Primates: cladistic diagnosis and relationships

Primates: cladistic diagnosis and relationships

introduction The past twenty relationships Novacek,

years

have witnessed

within

Mammalia

1982, 1986). The reasons

general

increase

(1966),

of interest

the marked

(Lillcgravcn molecular

rt al..

1979;

tree,

hut

Savagr

widespread Glircs

agreement

(Luckctt

ordrr

this rebirth,

in mammalian

comprises,

current

state

and this presents

influential

publications

promoted

th(,

‘fhcse

view trends,

trends,

as seen

progressive

abbreviation

prop;rcssivc

expansion

thr

should

tx. united

in the Supcrordrr

has rc,maincd

a prohem

for morphologists

stage,

thcrc

a historical

holdover.

in the 19.59 hook Thu .-lntrc.&lls Primates

clear-cut,

modern

tree with

clahoration

is best idcntifiahlc

shrews reduction

ol‘ the brain.

defined

In a series q/.\lnn,

is

a lemur-like

stage,

a monks\-like

what the

unique

include,

‘I‘hc 01‘ \‘crc

Lr Gras (Ilark

in terms

shared

to man,

about

relationships.

of a suite

ot‘

specializations. for

instance..

the

of the apparatus of smell, and the* A definition of Primates hased on

progrcssi\rr changes in features lent support to the strp-wise‘ gradistic c.volution also promoted by Ix Gras Clark: that PINrou/r tu man, primates trccx shrew-like

onr.

‘l’hcre is no consensus

for anal) scs uf higher-levrl

mcasurc,

of the snout and

of the mammalian For

and rodrnts

prohlrnls

Order than

from

pattern

of numerous svstematists

reached.

relationships.

culminating rather

fossil rccurda

the appearance 1982). Mammalian

have* hecn

1977; arc thy

the bvork of Hcnnig

and Cenozoic

of the branching

Primates

is. in large

that

iGoodman.

Szalay,

and Lraricd. Included

1985).

the Order suprrordinal

of affairs

evolutionary

la~gomorphs

in the superordinal 1975;

following

lYN3), and

phylogeny of corigrucncc

& Hartenhcrgcr,

‘l‘hroughout intrrcstrd

that

arc many

of the hlesozoic

on most aspects

points

ofintcrest

XlcKcnna,

systcmatics

& Russell,

tbr assessing some

rebirth 1971:

for this rrhirth

in our knowledge

arc, as yet, far ii-om agreement ordinal

Valcn.

in phylogenclic

increase

techniques

a rrrnarkahlc

(Van

stage.

view of primate went through a

etc. ‘The trends

and grades

01

2

J. R. WIBLE

Le Gros Clark’s

writings

many

continue

of whom

evolutionary Attempts

systematics.

had

primates

to define

several

arc either

as such,

cannot

shrews

from

order,

(synapomorphies)

for

of the primate

of the Paleogene Recently, definition

(1986)

boundary

hard-tissue (i)

fossil record

Martin

of the Order

primate

research

Primates.

Well-developed,

divcrgcnt

Elongated

distal

segment

(iii)

Relatively

large,

convergent

(iv) (v) (vi)

ethmoid

distance

relative

Petrosal

bulla.

Relatively Dental

of Hennigian

(tree

Valen

shrews)

and

acquisitions

With

and,

the removal

of tree

orbit and

this list ofsynapomorphies

of the order.

Nonetheless,

attempting

was required

this

to delineate

for analysis,

the

while the bulk

was teeth. his 1968 articlc

He specifically record.

His

that are applicable

present;

than

as Van

in the wall ofthc

offered

as a definition

material

has updated

(ii)

bar

Martin

for the paleontologist

Cranial

in the fossil

features

than

satisfactory order.

order.

of these

bone.

rather

five shared derived characters characters involve hard tissues

ofa plate of the ethmoid

from the petrosal

for future

by tupaiids or independent

to identify

two

of primatologists, grades

He concluded,

shared

in the same

on

the inclusion

was not very

boundaries

groups

Only

bulla formed

as a program

these

Primates.

an auditory

(1968).

(plesiomorphies)

went

in fossil forms:

“definition”

retentions

generation

of adaptive

1986; XC below). using the methodology

Primates

the features

Martin

identifiable more

that

be used to unite the

in terms

first was by Martin

earlier,

primitive

the current

order

to delincate

The

years

influenced the

et al., 1983; Gingerich.

clades (MacPhee have been made

phylogenetic (1965)

have greatly

AND H. H. COVERT

definition

to analyses

hallux

with

exposure

of modern

primates

phalanx

the six

arc (p. 20):

in the foot. index

interorbital

possible

detailed

includes

These

(viz. low calcaneal orbit

more

of delineating

of fossil remains.

with restricted in the

a much

the problem

a flat terminal

of the calcaneus orbits

to provide

tackles

value).

distance.

(depending

Postorbital

on interorbital

to skull size).

large

braincase.

formula

Sylvian

sulcus

on endocast.

maximally 2.1.3.3. ---_-_--. 2.1.3.3.

Premaxilla h4olars Martin record.

upper

incisors

arranged

cusps.

Lower

noted that this list is not without For example, a number of Eocene

grounds

retain

identifies many

short;

with low, rounded

four

premolars

as synapomorphies mammals

Paleogene certainly Paleogene

that

collections.

they

above

arc

Likewise,

useless a feature

below.

that

the

Late

The years

between about

the publication the delineation

of Martin’s

longitudinally. talonids.

the and

primates

such as a relatively

of the Order

that

arc so vague

for delineating

Cretaceous-Paleogene

than enlarged

when applied to the primate fossil that are clearly primates on other features

the

while

so-called

Controversy

in most

complete Martin archaic

because there is no evidence primate features.

two contributions

he of so

non-primates

braincase,

so few relatively notwithstanding,

plesiadapiforms,

Primates.

molar

characteristic

from

large

because problems

primates, should be excluded from the Order Primates, these forms had any of the above listed six hard-tissue lively debate

raised,

WC add

primates

true, is problematic for the paleontologist, mammalian crania are known. These

concluded

transversely with

problems mammals

and

of modern

more molars

have witnessed has concerned

that a very not

PRIMATES:

DIAGNOSIS

only discussions

of specific

orders.

Cartmill

( 1972, 1974) argued

shifts,

that

ancestral

taxa and features,

this

criterion

and descendant

criterion

of monophyly”

specific

habitat

from

(1972,

an ancestry

that

Cartmill

Plcsiadapiformes shared

postcranial

adaptation branches

involved

Some

Plesiadapidae,

(including

Anthropoidea).

They

concluded

with

ancestral

on insects

other

primatc

in the

terminal

to classification

(1983)

identified

like that

three grade

Picrodontidae,

Omomyidae,

the

because

12 1).

and Dermoptera),

Lorisiformes,

from

adaptation

a plesiadapiform

Paromomyidac,

differed included

the order

“the

approach

relationships:

Tupaiidae,

Lemuriformes,

p.

et al., 1975, p.

co-workers

that

et al.

MacPhec

phylogenetic

Mixodectidae,

(1972,

to a gradistic

Clark.

Carpolestidac,

Apatemyidae, Tarsiidae,

returned

not

to an arboreal

1972) within

predation

forests”

1974;

(Szalay

of an arboreal

between only to the

& Decker,

by these authors

proposed

visually-directed

by Lc Gros

by unresolved

and

adaptive

synapomorphies

adaptations

and

in Simons,

group

of tropical

have

for primates linked

grade

strata

primatologists

promoted grades

nocturnal,

Szalay

indicative

that

(Szalay

terrestrial”

shift proposed

& Tattersall

specializations

of the lower

habitually

1974).

subject

as primate basic

for defining

major

boundary

of caprice,

co-workers

but also “the

(1972,

excluded

a matter

and

reflect

taxonomic

identifying

and adaptive

(sensu Simons

Cartmill

Szalay

of the criteria

should

of any

largely

was probably

boundary

by

boundaries

approach,

specializations,

165). Yet, the primate

they

p. 98).

a similar

that

primates;

becomes

postcranial

postulated

but also discussions

that ordinal the “placement

groups

et al., 1975) followed

Szalay only

without

3

AND RELATIONSHIPS

primate

(including

Microsyopidae,

a prosimian

grade

and Adapidac),

(includirrg

and an anthropoid

that:

The search for exclusive features diagnostic for primates might be better replaced by attempts to formulate and account for the significant grade boundaries found within the order. Only two major grade transitions, delimiting the upper and lower boundaries of Prosimii, can be defined at present. ‘Phe antecedent plesiadapiform grade cannot be precisely defined, and it may even be polyphyletic. Polyphyletic origin of a grade damages its taxonomic status but enhances its rvolutionary significance, because a grade boundary crossed by several parallel lineages is correspondingly more likely to reflect adaptation under pervasive selection pressures rather than historical accident (p. 511 i.

