CALIFORNIA STATE UNIVERSITY, NORTHRIDGE

THE PHYLOGENETIC STATUS OF SIVAPITHECUS:

A CRITICAL ASSESSMENT OF DIVERGENCE HYPOTHESES
AND A PROPOSAL FOR THE DIVERGENCE DATE
OF HOMINIDS AND PONGIDS
A Thesis submitted in partial satisfaction of the
requirements for the degree of Master of Arts in
Anthropology
by
Deborah Elaine Elman-Whitchurch

January 1988
The Thesis of Deborah Elaine Elman-Whitchurch is approved:

Dr. Keith Morton

Dr. George Fisler

Dr. Bruce Gelvin, Chair

CALIFORNIA STATE UNIVERSITY, NORTHRIDGE

i i
 ACKNOWLEDGEMENT

First, I would like to thank the members of my

committee for their time, guidance, and patience. Second, I

would like to thank my good friend Elaine, who gave her time

and effort to inspire me when things got tough. Third, I

would like to thank my husband, Ed for his patience,

support, and hugs. And final !y, I would like to thank a! l

of my friends and family for their love and guidance. I

share this achievement with a! 1 of you.

i i i
 TABLE OF CONTENTS

Page

ACKNOWLEDGEMENT i i i

LIST OF TABLES AND FIGURES vi

ABSTRACT vi i

CHAPTERS

I. INTRODUCTION 1

A. Problem Statement 2

B. Problems with Divergence Dating 3

1. Dating Techniques 3
a. Potassium-argon Technique 4
b. Paleomagnetic Reversals 7
c. Fission Track 9
d. Faunal Correlation 9

2. Underestimation of Divergence Dates 10

C. Phylogenetic Analysis 14

II. The Sivapithecus Problem 18

A• 1 930 to 1 960 20

23

37

1. Analyses Based on Paleontological

Evidence 37

2. Incorporation of Molecular Evidence 50

3. Use of Molecular Data in

Paleonanthropology 51

III . The 1 980 / s 57

A. The Early Divergence Hypothesis 63

B. The Late Divergence Hypothesis 70

C. Sivapithecus is an Ancestor to Pongo 80

D. Ramapithecus and Sivapithecus as

Sympatrlc Populations 87

iv
 IV. Criticisms of Current Ramapithecine
Hypotheses 92
A. Early Divergence Hypothesis 92
B. Late Divergence Hypothesis 95
C. Sivapithecus is an Ancestor to Pengo 97

V. Conclusion 100

BIBLIOGRAPHY 107

~--
 LIST OF TABLES AND FIGURES

Page
TABLES

I. Revision of the Dryopithecinae 28

FIGURES

I. Phylogeny of Large-Bodied Miocene Hominolds 31

II. Phylogeny Supporting a Relationship between

Ramapithecus and Australopithecus 32

I I I. Three Alternative Hypotheses of Miocene

Hominoids 48

I ll
1 • Cladogram of The Early Divergence Hypothesis 69

v. Late Divergence Phylogeny 74

?I. Cladogram of The Late Divergence Hypothesis 79

V.I I. Cladogram of a Pongo Linkage to Sivapithecus 86

\' i
 ABSTRACT

The Phylogenetic Status of Sivapithecus:

A Critical Assessment of Divergence Hypotheses

and a Proposal for the Divergence Date

of Hominids and Pongids

by

Deborah Elaine Elman-Whitchurch

Master of Arts in Anthropology

This thesis is an attempt to estimate the date of-

divergence between the Hominidae and Pongidae. In order to

determine this date, the phylogenetic status of the Miocene

hominoids, Ramapithecus, Kenyapithecus, and Sivapithecus,

wil 1 be analyzed. If Kenyapithecus from the Early Miocene

of East Africa is included with the adaptive radiation of

Miocene hominoids classified as the sivapithecines, then the

earliest geological dates for African members of this

radiation are earlier than the dates for the Asian fossils.

Since advocates of the "early divergence" hypothesis <Middle

Miocene, ca. 15 Myr) consider sivapithecines to be a dental

vi i
hominid, addition of the earlier form Kenyapithecus to this

complex would, by inference, push back hominid divergence

close to 17.2 Myr. If Sivapithecus is not a hominid, as
postulated by supporters of the "late divergence"
hypothesis, hominid separation may have been as late as 5.5
Myr. However, another key element in predicting divergence
dates is the likelihood of underestimating the actual
divergence, as genetic changes are not immediately
distinguishable in the fossil record. A critical review of
the literature on sivapithecines suggests the following:
(1) Sivapithecus and/or Ramapithecus was not a hominid, (2)

the divergence of hominids and pongids occurred within the

radiation and evolution of the species within th~ genus
Sivapithecus, (3) the species of Sivapithecus which would be
ancestral to the hominid and pongid lineages, respectively,
were not distinguishable from the parent genus until they
evolved beyond the range of variability for the
Sivapithecldae, (4) the species that evolved into the
hominid line was not necessarily the youngest species, and

<5) the divergence of the Hominidae and the Pongidae may

have occurred 7.5-10 Myr.

vi i i
 Chapter I

Introduction

The purpose of this study is to discuss hypotheses

concerning dates for the divergence between the hominids and

pongids during the Miocene and the phylogenetic status of

Ramapithecus and Sivapithecus. A critical review of

published views wil 1 be presented and each viewpoint wil 1 be

treated both individually and with respect to its

interrelationship to the others. In conclusion, elements of

several views wil 1 be combined into one hypothesis.

