

bypassing di\wticulum.

and

the, precocious

characters.

I+

[as

the

in

sittcc

anthropoid

as

and ccrtainl)

not as a s),napomorph!.

its course

dissimilar the

autapomorphic this process

condition among

primates

t)y Folding.

high birth weight. hcmochorial

placcntation.

.4 f’caturc that unite ofa

mcldcd

alrcad~ (Schb\xrtz.

is distcndcd and a marked

scannitig

clrcTr.on

Dagosto,

1987) rcl.ral

galagids. 1984). :\]I.

of‘ tltr. slrcpsirhinc specifically, galagids-hints distit1cCx.c

and

In

it cott\ygc*s Toniu.c is

but cotnpl&ttg

( I 98(i) citc’d it rcbti\~c~l!~

this simply

to hv :I c.orrc.latc. of‘

and TarJiut is the prcscttcv share

an angular

h\ pocuttid

on

margin that is rlon,qatc,d attd rc.lati\.t’ ~ttlar,qcmcttt. Further,

lo\vc,r tvcch of‘ Tortiud jwni/u~ (Rlusscr matches

anatomica

c~lottgation

pcJsscssic)n

on 31’ 2

a RI, 1~rot”‘““id-rnclacr~tiid

al\YYJlar

in this spccics

anoth(*r

na\,icular

cla~v. \vhosc

that

is totall)

:Cllo

lorisids

at an afTinit)- of Tur\ilo with grooming

by ca\.itatiott hypothesis,

1T1Ot’(‘O\TI‘,

of the* ;intcGr

toorhctrmh.

mammals. Alorco\w.

~a11 f’acing on the talonid,

that “I, ” morpholog)-

the calcancal

that

of‘(~ti(‘) ‘I;miuJ ~_yric,/zln shot\4

I~uccall~, LI prcmasillar~ clc,qrw of‘orbital li.ontation

micrographs

In sum. it apprlars

ttcccwit)

;ttnniugc~ttcGs

“itt c’cmcc’rt

ws LI homologous

in a manttcr

IIOMvvrr. cotlsidvrc4

at the hasc and fi)rmitt,q a stwp

hl, 2 that dct\vttward,

at-c all

aggrcgatc

discussctl.

chvirogalrida,

crista

pr~h)~~~Jcm~3

himwlf.

at1

of this ca\it)

iikc, a \ atic.0. of-other

ot‘ tLnctiotta1

(in 3 sartiplc

allantoic

sralk.

t hart

hc rrgardcd

Jixtnatiott

(If the tarsicr/atttltro],c,id

ltiat

Lwtrncggcr

out

in hcgittttittg

In support

[ 1973) finditty

Ixutcxneggcr‘s

and that.

purely

hod!-

rather

of‘hap1orhittc.s.

attd amniort

fi-om that of‘anthropoids. anthropoid

cumplcs

]&~cc?ita

01‘ an

[ 1!)73, p. 171)) argues. “amrtioRc,ttc,sis [emphasis added ] tht,

I,uckrtt

‘Tonius hqitts

kvith

out that, in tltr dv\~clopmc~ttr

primates] wtr\ci!alr~ cavit).“, the dc\x~lopmcnt

characteristic,

rodents)

of ;t chorioa1latitoic

as in 7’ct~vius”. cannot

ot‘ plawntal

\xious

IJ~YW

Howc\x.r,

which nr\w4iclcss drvclop antniogcncsis and placcntation

of‘ ‘t mcwdcrmal

amniotic by folding,

ha1.c.

uwrus.

dcvclopmc,ttt

f’unctiottal

devrlopmcnt of a prirnordial with definitive amniogcttc~sis

on

but

rudimwtar)

diffcrcnbtiott

Further.

sew

and

cdctttatcs,

cs~abhshmc~tit

‘l’hr!, thus form a single

of‘ittdcprndvnt ca\.itatioti

stagy.

pole, oi‘thc u(c~ittc.

to fillding,

than simplex.

(1975, p. 179) has poitttrd prCC(JCk)US

the

choriovitcllinc

correlated.

rather

bicornuatr uteri havt of hlastocyst implantation,

I,uckctt

placc!tita,

the

dasypodids,

ca\itatiott

placc,rrtal

c~f Tmiu.~ from

DifYcrcwws

to thv mw~mc~trial front

of‘s hicornuatr.,

(e.g.,

lack of‘s citorio\~itcllin~~ placc~nta.