More

recently,

is more

Gingerich

informative

what

WC know

than

a cladistic

than

(and

approach

argued

do not know)

of the literature

regarding

about

by MacPhee

approach

“A grade

the phylogeny

to delineating

classification

primates

appears

of early primates

to express

more

accurately

been

expressed

p. 40).

reveals,

the delineation

advocated

that a gradistic

on clades.

(1986,

classification”

As this survey recently

(1986)

one based

vastly

different

opinions

of the Order

Primates.

et al. (1983)

and Gingerich

have

At the extremes (1986)

are the gradistic

and the Hennigian

approach of Martin ( 1986). The former essentially argues that we are unable to delineate the primate order on the basis of a suite of shared derived characters and that it is unlikely that we will bc able to do so in the future. includes

not only Plesiadapiformes

Microsyopidae,

Apatemyidae,

These

(sensu Simons Mixodectidae,

authors

figure

a history

& Tattersall

Tupaiidae,

of Primates

in Simons,

and

that

1972), but also

Dermoptera.

In contrast,

Martin argues that we can and should offer a concise definition for modern primates and that fossil forms which fit (or nearly fit) this definition be considered primates. Martin explicitly following, cvidencc

excludes

the Plesiadapiformes

we review behind

the

the evidence various

upon

hypotheses

and Tupaiidae which

these

of phylogcnetic

from

the Order

hypotheses affinities

Primates.

are based, between

that modern

In the is, the and

J. R. WIBIX.

4

archaic

primates

(cladistic)

and related

approach

euthrrian

for reasons

ANI> H. H. (:OVER’I

groups.

enumerated

Methods According

to Hennig

(1966),

relationship

or phylogeny.

evolutionary

novelties

ancestor

and

Natural

groups

are

usually

characters

cannot

be used

to one

to recognize

synapomorphies

tree. are

alone,

termed

discussion see Wiley, 198 1) The entire organism, including should

portions

and extant

already

been the most intensively that WC concentrate. in particular, fossils.

Polarity

of

(apomorphous) the group

under

relationships ordinal

other

Cartmill,

among

cartmill

(1974,

the

phylogenetic

evolutionary

presented

the

C:artmill indicates

characters that

(see

Szalay

is, derived

the taxa affinity; trrnds

have

is the pool from which With

over the last years,

and it is here

ofthcse

anatomical many

assumes

1980: Novacek, have

a unique

relatives

of

outgroup

of the eutherian

in allying

to exhibit

derived

(Maddison

that

pattern

feel compelled

demonstrated

analysis

in the closest

method

regions,

of the relevant and

of‘outgroup

the branching

which

regard

and pes have

not studied

this

by

further

the basicranium

is that found

& Drawhorn,

characters

groups

littlr

to rely

homoplasy

1986; MacPhee or rrlatively

&

unique

studied. argues

that

they must

characteristic

example

been

(for

(plcsiomorphous)

howr\pcr,

12:e therefore

which

p. 442)

WC havr

and unfortunately obscure.

groups

such as this, it is only those

descriptions

state

branching

not recognized

that can be sampled.

the method

that the primitive

of shared of characters

as evolutionary

those

oflife,

orders,

primitive

using

As a drawback,

eutheriwns

1986),

distribution merely

assumes

are well resolved, derived

their

is determined

states

polyphyletic

of the anatomy

because

and arranged

for subsequent

and mode

on published

or cladograms.

originated

in a review

ruthcrian

portions

primates,

tree is still somewhat

on those among

studied

study.

or

well known

and related

characters,

statps,

et al., 1984), which

somewhat

they

of

a common

distribution

of organisms,

unavoidably

We rely heavily

for Paleogene

groups

paraphyletic

Yet,

primates

because

no information

its development

be sampled.

of the organism

to extinct

diagrams are recognized

hierarchical

sharing

through

(plesiomorphous)

groups

Unnatural

inherited

the

or clades)

by the

provide

is genealogical

through

of branching

groups Primitive

and

of organisms

recognized

characters)

another

natural

ancestor

in the phylogenrtic

are

in the form

(monophylrtic

(synapomorphies).

in an earlier

characters

for the classification

relationships

depicted

to follow the Hcnni-ian

and materials

or apomorphous

relationships

derived

points

(derived

oforganisms

in hierarchical

novelties

the basis Such

In this. we choose below.

of the

differing

“taxonomic also mark

boundaries important

of a radiating adaptive

higher

shifts

must

adaptive taxon.”

postulated

reflect shifts

more

Above,

for the

than

that underlie wc have

primates

by

(1972, 1974) on the one hand and Szalay el al. (1975) on the other. This example the subjectivity involved in choosin,? which features to analyze for the

reconstruction

of “adaptive

shifts”.

We consider

the delineation

of taxonomic

units and the

adaptations of those units as separate questions (Lauder, 1981). One cannot about the initial adaptations of a group without knowing which organisms

ask questions are included

and which forms has

include

between

arc excluded. The delinc;ltion of hi,qher long been problematic for systematists,

our knowledge

of modern

taxonomic in part,

and fossil organisms.

groups

that

fossil

because thrrr is disparity \Yithin Hennigian classification,

PRIMATES:

two major

approaches

facilitated

by introducing

community” members ancestor closely

1981; Ax,

& Rosen,

members

to a particular

community

1977) equate

will rvaluate

taxa

have

the results

group

a crown

the name

1986; Gauthicr

surfaced.

A stem than

group

The discussion

lineage”,

lineage

and its stem lineage.

of a living

of these

and “closed

includes

to its living sister

group

~1al., in press)

of our rrview

“stem

is

descent

1). ;2 crown . group constitutes the living to hc descended from the closest common

1985) (Figure

of the group.

.I

RELATIONSHIPS

‘&crown group”.

crown

includes

(Gauthier,

higher

the terms

AND

and all fossil taxa thought

of the living related

descent others

for naming

(Hennig, of a group

DIAGNOSIS

with

restrict

in light of both

t-descent

the fossil taxa more

group.

Finally,

Some authors

the closed the name

a closed

(Patterson

descent

community:

to the crown

group.

Ct’c

approaches.

community-(

Figure 1. Cladogram illustrating the terms “crown group”, Listem linra,ge”, and “closed descent community” (Hennig, 1981; Ax, 1985). The crown group B-C includes the modern taxa B and C: and their closest common ancestor. LX.The stem lineage ofthr crown group B-C includes the outlying fossil taxa mow closely related to B-C than to its modern sister taxon A. The closed descent communjty &B-C: includes the crown group and strm lineage.

MacPhee

et al. (1983)

of
clades

a rationale

plesiadapiform postulating “pervasive Although

for delimiting

(praesimian a grade

how a group

and Gingerich in delimiting

without

of organisms

“Definition” in systematics

(Ghiselin,

within

no delimiting

Primates),

equivocal 1984).

primates; cannot

throughout

our usage

at most,

followed

the desirability

do these they

What

Moreover,

features

term and should “In conventional

about

Nowhere

be defined.

for its limits?

was employed

claims

boundary.

grade

any rationale

of the Order is a highly

grades

for which

make various

primate

of Gingerich)

selection pressures”. the term “definition”

the definition

(1986) the

that

the

is the purpose

admit

of

we do not understand

arc known

can somehow

the introduction that

for a name

to apply”

(ibid., p. 105). We do not proceed

necessary

and sufficient

attributes

to define

taxa in systematics,

applies

that

to define

groups.

is, by observing

We recognize that

they

natural share

(for example.

of the authors

and sufficient

above,

reflect

discussed.

be replaced with the term “diagnosis“ logic, a definition states the attributes

nrcessary boundaries

authors

groups

derived

through

characters.

by assigning

that is, we do not set the the methods The

term

discussed “diagnosis”

to the “brief listing of those characters which differentiate a taxon from related do not define taxa in the and/or similar taxa” (Wiley, 1981, p. 377). D’la g nostic properties conventional logic sense, but are useful for identification of taxa.

b

J.

R.

WIBLE

AND

H.

H.

COVER7

Results Euprimates

monophyb

The Euprimates includes offossil been

of Hoffstetter

the modern

lemurs,

forms,

including

proposed

in the

(Martin,

euprimates

soft-tissue

features

galagos,

the Eocene past

1986; Gingerich,

2). Modern

( 1977)) the “primates lorises,

adapids

(Gingerich, 1986) holds

and omomyids.

that

aspect”

monkeys,

Euprimates

from other

of Yimons

el al.,

contrary

1978),

mammals

views have

current

is a monophyletic

1986), but most of the soft-tissue

( 1972))

apes, and man and a range

Although

1976; Schwartz

are distinguished

(Martin,

of modern

tarsiers,

by a number features

consensus

group

(Figurc

of hard-

cannot

and

be identified

Figure 2. Cladogram illustrating thr generally accepted hypothesis of relationships within the monophyletic group Euprimates. The unresolved trichotomy reflects the debate concerning the affinities of the living tarsier Tarsius. Some authors (Szalay & Delson, 1979) ally Tarsius with omomyids; while others (MacPhee & Cartmill, 1986) make it the sister group to the anthropoids.

in fossils.

A survey

of the distribution

Mammalia

reveals

others

relatively

are

euprimates,

eight

that

only unique

(Table

are identifiable

Adapidae

and Omomyidae

exception

is the orbital

of hard-tissue

one feature

1).

Of

unique

these

in representatives

(see Szalay

exposure

features

is totally

& Delson,

of the ethmoid

one adapid,

the Cambridge

skull of Ada+

the absence

of this

feature

in other

adapids

distribution

among

modern

lemurs,

lorises,

nine

derived

omomyids

galagos

and

euprimate

tarsiers

groups, 1986). The

thus far in only

& Martin, and

eight

of modern

& Cartmill,

has been found

(Gingerich

across

while

features

of the two Eocene

bone, which

euprimates

mammals,

1979; MacPhee

paririensis and

of modern among

1981). Given

its somewhat (Cartmill,

variable 1978),

an

exposed orbital ethmoid does not represent a very efficient diagnostic character for Euprimates. Recently, MacPhee et al. (1983) h ave raised doubts about identifications in fossil forms of the one wholly unique euprimate character, a petrosal auditory bulla (Figure 3). These

PRIMATES:

liable I

Derived Features

universalI>-

DIAGNOSIS

AND

RELATIONSHIPS

features

diagnostic

for Euprimates

present

for

ruprimatcs

extant

and

unique

for

mammals OSSFOUS auditory hulla completed h! an ~~t present for extant cuprimatrs and unusual for mammals intratympanic portions of the facial ncrvc and carotid circulation enclosed within canals by outgrowth, of thr pctrosal (MacPhee, 1981) convergent orbits - postorbital bar flattened nail on the hallux Features nearly universally present lix cxtdnt ruprimate groups and unusual for mammals exposure of ethmoid (OS planum) in the urblt ((Iartmill. 1978) distinct lateral trochlcar margin of the humrrus (Szalay & Dagosto, 1980) opposable hallux anterior (distal) elongation of the calcaneum (blartin, 1979) well-developed groove for the tendon of flexor digitorum libularis on the plantar surface of the calcaneal sustentxular process (Szalay & Drawhorn. 1980)

foramen

ovale

petrosal (cut) ‘tegmen

bulla

tympan’

epltympanic

crest

epitympanic

recess

facial

canal

stylomastoid

carotid

internal

canal

y

foramen

postdrior

carotid

toramen

Figure 3. Diagrammatic representation ofbasicranial structures in an adapid euprimate (modified after MacPhee & Cartmill, 1986). The petrosal bulla and ectoty-mpanit are cut and the middle-ear ossiclcs removed to expose the hard-tissue contents of the middle-ear space. Not labelled is the stapedial artery, which arises from the internal carotid on the posterior aspect of the promontorium of the petrosal and runs anterolaterally within a canal (a portion of the stapedial canal passes through the epitympanic crest). The portion ofthc tympanic roofimmediatrly in front ofthc rpitympanic crest is derived from the trgmen tympani, an element of the dewloping chondrocranium land not properly an element of the adult skull). Abbreviations: AS-alisphrnoid; BO-hasiorcipital: EO-woccipital; SQ-squamosal.

8

J. R. WIBLE

authors

specifically

is a general

address

issue

development

here.

an osseous

sutures

develop

et al.,

(MacPhee examples

among

expanded

tympanic

modern

complete

fusion

the modern MacPhee

ofthe

because

euprimate

that

only a single

omomyids

have

petrosal,

(i.e.,

an

the appearance

On

elcmcnt

bulla

independently

as do

is a pctrosal

in basal

concepts

01

hulla,

cuprimates)

hand,

is

if adapids

secondarily

fused

of a petrosal

in adapids

half

hold,

ha\~c a petrosal

hulla

(e.g., the appearance

rodents,

fi)r nearly

current

the other

an bone

among

omomyids

and omomyids

or cntotympanic

of either

we still

given

of a petrosal

lift>”

numerous

entotympanic

is characteristic and

rodents)

cites

composed

iYonethelcss,

solution

distribution.

arc required

and the loss of that

hulla

in adapids

2). If adapids

cctotympanic events

bulla

(1931)

is not a rare cvcnt;

1985).

parsimonious

for the observed

then multiple

euprimates

most

some

in postnatal

or an independent

and pctrosal

the auditory

as a pctrosal from the ahscncc

the petrosal

Klaauw

hut there of prenatal

cstahlished

(for example,

with

auditory

& Parent,

(Figure

event

to account

an bone)

analyses

composition

mammals is valid;

ectotympanic

it is the

hullar

In fact, such fusion

(Lavocat

interrelationships

required

p oint

(see below).

through

be properly

about

where

petrosal.

of the expanded

et al. (1983),

can

seamlcssly

(the cctotympanic

superfamilies

outgrowth

Their

mammals

ring

fuses with the surface

fuses

primates

it is only

of extant

which

p. 510).

archaic

that

bulla

a number

bulla

1983,

argue

to infer anything

in adults;

a non-petrosal

H. H. COVER1

in certain

They

auditory

“It is not possible

of bullar

then

this problem

involved

that

outgrowth.

AND

and

with

the

bulla in basal

and omomyids).

Plesiadapiformvs The plesiadapiforms, Late Eocene or partial been

the archaic

fossil forms,

crania

assigned

dentitions

are known

major

Monop/&y.

there

Firstly,

authors

carpolestids,

and

apatemyids). effective

saxonellids;

of the

identifiable

literature

shared

such

height

(Szalay,

groups

of Paleogene

derived

derived

of archaic features

are that

features, of stylar

1969). Such

trends

cutherians, primates

from those

about

known

primates

reveals

as reduction

the archaic

cannot

of the various

thought

to be closely

related

to archaic

without

microsyopids

(or apatemyids).

with

must

to group

be distinguished Paleogenc

primates, (Wood,

by discrete,

groups

Instead

and

(3) postcranial.

teeth, A

of distinct,

dcntitions reduction

consist

but occur identifiable

Plesiadapiformes,

01

of trigonid in other

1962). Perhaps

the

shared have hccn with

or

taxon.