Mastoc)-st

atnniogc,ncsis

possession

mammals

cavity,

hcmochorial

ofthe

during

placcntar that characteristics

of a hctnochorial

perfbctly

amniotic

of.a discoidal

anti

1~. 4tc.a arc’ ;I small ftx~,

the unite

that

rcy$tt, exists

lorisoid all li\.ing

that

ofthc.

;tnklc

mOrphOl(Jg)-

in chcG+&ids

group.

B

latc.ral tooth

Sintilarl\~.

str~psirhirt~s--and

and most the

highI>, perhaps

also adapids too (\wn IGtcrtin,qs~~ald, 1979). is also fbuttd in Tmiu,. ‘I’arsicrs actualI\ possess 1wo qroomitig claw3. on the- second and third pcclal di,qits: thtl 1attc.r. fi.aturc, is plausihl) In addition,

itttcrprctcd

as an autapornorph!-

‘distal tihio-fihular

lorisifbrm primates. hlolrcular (1986) cites the immuttodif~usiori

fitsion

adcld

to the,

~),tta1)o”“Jt‘l~tt)

with strqxirhirtcs.

c.xists in 7imiu\ as WCII as itt tttc’ ~on~paral)l\-si~cd

data ma) point in :I similar diwction: although ;2icllo studios ofI>cnc rt rti. ( 1!)70) as d~~rnonstratittg thv uttiC) of‘

INTEGRITY

?iirsiu.~ and .-\nthropoidca. studies

ofhemoglobin

of which

the same

scquenc~s

chromosomal Of ChrOlnOSOlna~

c‘rtr(.,ic-trrtdn/ll\. Hopefully si,qriific,ancc

. hroadvr

of this striking

grOup

(Baba v/ CT/.,1!@21later concluded

is enc.

hut

study

b)

h(JmOhgy

comparati\x~

comparisc)n.

.3

STREI’SlRI1ISI

that their data \icltl

Tnrriu.r + Anthropoidca

rccrnt prrlimin,iry aStolliShin,~ anlOUnt

OF

Tiniut

thrc~~ phylogc‘nics + Strrpsirhini

1’001~111al1 h’l\\~‘(‘ll

PI

N/.

ofcqual is another.

( 1985)

?i!~.\i~\ and.

an,ll\ SC.5\\,ill rlucidatc-

from

their

likclihnod. l:inall\.

;I

drmonstratcs 211 cspcciall~Ck/n,g~~ the ph) Ioq:-cnc‘tic

34

J.

this taxon couple

Leptadapis,

features

prcmolars,

quadratc check

lower

lower

premolar

Ml-z, teeth

Other

of

the

group

is certainly

three,

Smilodectes would

its allies; shared

holarctic

linked

into

hypocones;

a mctastylid

and

and lower last

ringing

ofuppcl

and

occlusall)

on the last

dental

the cheek

locvcr

with

notch region

Pe&odus

together

allies. If, then,

there

forms

spatulatc

or Notharctus (stout

seemed

cladc

regions,

arcuatc molar

at present

Adapis and WC delineated incisors,

quadratc

molar

However,

as opposed

various to crests

in

against

a

on M,, argue cogent

molars,

of lower

arguments

h/I, cristids

can bc

ohliquae

mcsostyles,

which

development

and

expansion

of

that ~Smilodectes, h’otharctus and

than any one of them does with ,-ldapis or its

In the course

1985), upper

on cusps

and a Pe!ycodus cladc,

to distinguish

expanded

stylar

clade

of Primates?

& Tattersall,

and mesially

Table 3

conclude

on M,.

more

the

to izdapis and

form of the upper

on the lower last premolar;

of upper

WC therefore

setting

have

and

folds on Ml--:‘. Of the

to he a paraconid

In any cvcnt,

(e.g.,

generally

1979),

the accentuation

the emphasis

on M1_2, development

exist an ildapis

and

allies

to bc rclatcd

the quadratc paracristid

cresting

Gingcrich,

of protoconr

cusps,

proper.

form a more plausible

that

rmphasizrd

buccal

of what appears

the adapids

on Ml-‘).

the wider (Schwartz

include

its

ATotharctus are

and

to have the bc.st claim

Smilodectes with Pe&codus (both

for pairing

meet the metaconid) hypocone

possession

anteriorly-directed

and the presence

close relationship

of entoconid

appear molar

Smilodectes

and

and Smilodectes and its allies

PeJvcoduJ (e.g.,

from

of Smilodectes, including

features

teeth,

A’otharctus

include

1986).

synapomorphies

the somewhat

including

Eocene

1984,

otherwise

apparent

crests

and

what

features

its close relatives

of European the Pe(wodus

from

a set of characters

that

check teeth with compressed quadricuspid

of each

arc the affinities

of describing

hII-

included

buccal

M.ith rather

cusps

broadI>,

Distribution of various morphologies among extant strepsirhines and “adapids”

Lcmur’s

Lepi.