Relationships. Are some archaic primates the closest relatives of ruprimates? three sorts has been used in the past to ally archaic primates and euprimates: basicranial,

possibly

and

features.

that over the years

If so, then

is not a monophylctic

(and

to dental

cusps,

rodents

primates.

plcsiadapids,

from jaws

primate

to archaic

paramyid

primates.

in the archaic

microsyopids

bulbous

(1) arc

of the ruprimatcs?

picrodontids,

exclusively

of archaic

& Gingcrich,

be addressed:

are wanting.

more

arc not unique

for instance,

have

are rcstrictcd

diagnoses

lower,

postcrania

in association

relatives

1986) add almost

diagnoses

shelf,

numerous

primates

necessarily such

Whole

1979; Gunnel1

paromomyids,

to

and teeth.

found

the closest

what

of Late Crctaceous

from jaws

been

& Delson,

(Gingerich,

forms

for archaic

trends,

dentitions

others

most

have

or all of them

1979) include

array

and although

(Szalay

is disagreement

& Delson,

Because

diagnoses

perusal

concerning

(2) are some

(Szalay

a diverse exclusively

postcrania

Plesiadapis

questions and

Some

primates,

for only one genus,

1987). Two

include

are known

for only live genera,

to archaic

they monophyletic

primates,

most ofwhich

Evidence (1) dental,

of (2)

PRIMATES:

[ 1) Dental the basis

evidence:

of the sorts

numerous

authors

as a group, of dental

(Gidlcy,

similarities

in the molars

particular,

the plesiadapid

argues

that

interpret

the

them

P(esiadapis

or possibly

primitive

link

because

with adapine

have been linked In addition,

plesiadapiforms

and primitive

thcsc

forms

because

of the derived

the most

primitive

I,); Cant&s.)

notharctincs

share

absent

br

through

plcsiadapid

ol‘d~i\~-d

features

A second

into-rpretation

group

bctwccn

Plesiudupis

wcrc present

of these

of Euprimatrs and

primi tivc omomyid until a detailed

forum

ancestor

similarities

( T&zardinn),

character

analysis

primitive

of omomyid,

for a detailed

discussion

adapinc

some

(:In)

unknown

would

rcjrct

and skeletal

is that Ploiudupis this hypothesis,

and this

fcaturra

cuprimate

features

of the dental

morphologic-s notharctinc

of these forms is completed,

is

similaritics

and nrcessaril)

and adapinc

and

01‘ thv and

or Plrsiadapidac all dental

notharctinr.

(Donrussellia).

of the molars

is that

4(a)].

in plrsiadapida.

4(b) 1. Under

Cuntiu~ arc retained

in the common

This is not the proper

molar

[ Figure

shared

lacks the upprr

cranial

to

for hypotheses

Fiqurr 4. Implications liir phylogcnetic relationships acccptiny that thr molar similarities plcrladapid Pk~s&zpi.r and thr notharctinc Cantzu! arc synapomorphirs. Either: (a) Plesiadapidar Y~~tharctinac arc sistrr taxa; or it11 I’lcsiadapidac is thr si\trr group to Euprimatrs (SW trxt).

the sister

in

( 1986)

it is difficult

[Figure

if not all. rcscarchcrs

a number

cuprimatcs

to notharctines

known

Most.

euprimates,

one interpretation

Cuntiu\ must

and

molar

h)

are very striking

implications

plcsiadapiforms,

is the sister group

retained

and omomyid

many

in other

that

on

noted

Cuntius. Gingcrich

are so minute

have specific

For one, because

Pkriudupis

adapid

to euprimatcs

as has been

1986). there

Such statements

Plrsiadapidac

plcsiadapid,

hvpothcsis,

bctwecn

between

first prcmolars

primates to above.

and the notharctinc

Cuntius arc lacking

and

phylogcnetic lower

resemblances

9

REI.AI‘IONSHIPS

1935; Gingerich,

advanced

Plesiadupis

relationship.

by Pletindupis

alluded

1923; Simpson, of some

AND

the archaic

trends

as homoplasies.

of phylogenctic

DIAGNOSIS

euprimatcs. of the most

( Cuntiu.r), but we belicvc

it is

prcmaturc to accept Cm&s as more primitive than thr other firms. More important, rrccnt analyses by Covert (1985) reveal that more primitive species of Can/&s (C. ejp.vi, (,‘. ralstoni, and c’. /orresi) do not exhibit the derived more derived species of Cantim. These similarities

(2) Basicranial Phtmmlemur

jepJeni

evidence: (Szalay,

basicrania

molar features shared by Plesiudapi.1 and must have been independently acquired.

have been dcscribcd

1972) and I
in detail

for thr paromomyids

(hlacPhee

tf ul., 19831, the

10

J.

Plesiadapis Microsyops

plesiadapid microsyopid Cartmill,

1986) agreed

primates:

a microsyopid

more

derived

most

substantial

R.

WIBLE

1963; (Saban, latidens (Szalay, that

there

type,

H.

H.

Russell,

is very

for excluding

types

close

type.

MacPhcc

It is generally

are two basic

which

COVERT

1964;

1969).

paromomyid-plesiadapid support

AND

et al.,

(Szalay,

of basicrania

to that

of ancestral

from

1983))

the, 8r

among

It is this observation

microsyopids

and

1975; MacPhee these

archaic

cuthcrians,

that

and

has provided

Plcsiadapiformrs

a

the

(Szalay,

1975). Do either of these two archaic primate those of euprimates ? Two basicranial characters

listed

of the carotid auditory have

in Table

1: a petrosal

circulation

bulla

a slight

and

is not known ridge

along

nerve.

1981; MacPhee

et al., in press).

1986; MacPhee portions ventral

of the carotid surface

opportunity from

of the petrosal

to study

the Middle

12363) collected J. G. Eaton

and facial nerve; (Szalay,

Eocene

is in preparation).

these structures

(Geological

Muscum,

(a full description The basicranium

is more problematic.

Closed

between

these

two views

canals

characters

indicate

that these vessels

Although

were probably

the identification

the intratympanic

routes

of arterial

canals

of Wyoming

No. and

exhibits

but we agree in adult

with

carotid

and

circulation

MacPhee

have

paromomyids

is questionable,

of both

& Cartmill

and stapedial

there

paromomyids

arteries

and plesiadapids.

is clearly

of the facial nerve in either paromomyids osseous auditory 1963). E vi d ence for a complete in basicrania

paromomyids

but more recently, MacPhee rt al. Having not studied all specimens,

portion

plesiadapids (Saban, with the petrosal is apparent

the same suite of

of the carotid

for the internal

vestigial

had the annectens

by M. ,J. Novacek

in known

for portions

is not possible,

(1986) that the size of the possible

for the intratympanic

University

of this specimen

is not a

(Novacek,

ran in open sulci on the

of this specimen

been described (Russell, 1964; Szalay, 1972, 1975), (1983) have argued that these “canals” arc imperforate. a choice

clearly

cuthcrians

0 ne of us (J.R.W.) has recently skull of the microsyopid Microsyops

characters described above for M. latidens. Identification of the two euprimate basicranial plesiadapids

Paleogcne

(Szalay, in modern

1969).

preservrd

of Wyoming

by J. G. Eaton

of the petrosal

1986). Yet, this ridge

of other

portions

An osseous

plate develops

in M. latidens are canals

Also lacking

a beautifully

features.

with

cuprimate

1969). .blicros_yops latidens does

the petrosal

& Cartmill,

the

the intratympanic

lack both

(Szalay,

in basicrania

circulation

around

resrmblances

among

of the promontorium

to the area where

nor is it uncommon

unique

included

Microsyopids

aspect

euprimates

(MacPhee,

are

for any microsyopid

the medial

in position

bulla

types share

features

bulla and canals

facial

1969) that corresponds petrosal

basicranial

no canal

for

(Szalay, 1972) or bulla continuous and

plesiadapids.

the previously cited caution of MacPhre et al. (1983) concerning the However, identification of petrosal bullae in fossils is still relevant here. Unlike the adapid-omomyid example

discussed

above,

paromomyid-plesiadapid

parsimony bulla.

does not resolve

An equal

number

this problem

of steps

for the condition

is required

to obtain

of the either

a

petrosal bulla or a non-petrosal bulla fused secondarily to the petrosal in paromomyids and plesiadapids from ancestral conditions of an cntotympanic, cctotympanic, or no osseous bulla

(Figure

demonstrable

5). We agree in paromomyids

with MacPhee

et al. that the presence

of a petrosal

bulla

is not

and plesiadapids.

The only other region of the cranium that supplies characters allying some archaic primates to euprimates is the wall of the orbit. In most eutherians and in marsupials, the maxilla makes no contribution to the orbital wall; it is excluded by the large orbital process of the palatine

contacting

the frontal

and

lacrimal

(Novacek,

1986).