Indriids

Loris’a

Gruorning claw: 2nd pedal digit

X

X

X

X

Toothcomb

X

X

X

X

Ertotympanic: aphaneric phanrric

Position

upper

(“prcmolariform”)

(e.g., Cantius, Protoadapis),

by the common

of the upper

and

well developed

and to a

of Adapis

specimens

submolariform

mctaconid

independently

the compression

Eocene

of the

Schwartz,

to be descended

group

include

and Paradapis,

complete

a non-caniniform

Pelycodus and its allies

Washakius-see

within

Adapis, Leptadapis,

Ml-:’ bearing

distension

“adapids”

believed

made

TATTERSALL

on the more

ofAdapidae

with

I.

and MI_‘<.

Cercamonius),

other

Based

by cingula;

canine;

AND

fossil genera

forms.

characteristic

and

(e.g.,

SCHWARTZ

to the well documented

of less well known

rotated

H.

carotid

1‘

.-Idapis

,I”““.

P&r.

3’0th.

.Smilo.

1tnk.

unk.

nttk.

0

0

I)

INTEGRITY

OF

:35

STREPSIRHINI

Table 3 mfirrud

Lemur’s Promontory Stapedial

artery arrer)

Ascending- pharyngcal artcv ipharyngeocarotid anastomisisl

Lep1.

Indriids

Loris‘s

ildapis

Pt$C.

N&h,

sm.

sm.

sm.

sn,.

sm.

sm.

sm.

1s.

vest.

1%

vest: 0

le;.

I$.

‘4.

0

0

0

X

(I

0

0

Orbital

lionration

hr.

hr.

h-.

mod-mk.

hr.

hr.

hr.

RostrA

vlon,~atwn

0

0

0

mod.-mk.

0

0

0

P remsxillar\ margin Ethmoltt

undist.

cxposurr

0

undist X

undist.

diyt.

undist

“CC.

X

X

0

0

0

.Smilo

undist >

(Hqmr Palatine-frontal Cr)nt,l< t

X

3

L

(Hapa.)

\h.

sh.

sh. (lorisids.

Na\-icular

sh.

sh.

sh.

Ing (;.cmsc.) ‘nq

(lorisids. ‘l‘r
sh.hr.

Paracclnid

(;.crarx.

sh.

sh.

sh.

sh.

41.

sh.

I

sh.hr.

sh.br.

nar.lng

sh.hr.

sh.br.

“lip”

arc.

str.

arc.

“lip”

0

0

0

M,

MI_.,

II,\\

Paracristid

a,,t.

“ant.“

hl, trigrlnid

open

“open”

es.comp.

open

hc.

but.

mrd.

rs.med.

mrd.

mtd

med

bUC.

hut.

med.

med.

med.

med.

hut.

hut.

but.

mcd

“kinks”

“kinks”

spat. qu”‘I. quad.

red.

red.

Id.

transv.

transv.

quad.

transv.

tmnsv.

qu‘ltl.

hlh.

sctniwmp

Cristitl 11,

ant.

sh.hr.

obl.-med.

ohI.-med.

cump.

camp

~Miqua

M,

red.

(I

spat.

‘:!:::;’ (chviro’s] rx.mcd. ichviro’sj hi.cr.

Xl’ 2 \h‘qx

wet.

quad.

quad.

transv.

11 ssh
rect.

quad.

quad.

transv.

semicamp.

camp.

camp.

hulh.

I!pptr molar protocristac

br.

hr.

ex.br.

“1’.l-1”

hr.

Protoccme

br.

br.

hr.

nar.

br.