The

orbits

of the

PRIMATES:

DIAGNOSIS

11

AND RELATIONSHIPS

Figure 5, Character phylogenies accounting for the hullar morphology found in paromomyids. plesiadapids, and euprimates. In the examples shown here, an entotympanic bulla is taken as the primitive state, hut either an ectotympanic hulla or no osseous hulla as the primitive state yields similar results. The bullae of paromomyids and plesiadapids have no visible sutures with either the promontorium of the petrosal or rctotympanic bone. [Unlike the separate, ring-shaped element of lemuriforms and adapids (Figure l), the ectotympanic in paromomyids and plesiadapids is part of the bulla, as in lorisiforms, omomyids, tarsiers, and anthropoids.] In the examples here, the paromomyid-plesiadapid bulla might be either (a) a derived petrosal bulla secondarily fused to the ectotympanic or (b) a retained entotympanic hulla secondarily fused to the promontorium ofthe petrosal and ectotympanic. In the former, a petrosal bulla is a synapomorphy ofparomomyids, plesiadapids, and euprimates; in the latter, the petrosal bulla is restricted to ruprimates. ;\n equal number of steps is required to account for either.

Microsyops lundeliusi (Szalay,

microsyopids No.

12363)

archaic

exhibit

primate

that contacts contact

this

whose

pattern. orbit

the frontal;

the frontal

is known

the palatine

(Russell,

(Le Gros Clark,

1934; Russell, euprimates

most

modern

orbital

contacting

the frontal

very

unusual

one data).

(3)

Postcranial

specifically

that

are

Drawhorn,

1980). Most

Drawhorn

(1980),

The

associated

dermopterans

is the

digitorum

fihularis

reliability

of this

well-developed

groove

occurs

it does

in adapids

1960; Wilson,

only

distinguishes

presence

exhibit

free

1966),

and

however,

to Plesiadapis

according

Plesiadupis

and

euprimates

from

pronounced

groove

of all, although

relatively

concomitant

for the We have

unusual

increased

in c(r and

and dermopterans

feature,

of the calcaneum.

also found 1977; Szalay

tarsal

surface

presumed

(Szalay,

(tree shrews)

P. gidleyi),

and

of specializations are not unique

scandentians

rodents,

19856, 1986; Wible?

(P. tricuspidens

of the foot

process it is not a

insectivorans,

(Novacek,

a number

mobility

orbital

and euprimates,

derived

of a more

First

a maxillary

in lipotyphlan

of Plesiadapis

tarsals

for modern

The

Although

of Plesiadupis

appearing

with

on the plantar feature.

other

of the maxilla although

pattern

& Russell,

and some edentates

of these specializations,

lemurs). that

(Simons

feature

and calcaneum,

but are also known or flying

process

to the floor of the orbit,

1976). The same

ethmoid.)

derived

eutherians,

evidence.

euprimates

(colugos

of the

proboscideans,

the astragalus

euprimates,

but

among

unpublished

orbital

1978). (A n orbital process of the maxilla occurs in usually does not contact the frontal because of the

is a shared

hyracoids,

is confined

1964), omomyids

exposure

lagomorphs,

has a large

the only

(Cartmill,

euprimates,

intervening

in detail,

of Wyoming

Plesiadupis,

the plesiadapid

1964; Gingerich,

and some modern

M. annectans (University

1969) and

In contrast,

to Szalay tree tendon

doubts

among

muscle

shrews

& and

of flexor about

the

mammals,

mass

for

flexor

a

12

J. R. WIBLE

fihularis

also appear,

for instance,

AND

in some

II. I{. COVERI

lagomorphs

and rodents

(Covert,

unpuhlishcd

data). Second and more importantly, a well-developed groove for flexor fibularis dots not appear in the calcanca of all Plesiadapis. One of us (J.R.W.) has rcccntly had the opportunity to study the skull and associated partial skeleton of Plesiadapis cookei from the Early

Eocene

discovered specimen does

of Wyoming

by these authors

not show

any

Plesiadapis

and euprimates.

consists

archaic

shares

the

of shared

evidence

dental

primate,

several

not unique

derived

arc distinguished ~c.g., reduction pctrosal).

for flexor fibularis

features forms.

the

of the orbit

from microsyopids

by several

Derived

carotid features carotid

(c.g., canals

around

the internal

the evidence

from

the basicranium

\+ith paromomyids

shared

also

derived

and stapcdial

cuprimatcs

features

in that

hut again

thcsc arc

it

of thr hasicranium continuous

cuprimatcs

artcries,

Euthcria.

cvidcncr

and plrsiadapids

a bulla

from

a petrosal

with regard

this

for allying

and

paromomyids

distinguish

is equi\,ocal

primates

additional

artrrics,

was

of this

Givrn

character

with cuprimatcs,

primates,

and stapcdial

basicranial

fibularis.

to these
provides

and tarsus

the archaic

which

the only one prcsrrvrd.

of flexor

archaic

is unique

Plesiadapis,

Among

of the internal

and 69934),

(a full description

is not a reliablr

various

none of which

the plesiadapid

to these

for the tendon

for allying

trends,

87990 (1987)

l’hc right calcaneum,

excavation

groove

Nos.

& Gingcrich

is in preparation).

a well-developed

In summary,

of Michigan

in Gunnel1

pronounced

\.ariability.

One

(University

in 1986 and reported

with

chc

microsyopids

bulla).

to the affinities

H0wrvc.r.

of cuprimatcs

and plcsadapids.

,Superordinal relationships In the past allying

twenty

primates

years, with

morphological

a diverse

1986 alone, primates have srquencc data (Miyamoto scandentians

based

ans (Old World Cooper,

of modern

features

1986). For the molecular

that some

thrir

based

Euprimates.

primates

plcsiadapiforms

have provided

orders

support

fbr

groups.

In

and extinct

includes

dcfinc

usagr.

primates

(.wnsu Into) arc rcvicwed

studies,

current

Novacck

although

(Prttigrcw

data

Br

(Shoshani,

primates

is equi\.alcnt

stated.

it is not clear

explicitly

& Wyss

hypothcsrs

data

distance

the grouping

unless

to the paleontologist;

Several

1986), with mcgachiroptrr-

on immunodiffusion

Howrvcr,

may be included.

explicitly

Br Wyss,

based on ncuroanatomical

and neuroanatomical

taxon

what the grouping

(Kovacck

and dermoptcrans

1986), and with scandcntians

to the monophyletic

studies

euthcrian

been linked with rodents and lagomorphs based on protein & Goodman, 1986), with chiropterans, dcrmoptcrans. and

on postcranial

fruit bats)

and molecular

array

their analysis

(1986),

suggrsts

for instance,

of thr higher-lcvcl

do not

relationships

of

in the following.

Rodents and lagomorphs.

Similarities

plcsiadapiform rodents within

led McKenna ( 196 1) and Wood (1962) to proposc an origin for (rensu lato). McKenna ( 1975) later abandoned this hypothrsis. and

support

for a primate-rodent

‘I’hc most that

widely

rodents

Hartenberqcr, \Vyss,

primates primates

be grouped

with

of Paleoccnc

morphological

on data

for hlacroscclidca

systems

have

(Novacek, offcrcd

(elephant

rodents

and

has not appcarcd.

morphological

in the Superorder

rcscarchcrs

in press)

paramyid

from various

lagomorphs

scvcral

1986; Wible, anatomy

molars

from other

view based

1985). Additionally,

1986; Shoshani,

c.raniaI and postcranial (ilircs.

clade

held current

should

in the

Glires

systems (Luckctt

1982, 1986: No\.acck

considerable shrews)

support

is St &

fi-om

as the sister group

to

Molecular Albumin bats,

studies

carnivorans,

Sequence places

analysis

tree

advocates

Goodman,

1986).