Upper

obl.-med.

incisors

Upper molar hut cal cusps

.M-”


“V”

mdl-mtd

“V”

cs.hr

nar.

hr.

hk tu abbreumtions: L emur’s = Lemurids; Lepi. = Lepilemur; Loris’s = Lorisids: Adupis = Adapis-group; Pelvc. = I’e&odus-group; Noth. = Nothnrctus-group; Smdo. = Smilodectes-group; cheiro’s = Cheirogaleids; f. = foramen; x = prpscnt: 0 = not present;= not applicable; fus. = fused; unfus. = unfused; post = posterior; lat. = lateral; med. = m&al; cent. = central; sm. = small; lg. = large; vest. = \,cstigial: hr. = broad: mod. = moderatr: mk. = marked; dist. = distended; undist. = undistended. occ. = occasional; sh. = short; lng = long; nar. = narrow; str. = straight; arc. = arcuate; ant. = anterior: &I. = oblique; mrd. = medial; but. = buccal; camp. = compressed; mtd = metaconid; red. = reduced; spat. = spatulate; hi.cr. = high crowned; rtct. = rectangular; quad. = quadratc; transv. = transverse; bulb. = bulbous; “\“’ = “\“’ shaped; “I”’ = U shaped; ex. = extra: taxa in brackets = c~xceptions: Hapn. = Hapnlemur; G.cm j. = f&zlqo rnmicaudafuz.

36

J. H. SCHWARTZ

divergent

protocristae

compression

of which

of the first three

on the last

lower directed

paraconid

(see Table

conclusion

that Ad+

paracristid

almost

bears

short-crowned ancestor. among

radical

no obvious

If

the

lower

has

relationships

procumbent

a dental

comb.

Only

time

and further

whether

or not the lower anterior

dental

conformation

of‘rldapis

of the wider Apart must

Eocene

from

clade

the indriid-like

conclude

evolved

that

would

adapids.

The

“lemur-like”

consisting

these

link it with

may lie within

of the cctotympanic retention

(the

development), another

while

lying

characters

bulla.

hypothesis we have

within

the tubular

of the uniquenesses

tentative

within

and Leptadupis

as a whole.

can, however,

that

If: as suggested

above,

the lorisiform

including

bulla,

extension

which

plausibly

of the cctotympanic autapomorphic

within

it may

primate.

this larger

group

of Tursius relationship

as a primitive

in

the

course

be regarded

Figure

that

clad<

on the basis

the spatial

“captures”

the serves

The larger

the relationships

group,

we

no derived

be defined

in Tursius can be interpreted

of this highly

must will trll

was typical

shows

Strepsirhini,

to Strcpsirhini

with

the

of relationships been

group

relationship

ring to the bulla ring

offluubentonia

fossil discoveries

the PeLycodus-group

and the strepsirhines

specifically

parallel

Hudropithecus

of Smilodectes, lVotharctus and Prorycticebus

bulla

of the auditory

Strepsirhini,

cheek-tooth

of Smilodectes, a feature

of the molars

any particular

auditory

and

belonged.

in the two taxa,

taxa in a sister

of the configuration

forms

quadratcness

Pe&odus-group

ofthe

these

in parallel

features

only to place

to which

by

indriid

and an ancestor

have

the

orthal

subsequent

subfossil

likewise

lurks

share

a toothcomb-hearing

bc without

of the

antecedents,

ofcoursr,

relatively

from

the

includes

forms

suggested

not, however, dentition

problem,

of a

towards

which

The extant

Adupis

by an

absence

point

strepsirhincs

Adupis is descended

would

anterior

sign ofits

possessed

the

that

modification

strepsirhines:

shows

teeth.

and

dcntition

i\ serious

while

crests

also distinguished

lower dentitions.

comb,

buccolingual

and trigonid

lingually,

of living

to the latter.

anterior

it follows

molars

trigonid

to a group

dental

incisor

are accurate, Such

ofthe

lower

the

emphasized,

on the talonid

of the posterior

and particularly

strcpsirhine lower

opens

characters

an affinity

conformations

ubiquitous

morphology

that

3). These

is less

emphasis

as well as on MI_l,

Lepilemur and the indriids, in the respective

postprotocrista

lower premolars,

premolar

anteriorly

the

AND I. TATTERSAIL

3 provides

is consistent

of

as yet a very with

the

discussing.

Fossil “tarsioids” Numerous fossil primates, primarily with Tarsius in a “tarsioid” group justification material, American these

exists especially

for

this

association?

of the European

Tetonius (Gregory,

impressions

include

distinguished by their small size, have been associated (e.g., Szalay, 1976, and references therein). What Historically,

Necrolemur

1922), h ave appeared relatively

large

orbits,

features cctotympanics. In the postcranium, navicular, and in the case of the microchoerines support

this

relationship,

although

as early

overall

(Simons

to support

it: characters dental

as elongation

tibio-iibular

fusion,

1922 Gregory

from

1960) and

V-shaped

such as

impressions

& Russell,

arcades, of the

cranial

the North

contributing and

calcaneus

have also appeared pointed

out

to

tubular

that

and to the

similarities of Notharctus with Galago were more impressive than those with Tarsius, and Simpson (1940, p. 196) commented on the postcranials of Hemiacodon that “if these bones were judged entirely on their own merits to thr Tarsioidea”. But despite the general

probably nobody would refer them definitely dissimilarities from Tarsius in the dentition of

INTEGRITY

Figure

3. Tentative

microchoerines, some

way

The

tarsier

first question

molars

are considered

distinguish

them

from

preserved

small

teeth

Wushakius,

presumed

Chumashius

and

a cohesive

referred

Other

of the jaw;

by a common

to collectively

crista or its apparent

the

these

type

primitive

clade.

many

The

group

as omomyoids, derivative,

of these

with

enlarged

protocone

protoconids

and metaconids

melded

talonid where

basin.

This

postcranials

set of characters are known,

to the extent typical, for example, known astragali associated with morphology

otherwise

seen

likewise

the omomyoid

from

and forming

in lorisiforms,

latter which

could

as noted

is he

earlier,

wall facing bones

by on the

are elongate

The trochlear narrow and

not in Tursius. These

premolars

“group”

and does not

and Tursius. In cases

lorisiforms and navicular

Lower

had

of a prehypocone

a steep

calcaneus

if not the absence, large and distinct

known

are characterized

distinguishes

although

Omomgv.

best

to have

teeth,

teeth,

of most G&go and Microcebus. these taxa displays the derived

Tursius in the diminution, Tarsius retains the posterior lower molars. molars; lorisiforms lack paraconids altogether.

distinguished

This

molars

to

would

Loueina, Shoshonius,

fold. This feature,

clade as well as of Tursius. Lower bases

appear

anterior

anterior

that

tooth,

in Mt the presence

of the lorisiform

actually

allocated

Pseudoloris,

(e.g., anterior

of Chlororhysis.

is distinctive

at their

been

Anaptomorphus,

of diminutive

also shows

a full-fledged

forms

of a

molars.

“tarsioids”

have

in contrast,

include

similarities,

even one feature

an enlarged

specimen retention

the fossil

in

molar

the presence upper

the years

to detect

tarsioids,

forms

1961). These

three-cusped

over

in general

for example

whether

that

in possessing

at the front

distinguished constitute

lower jaws.

of simple

gmups.

the idea that all are relatrd

retentions,

is simply

However,

primate

via the similarities

we are unable

are distinctive

and extinct

form (e.g., Simons,

If all taxa

together,

as a clade.

Tetonius, Absarokius)

then,

living

all omomyids,

and presence

group.

the 1 arious

to persist

of primitive

to be asked,

a monophyletic

Tarsioidea

managed

in the form

on the lower

constitute

among

and almost

has

37

STREPSIRHINI

Omomys and the extant

between are mostly

paraconid

of relationships

anaptomorphids,

to the

morphology however,

hypothesis

OF

forms

facet of elongate are also

of paraconids on at least paraconids on all lower of lorisiforms

do not have

38

J.

truncated

heels

but are instead

Tursius and omomyoids. septum,

and some

ectotympanic

H. SCHWARTZ

orbital

elongation

is intrabullar;

synapomorphies

oval in cross section;

Relative

rostra1

and enlargement.

also distinguish

this larger

group

cristid

show

obliqua

and

that

meets in

the metaconid

a cristid

of Ml-2;

teeth,

M‘J broad

completely

lingually,

above,

with

derived

features cusps the

squared

the other

+ Strepsirhini. ofa tubular

Loueina, Anuptomorphus,

namely

encloses

with

interorbital

In Necrolemur the

in its development

on M t; MI_2 talonid

that

group.

argued

be autapomorphic

anterior

also characterizes reduced

of the Pe[_ycodus-group

to Pe&codus in various

similarities

incorporated

transverseness

small

as we have

that consists

As in the case of Tarsius, Necrolemur would extension of the ectotympanic. Some fossil “tarsioids” with

this configuration

frontation

this is consistent,

of the yet larger

Chlororhysis,

AND I. TATTERSALL

(e.g.,

poorly

talonid

corners),

and

and

an arcuate

distinguished basin;

extreme

belong

with

the

Pelycodus-group. The

of Washakius

affinities

Shoshonius appear

and

Smilodectes. Features

Pebcodus-group:

Ml_:! that meet the metaconid, molars,

mesostyles

distinct

protocone

are

common

and a mesially

present

a character

Summary The

relationships

firmly

within

established than

However,

with

the similarities

the primary retentions

factor in both

highly

severely

limit

clades

artery,

and in some by Aiello

for this relationship.