Of the features

analysis other recent

analysts

process

contacting primates

Rose,

for

Dac,tulopsi/a,

have

proposed

from

(Russell,

possihlc

8r Fitch.

clata

ha\.c

the

apomorph)

tandem

alignment hypothesis

but appears

1981; \\‘yss raised

Cy:

with rodents

this

in sc\rcral

et al., 1987) in

questions

about

the

the

brtllac

Civctt

that

wall

an \vith

Daubentonia).

occasions

but

198.56). In othct

primates

Particular

region

not

incisors

fcaturc.

has

in

is obscured

prcmaxilla

t)!

with (e.g..

prrsumabl!

a the

bc~cn

197 I).

ha\-c~ h(.cn used to support close aftittitieh I,cuprimatc,s and non-micros) opid

sigttific,ancc

in both groups’

dots

Zgnarius grgvbullianu~, this

enLtrgcd

(Gingcrich,

similarities

1976. and and

procumbent

this

its

posterodorsal

1969; the carpolrstid

is restricted

paromomyid

frontal.

mamm;tls

bullar

basisphcnoid (Butler, h)pothcsis with other

1976: Novacck,

prctnaxilla

ib thr the

and

lagomorphs;

Phenacolemur pagei, Gingerich,

1977),

cuprimatc

(1973).

and

A\licro.yop~~Iatidens, Szalay,

1976).

by Szalay

rodents

with an expanded

1961; Gingcrich.

in other

on sc\,cral

\\ith

prcmaxilla

contacted

iriSecti\YJranS

a pctrosal

fcaturc.

exception

have

to the auditory

in the orbital

lo these forms,

(hlac,da

scqucncc

drrivcd

/nsectirrorans. Basicranial

crinaccomorph

contribution

cladc is not unique

& Cartmill,

may

indcprndcntly

plcsiadapiforms)

in tandem

(Miyamoto

share

Although

1973; the paromomyids

One

prcmaxilla

Erinaceomorph

and cuprimates

beta

bats. and

combined

and cuprimates

maxilla.

authors

the microsyopid

fractures (Rose & Gingerich, postcrodorsal process appears acquired

and

WC do not find this phylogenetic

lie in an enlarged

the frontal (i.e.,

the frontal.

marsupial

process several

additional

nacirnienti, Kay

the

(1986),

of alpha

of the

but

lipotyphlan

of lagomorphs,

proteins

with 1985).

Glirvs,

euprimates, analysis

of scorn

Pl~sindupis

process

alignment

incisors

Carpolestes dubius,

which

orbital Also,

an

upper

archaic

contact

orbital

not contradict

a grouping

lagomorphs.

(Sarich,

of this tcchttiquc.

shares

Palaecthon

in our rev&v,

of tandem

power

procumbent

analysis

and lagomorphs.

but ally rodents

(shrews)

does

S e q urncr

supports

while

soricids

protein

1985).

for rodents,

& Goodman

groups.

and

ofrodents

for Glires,

the lagomorphs,

rodent-lagomorph-euprimatc

A maxillary

Plesiadapis

among

alignment

et al., 1985),

a derived

eutherian

resolving

(moles)

(De Jong,

a trichotomy

by Miyamoto

compelling.

shrews

relationships no support

A eyv-lens

of rodents

discussed

lagomorphs the

talpids

in tandem

(Shoshani

aligntnent

supports

provide

of the ol-crystallin

combined

tree shrews

views ofthc

data

probably

relatives

and

hemoglobins

dkrerse

distance

and

the closest

inscctivorans,

and

provide

immunological

is placed

on the large pctrosal

most ti)ssil and recent to a large tympanic

crinaccomorphs process of the,

component in addition supporttd this 1948; Rich & Rich, 197 1). N 01 ad ( 1982) tcntati\.el) shared derived features oftht, cranium ic.g.. n process of tltc maxilla

contacting

the frontal)

and

\vith

dental

rrscmblanccs

bct\\.c’cTtt the

carlicst kttobvtt archaic primatc%s. such as the paromortt),id PurgatoriuJ. and crinaccontorphs. Both Szala), ( 1977) and Xo\.acck (No\.lrcck pt oi., 1983) ha\.c, abandoned the hypothesis of ~losc primate-crinaceomorph

tics. Ho\ve\,er,

Szala)

continues

to argue

that “printiti\,c>

significant (basicraninl I sittii1aritic.s share some cn$ginatrcl and that it is “likel). that prin1,ttc.s and other archontans morphotype” with a basic.rnnilittt tiot v(‘t-)’ clitkrctit front that of frottt m adapisoricid inscctivoran, X-27). ‘I’tic pr~sutiicd the rrinaccomorph morphot) pc” (Szalav & Dc~lsott. 1979, 1’1’. 12.ith the

primates

dcri\.cd

frattlrrs

shared

b)

the

primate

( \V~I$USzala).

197.5) and

crinaccotriorl,tt

rrinac~c~rttorph

14

J.

morphotypes ventrally

in

are,

promontorium,

WIBLE

addition

bony canals

“shielded”

R.

AND

to

the

for the entire

by the internal

H.

H.

COVERT

petrosal

hullar

intrabullar

carotid

canal,

in ancestral

feature.

[Further,

modern

erinaceomorphs

state

(Wible,

Szalay more

eutherians

1986).]

& Delson recent

rotunda carotid

of the two cannot

as

out, because

of the basicrania

reveal

it appears

in many

features

cited

groups.

plesiadapiforms

by Szalay

1975) and erinaceomorph

carotid

mammalian

of non-microsyopid

that the other

(sensu Szalay,

of the single internal

be a derived of

point

position

internal

artery

analyses

& Delson

morphotypes.

Diacodon and Pholidocercus have insignificant

erinaceomorphs

rounded

(1979) and Wible ( 1984, 1986) (medial and lateral) were not

the loss ofone

or promontorial

a fenestra

and strepsirhine euprimates is likely a retained plesiomorphous promontorium of the pctrosal is of questionable value, A rounded

rightly

erinaceomorphs the primate

and, therefore,

the lateral

circulation,

and loss of the medial

artery (Szalay & Delson, 1979). The analyses of Presley have shown that two separate internal carotid arteries present

contribution,

carotid

Finally,

and

extinct

are not shared

by

First of all, the Eocene

petrosal

contributions

to the

auditory bulla (Novacek et al., 1983; MacPhee & Starch, 1987; MacPhec et al., in press), suggesting that these elements may have been acquired independently in other forms and are not part carotid

of the erinaceomorph

circulation

Diacodon (Novacek in press)

and

morphotype.

are absent

Secondly,

in paromomyids

bony canals

and plesiadapids

et al., 1983), and Pholidocercus (MacPhee

are not part

of either

the primate

for the intrabullar et al., 1983),

(MacPhec

& Starch,

(sensu Szalay,

et al.,

1987; MacPhee

1975) or crinaceomorph

morphotype. Archonta. Gregory’s (1910) include bats, dermopterans, plesiadapiforms).

This

in Plesiadapis,

tarsus

Drawhorn,

bones

grouping

of bats

Recently,

and

have

previously (1977; Szalay

sustentacular

and

microchiropterans Novacek & Wyss

additional

synapomorphy

penis is enclosed abdomen, caution

but in recent that

considerably

although within

presumably

a derived

members reverts

does

published

cladograms

not

genital feature

external pouch

is unique

of the group,

to the more

linking

have

restudied

megachiropteran and

a distinct

medial

appear

in any

is free and to

recent

including

some

generalized

sheathed

&

euprimates has

In most

astragalus euprimates

confluence other

pendulous.

ofthe facet and an

eutherians,

Novacck

Archonta,

by

eutherian,

the sustentacular with this fcaturc recent

the to the

& LYyss

conditions

microchiropterans

&

the tarsal

that forms a sling in close attachment

the penis

of the

of the Archonta

Plesiadapis,

genitalia.

features

1977; Szalay

1980, 1982; Cartmill

(echolocating bats), which lose (1986) tentatively support Archonta

The supporting evidence is not overwhelming, only hypothesis for the higher-level relationships based on uniquely relationships within

(1986)

in the

1980). This feature,

archontans this

1980). Critique

tree shrews,

facets,

data

of bats (Novacek,

feature

dermopterans,

of the male

in a sheath-like

derived (Szalay,

immunological

& Wyss

to

and non-microsyopid

by shared

& Sarich,

(1975)

by McKenna

and dermopterans

Novacek

astragalar

including altogether.

penis

(Cronin

found

revived (euprimatcs

supported

for the inclusion

& Drawhorn, distal

been

and transferrin

however,

for only

was

and primates

tree shrews,

and dermopterans

1980).

described Szalay

has

euprimates,

on the lack ofevidence

MacPhee,

Archonta

tree shrews,

1980) and by albumin

to tree shrews centered

S u p erorder

where

vary the

condition.

but Archonta currently of primates (euprimates

represents the and PIesiadapis)

derived morphological features. i\ccepting this hypothesis, can the superorder be relined further’ .’ Figure 6 shows two previously of archontan interrelationships. The two differ radicallv in that

PRIMATES:

DIAGNOSIS

AND RBLATIONSHIPS

(b)

(a)

Figure 6. Prrviously puhlished hypotheses ofarchontan int~rrrlationships. ia) From Smith 8 Madkour (l%O), hascd chiefly on features trf the pt-nis. (b) From Novacck & ‘i2:yss i 1986), based on cranial and postcranial features. Ncithw sft of authors sprcificall) discusses thp relationships of tht- archaic primates, although the distribution of characters discussed by Novawk & M’yss suggrsts that sonw plcsiadapiforms may hc includtd within primatrs.

Figure

6(a) depicts

Chiroptcra

as diphylctic,

while

in Figure

Bat diphyly, as proposed by Smith & Madkour (1980), Cooper (l986), is supported principally by differcnccs and

microchiropterans

cuprimates wings

and

of mega-

unexpected, within

Order

overshadowed mcga-

than 50 million (Novacek,

Wible

dermopterans

unique

& Kovacrk, 1980).

identified

and lost secondarily is an alliance

ofChiroptera

support

from

of both

hypotheses

(I) A grouping

of archontan

of cuprimates

shared

derived

Tahlc

I : a postorbital

cranial

clade

features

and

bar and canals

around

is completed by an rntotympanic features shared by cuprimatrs

in thr basisphenoid

1981) (Figure carotid

artrr)’

from outgrokvths

that is converted

anterolatcrally 3). The enters

anterior thr

and

Included

artcry

expanded

Archonta [Figure

offered

6(b)].

arc

immunological

data

cuprimatcs,

(1986),

to

1982; Wihle. and

and Pettigrew

or some

subset

by Novacek

& \Vyss

This relationship

postcranial

anatomy

8r

thereat

seems

( 19863 sccur(*.

(see also Novacck,

for the following

tree shrews

carotid artery Other derived fi)ramcn

albumin

for Archonta

is acceptrd

and stapedial

greatly

are restricted

evaluation

of sevrral

interrelationships.

of the cranium.

formed

differences

that

(see below). within

and Dermoptcra

features

of these groups

1979; Novacrk,

(19801, Pcttigrcw

in the

but are not

slight

of mcgachiropterans,

may be plesiomorphous

1986). A Chiroptera-Dermoptera alternative

& Madkour

of relationships

features

Valen,

1978),

evolution these

and by supporting features

in microchiropterans

The only refinement

Moreover, (\-an

Differences

(Strickler,

and postcranial

eutherians

derived

by Smith

recorded

years of independent

in press)

The

hats are monophyletic.

in microchiropterans.

been

1985a).

cranial

among

( 1986) alternatively

Cooper

have

Chiroptera

& Sarich,

features

are absent

the more

and microchiropterans

(Cronin

derived

that

by the many

1984, in press;

with

by

and microchiropterans

given

the

and

dermopterans

6(h)

Pettigrrw (1986), and Pettigrew & in the flight apparatuses of mcgathat megachiropterans share with

[E’igurr

the intratympanic of thr pctrosal

carotid

by srvcral

synapomorphies

portions

from

of the facial nerve’

(the canal around

into a long tube and a tcgmcn middle-car

foramrn. the opening usually a simple

is

is supported

the internal

clrmcnt in tree shrews) ( MacPhee, 198 1). and trcse shrrws include an anterior carotid

to cover the entire

hraincasc,

7(a)]

are two cuprimatc

ossicular through opening

tympani chain

which cithcr

that is

(hlacPhrc, the internal \zithin the

J, R. WIBLE

16

AND

fI. H. (:O\‘ER’I

(a)

(b!

v v I’

4)’

(d)

(cl

basisphenoid or foramen appears brachycricine 1984). l‘he forward ofthe

between the basisphenoid and to be somewhat unusual among

crinaceomorphs tegmen

(Rich

tympani,

from the anterolatcral tympanic

and cuprimatrs.1 and strcpsirhine an incomplctc

roof, is typically In addition primates osseous

portion

1971) and some

of the mammalian

of the auditor!,

small in cuthcrians character&d

bordering

microchiropterans

capsule

portion

1937), but not in tree shrews

tympani margin

prqjecting

to fbrm the posterior

by the presence

thr anterior

(CViblr.

chondrocranium

(De Beer,

to its large size, the tegmen

is further septum

t(r Rich.

an element

pctrosal. A tubular anterior carotid other eutherians, although it occurs in

in modern ofan

tree shrews

cpitympanic

of the epitympanic

crest, recess

I98 I : Zcllcr, 1986) (Figure 3). <\rnong within which the stapcdial artery runs (AlacPhr~, other archontans. the tcgmcn tympani is lost in dermoptcrans (Halbsguth, 1973) and is a

slrndcr,

vertically-oriented

press:

W:ible

tegmrn

rod

& Novacrk,

tympani

have

a hat

in chiropterans,

in press).

not brcn

:1 tubular

synapomorphy

anrcrior

carotid

for PleJindopis

idcntifird

(1Vihlc.

foramcn

or othrr

I!##.

and

archaic

in

cspandrd

primates.

The

tympanic roof is relatively broad in Pkrindapi~ (SalIan. 1963: Russell. 1964) Phetmolemur (Szalay, 1972). IIUI sutures dcfirtiqq the limits (nf’cltc pctrosal contrihurion not c\idcnt. (2) To ally euprimates maxilla

must

discussed

above.

not c-c)n\:inccd cuprimatcs

Gi\rrn

mot-c’ hcavil\-

7111i 1, the sharrd

is not unicluc

gr(JUping

orbital

tltc c,rtprimatc-trc-cx

thrsc.

shrew

is

nicw

tltc

01’

and PIpriadnpi\, \VCarcs

to cltprimatrs

fi)intis

prorcss

s~napomor1)hic.s than r)tw q-oul’ittq

twdvcd

tree shrews.

ottI!- prcsumcd

1Figure

than

that this fraturr

that a cladogram

and

(3) l’hc bats

I’le.riatia~i~ [Figure

and

lx wcightcd

and arc’

7(c)]

clcrivcd

fcaturc

is an auditory

hulla

idcntific,d

formed

ti)r trw

from

sltrcws.

indcpcndcnt

dcrmoptcrans.

wtotympanic

attd clcntvnts

1977; Cartmill & hIacPhw, 1980). Yet. the hrm~ologics of the c.tttotympattics itt groups arc uncertain: d~~~~clopmcntall~, trc*r S~W\V> Itit\‘? only OIIC wtot)mpanic~

(Novacch,

thcsc

c~lcmcttc (hlacl’hcc. (\VitJlv.

1981; Zcllrr,

alwrttati\~c,

,qi\.cv

Xcnnrthra,

thr

prcsrnc-c

Carnivora,

of ctitot)

I’rttigrw

nic,~arliit~optcratts

B (1oopcr.

1986). Hocvc\w-~

distrihrttirm

of thcsc

charactcrx

character

conwrns

tit<, urethra.

In the state

corpus

Sp(JIl,qiOsUni

prni,.

In

is

contrast, is highly

homolo,qirs

oi‘ this variahlc.

cx)rpus

not

In fact,

PrItiq-rb

aftinitirj. Earl\

Ttw 1:occnc

l~laqiomrnid

basis

clcntitiotts

a

vu)

pctrosal

against

larqc

for

‘1‘1 1~

ot%‘yomin~

drrmoptrtxn

contribution

cotttrihution

which

Ixcauw

19861.

horn that

thr tcgmw

Yc,r. rhc,

of rspansiotl

1101

rxpt~d~~~l.

drscriptions

may

ot’ rhcw, ph~loc?_c~ttc~1it~

thrsc ha\,c

tixtnh

known,

to tht.

dcrmo~)trtxtt

dcrmoptrtx-like arc

‘1‘1~

in tlt(, \.isttal

asscssin~ allird

i\

& hladkour.

tarxals hut

irt

t;)r \vhictt

hrlrl to tx fossil drrmoptrrans of’ tttc hliddlr Eocrnr .\licto.~w~p\ .. aho\x~ wxxxLs details of~hc lxtsicrxtturn

In particwlar.

pctrosal:

of

dcntitions

12363) discuswd

tYom rhc

I 1WI) hw micros)x)pitls

[plagiomcnicls discovrrrd skull

afftnitics.

lwtilrc

of‘ tltc,

tltv c3~rptls

patlibvays

I)rt;tilcd

is the occ‘urrww

rawly

portion

1)~ Smitlt just

c~trwt.

of‘ wrious

c~uthrr~iairs

microsqpiti

arc not known

1977)].