Those

In contradistinction there

non-anthropoids.

exists

in

In the widest

and the Pebcodus-group,

“lemur-like” the specialized

configuration ear region

other

and

context,

crista melded

which

a problem

evaluation

the diminution

have been primitive

apparent a suite

suggests

region,

derived

of a

potential carotid

and lack of a

with the anthropoids,

the living

Tursius, are united specifically, Tar&

list of

in support

suggesting

affinity

strepsirhines,

ancestor

and tympanic

provide

of the internal

synapomorphies

of Tarsius is also plausibly

fossa,

that few provide course

from a common

bulla

tarsier

of the

(1986)

of the auditory

that

of the

of its phylogenetic

re-evaluation

of characters

it appears

due principally

of the nasal

Our

& Cartmill

features, cavity

are descended

of the auditory

associated.

represent

apomorphies

that do are the prebullar

this larger group all extant taxa, including claw on the second pedal digit. Yet more prehypocone metaconids

for

by MacPhee

tars&

clade

traditionally

structure)

The

primates.

accessory

1 arc reasonably to this is the linking

actually

problem,

primate.

available

to these the

possess at large.

to link the two.

more difftcult

and of other

of an anterior

it has been

the lemurids,

for example

(1986)

in Figure exception

and mandibular

of this

with

relationship,

development

however, adapids

cases,

“tarsioids”

with the Indriidae/Daubentoniidae

with

potentially

synapomorphies

cited

rhinarium.

a much

on

on M3. In the upper

toothed

as reflected

which

and thus do not serve

characters

Tursius-anthropoid evidence

in cranial

nature

of the

obliquae

of the Peivcodus-group

One possible

with

Lepilemur

represents

the

apparent

characters

(notably

autapomorphic

relationships, false

the family

in grouping

Tursius, of course, to the

group

Megaladapis

relative

obliqua

anterior

member

cristids

and conclusions

the strepsirhine

Lemuridae,

another

taxa include

cristid

distinctive

on the basis of morphology.

of Lepilemur and its subfossil rather

directed

on M 1. All small

folds on Ml-2,

to lie with

to all three

ring.

with

plus the

that possessed As discussed

from this condition.

the

above, Within

by the presence ofa grooming and the lorisiforms exhibit a

on Ml-2, ovoid lower premolars. lower molar protoconids and at their bases and forming a steep wall facing on the talonid, buccal

INTEGRITY

expansion

of an angular

the prcmaxilla, frontation.

hypoconid

some

Othrr

anterior

features

on MI_?, downward

rostra1 that

elongation,

appear

tarsicr-lorisiform

link include:

loss ofthe

of the calcaneus

and navicular

(except

degree

of chromosome

Characters

affinities.

the evolutionary

bc the obvious name) than one suggesting knowlrdge

the

phylogtanetic dcservrs

suggestive artery

orbital if not

in adults

cmrirmdatus

in G&go between

that point

apparent

inconveniently

autapomorphic

but

nesting

this

form

with

group,

of and

of a

elongation and a high

G. crassicaudutus).

and towards

cladc

7hrsiu.t

demonstrative

ofthis

s) napomorphies

of the non-anthropoid

margin

enlargement

and in Iorisids),

the tar&r

in the tarsier

The fact that hierarchy

be

of the alveolar

relative

both anthropoid exist at different

(for which

may well suggest that a hypothesis emphasizing a tarsier-anthropoid cladc. (:lcarl),. at the

placement;

serious

to

cxtcnsion

and

stapedial

(at least

may thus be discerned

non-anthropoid within

homology

39

OF STREPSIRHINI

continues

the lorisiforms

Prosimii

and levels would

these is more robust current state of our to elude dcfiniti1.c is a possibility

that

consideration.

Acknowledgements We

thank

Lawrence Fred Grine, manuscript. Anthropology,

the Martin

Richard

Lounsbery

for their

and several

Foundation

invitation

anonymous

This is Contribution American Museum

fill- its support,

to participate rcviewcrs

and

in this symposium.

kindI)- commented

no. 20 of the Lounsber) of Natural History.

Fred

on an earlier Laboratory

Grinc

Peter

and

Andrews. draft of the

of Biological

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