(LItlit crsity

that argur

8~ I)ra\\

spcculatc

for this sprculation

localities

(ROM, CyrSimons, nntw/m

fit.’ Sxalav authors

oth

i 1980). th(.

of the penis. dc,grw

\tudicd

to sonic

& C~oo+.

thr

surrounds

Sr hbdkour

the ql;ttts that

I)( tttc,

and or(lrrs.

that

to tlw proximal

mic,t-o~hit,c,l,tc,ratls

fi~r

of tbrms

nic~:;icltiroptcratis,

itlto yivvn

in tliv glans

complrtcd

thcsc

crpand~d

must

in particular.

tissw

1,) Smith

arid

(x)iicrt~ti the* projcctic~nx

~~‘lwrc do tnicrosvopids Arcl~ottta.

and

ttumtx-r

is rrstrictc,d

l!)Xt?

ly, thcsc fbaturcs

clic crccrilc

for Ertthcria

qucstionahlc.

is prcwnt

hr

for :\rchont,t.

Pcttigrc\v,

hcrc,. Ixxtusc.

spottqio~um. and

rr. in most

1986:

should

I c.,q..

ourgroups

1980:

IJat mottopIt) cxulicltt

dcrmoptcratts.

mm

characters

archontan

till- a ratltc-r lintiwd

cqanizcd

vascularixd hlorw\

(Pcltigrcw,

char-actors signiticancc.

is knwvn the corpuc

M-C]]

in w\w;tl

to maintaitt

tar l)r primitive

fcaturt.

spoti~iosuni

ttcut~o;tttatomi~~~l sytcxi

h&f

tta\x, (\\(I 19841. .\a :it1

mcrcl>~ may tjc primititc

&I .2lndkour.

\t’c sugyst

in cuprimatrs.

spott~iosum sotnc,\vhar

(Smith

in microchiroptcrans.

‘I’hc- pcnilc

mpatiic\

&c%sc c’lrmrnts

and

lost scxvndaril!,

:tncl nrirrocltiroptc~ran:,

with thr dt,rrncllJtc,r;ttt-clliroljtcran cladc [Figure, iid) 1ib p~~nilc and nc~~tro;tttatontic;tl c~haractcrs 01’ c.trprin1;ttc.b.

(4) The grouping of ruprimatrs chicflv , hvI dcrivrd

dcrntoptcrans.

tl~vmoprwarts

ha1.c up to t;)ur i I*;laau~\. 1932; \\‘ihlc.

Leptictida),

supported

thr

1986).

1984). and mcgacltiroptc,ratls

mod~w~

tymlxttli

arc

thr rOOf‘(Jf the t) mpanic. dcrmoptrrans

is :il)sc.ttt. .Ilthouqh

haw

ca\.it!

virtualI\

tltc mastoid

~JOII

has

no ion

18

J. R. WIBLE

ofthc

auditory

capsule

1969, Figure and has canals

for a major

1984, in press), into the hraincasc

The affinities tympanic

artery,

euprimates,

aspect

dermopterans,

these

pneumatizcd

the arteria

appears

to be absent the squamosal remain

is reminiscent

(Hunt

diploctica

orbit

come close to the pattern

Until

postcrania

found

magna,

muddled.

crest

is lacking.

reconstructed

(see Szalay, processes

& Berth, and veins

are 1980)

(\2:iblc,

Other

petrosal

portions

in tree

of

to the

shrevvs

of the basicranium

cutherians

dcntitions

from

foramen

contribution

tympani

for primitive

with microsyopid

its usual

is lacking.

The large tegmcn

( 1980), we SW no basis

Dcrmoptera

because

and mastoid

expanded

in association

& Drawhorn

in this form,

of the

yet an cpitympanic

with tither

is highly

between

of microsyopids

roof

of Szalay

this process

cranial

on its posterolateral

in recent

data). Evidence ofpncmatization is not apparent in ;M. annec~ans, nor are there canals. In fact, the arteria diploetica magna, a derived euthcrian character

unpublished any vascular (Wible,

flat eminence

the arrangement

In Cjnocephalus

not equivalent.

entry

has a broad,

17) resembling

AND H. H. C:O\‘ERT

and

by Novacrk

(1986).

the speculations

confirm

the cranium

and

for linkin,g

these

forms

or Archonta.

Conclusions The following (1)

conclusions

Euprimates

characters

of the

distributions

derive

cranium

among

Plcsiadapiformes

and

diagnosed

is the only higher-level

grouping

derived

characters.

morphological

have been found

unites

Archonta,

supporting

Microsyopidae

(4) Hypotheses

of

(5) Following Order

demonstrable;

is

not

of shared

derived

relatively

unique

that

exhibits

which exhibit

(and

the criterion include

cquivpocal. Therefore, the criterion group and its stem lineage cannot

shared for

character

Paromomyidac)

that within

with

either

‘The Euprimates-Scandcntia

in a number

homoplasy

among

several of which appear to is supported by a shared ofothrr

other

eutherians. cuthcrian

Relying orders,

the

is preferable.

for restricting

is equivalent

tarsal

ha\c archontan affinities. Archonta ally Euprimatcs

is also found little

by unique

the only plcsiadapiform

the unique

of Plcsiadapidae

a within

and Chiroptera)

that is diagnosed PlesiadupiJ,

group

of the basicranium.

derived characters of the cranium, The Euprimatcs-Plesiadapi.r cladc

hypothesis

it may

Euprimates

within

demonstrably

Dermoptcra,

(Paromomyidac-Plcsiadapidae).

of the orbit,

Primates

or

characters

Scandentia,

with dentitions?

the inclusion

by shared Eutheria.

Euprimatcs-Scandentia the

unique

form a monophyletic

derived

The plcsiadapid

relationships

characters

shared

does not demonstrably

or Plesiadapis

character

Microsyopidac)

and Plcsiadapidac

including

postcrania

on derived

analysis:

by a suite

without

(Euprimatcs,

which

derived

cladistic

diagnosed exhibiting

by several

Archonta

cladc is supported be unique among

group

postcranium

or (with Paromomyidae

(3) The Superorder

Scandrntia

the foregoing

Eutheria.

(2) Plesiadapiformes monophylrtic group.

Archonta.

from

is a monophyletic

higher

to Euprimatcs. Plesiadupis

taxonomic The

groups

ruprimate

to the crown stem

lineage

(Paromomyidae-Plesiadapidae),

of equating higher taxonomic be meaningfully employed.

groups

but with

group, is not this

the crown

Acknowledgements The authors (University

thank T. M. Bown (U.S. Geological Survey, Denver Station), P. D. Gingerich of Michigan, Ann Arbor), and M. J. N ovacck (American Museum of Natural

is

PRIMATES:

History,

New

comments Clark,

York)

for access

on various

J. A. Hopson,

to specimens

and

four

from

Science

AND

anonymous

in their

Foundation

care.

grant

For helpful

we are grateful

R. D. E. MacPhee, reviewers.

19

RELATIONSHIPS

of the manuscript,

J. M. Humphreys,

R. T. Zanon, National

portions

DIAGNOSIS

The

senior

M. J. Novacek, author

discussions

to M. Cartmill,

and J. M.

L. Van Valen,

acknowledges

support

DEB-8208797.

References hx, P. (1985). Stern species and the stem lineage concept. C’la&itirr 1,

27%%7.

P. M. (1948). On the evolution of the skull and teeth in thr Erinaceidae. with special wfercnce trj fossil material in the British Museum. Proc.
Evolution

ofF/i,ght.

Mon.

Cal.

Acad.

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