The hypothesis derived from this study estimates a

range of 8-10 mil lion years <Myr) for the divergence of the

hominid and pongid lineages. In support or this estimate,

the phylogenetic status of Miocene hominoids, particularly

Ramapithecus, Kenyapithecus, and Sivapithecus, wil 1 be

discussed. This study contends that the divergence of

hominids and pongids occurred among species that were

evolving within the Sivapithecus radiation. Therefore, the

species of Sivapithecus that would evolve into the hominid

and pongid lineages, respectively, remained submerged within

the parent genus until they evolved beyond the acceptable

range of variability for the Sivapithecidae into the

Hominidae and Pongidae.

1
 2

A. Problem Statement.

In order to determine the divergence date of hominids

and pongids, the phylogenetic status of Ramapithecus and/or

Sivapithecus is analyzed. This is a vitally important

element of the controversy over the divergence date of

hominds and pongids. Whether Sivapithecus is considered a

hominid or not based upon fossils from Buluk, Kenya <Leakey

and Walker 1985; McDougal 1 and Watkins 1985), and

Haritalyangar, India <Sankyhan 1985:573), will indicate that

the hominid-pongid divergence occurred between 17.2 Myr and

5.5 Myr, respectively.

This thesis will concentrate on interpretations of the

position of Sivapithecus in human evolutionary history and

the effect of its position upon estimates of the length of

the hominid and pongid lineages. The study consists of five

chapters. The first chapter analyzes a variety of

paleontological dating techniques, their problems. and

examines the topic of underestimation of divergence dates.

The second chapter discusses the historical background of

sivapithecine analyses spanning the 1930/s through the

1970/s. Chapter III presents a review of four current

hypotheses on the phylogeny of Sivapithecus. The fourth

chapter analyzes each of these hypotheses with respect to

its effect on a hominid-pongid divergence date. The fifth

and concluding chapter proposes a divergence of 8-10 Myr

based on an evaluation of the views discussed in the

previous sections.
 3

B. Problems with Divergence Dating.

Until twenty years ago, most research on the divergence

of hominids and pongids was conducted by primate

paleontologists and paleoanthropologists. Analyses and

interpretations were based solely upon the fossil record.

De Bonis <1983:628) summarized two general problems which

paleoanthropologists deal with in the study of the Miocene.

For this time period <between 22 and 8

M.y.a.) paleoanthropologists face two types
of problems. In the first place, it is
necessary to search for the ancestral forms,
or more precisely for the sister groups which
mark the history and phylogeny of the various
lineages starting backward from the modern
groups. Second, it is important to establish
the most probable date for the divergence
between the groups being considered, or, at
least the time period from which one first
notes that such groups are separated.

Two aspects of dating techniques and their problems,

will be discussed: <1) A description of dating techniques

along with some of their inherent problems and <2) the

underestimation of divergence dates by these techniques.

1. Dating Techniques.

A variety of techniques are used to date events in

paleontology as well as in historical archeology. Fossi 1 s

from the Middle and Late Miocene are dated by associating

them with the dates derived from materials in the

surrounding deposits. Relative dating of the surrounding

geology, flora and fauna provides dates for the fossil. In

contrast, chronometric or absolute dating of the surrounding

deposits provides a numerical age for the source deposit.

Of the variety of dating techniques currently in use

and/or being developed, several will not be discussed in

this thesis. Radiocarbon, obsidian hydration, amino acid

racemization, electro spin resonance, and dendrochronology

are only reliable for dating recent events. Obsidian

hydration and radiocarbon may, under favorable conditions,

have an upper limit of 75,000 years before present (b.p.).

Thermoluminescence <TL) and electron spin resonance are

limited to pottery and hearth stones <Curtis 1981).

Recently, Vogel <1985) has applied TL to sediments in South

Africa. The dating range for TL seems to be only a few

hundred thousand years <Vogel 1985:190). However, the

techniques that will be discussed, including potassium-argon

<40Ar/40K and 40Ar/39Ar) ratios, paleomagnetic reversal,

fission-track, and faunal correlation, are more widely used

for dating the Miocene.

a. Potassium-argon Techniques

Fossils may be dated in the geological context of the

area in which they were deposited. Since 40Ar/40K (K/Ar)

and 40Ar/39Ar techniques can accurately date the time of

deposition of volcanic material, it is fortunate that many

Miocene and Plio-Pleistocene fossils are found in

association with such material. Many of the East African

fossil bearing localities such as Hadar, Ethiopia, and Lake

Turkana, Kenya, have been dated by the potassium-argon

techniques.

Potassium-argon dating provides the most reliable

chronometric dates older than 500,000 b.p. The

 5

potassium-argon technique measures the amount of argon gas

trapped in many types of potassium-bearing rock. Potassium

decays at a regular, measurable rate. At the same time.

argon gas accumulates as the potassium decays. Under

control led circumstances, a rock containing argon gas can be

dated using the ratio of potassium 40 to argon 40 <Miller

1972).

The 40Ar/39Ar or step-heating variation of

potassium-argon dating was independently developed in 1962

and 1965 <Curtis 1975) and has gradually replaced the

conventional K/Ar method because of its greater accuracy.

40Ar/39Ar measures the potassium and argon gases at the same

time and location of the sample while K/Ar measures these

gases at different loci and stages. The step-heating

40Ar/39Ar method can yield "extremely accurate multi-point

potassium-argon isochron ages and can prove the presence or

absence of excess argon or argon loss discrepancies" <Fitch

1972:85). The presence of excess 40Ar is a primary source

of error in K/Ar dating. The disadvantage of the

step-heating method is that it is very time consuming and

difficult to use. requiring between six to twenty-four

heating steps and argon ratio calculations.

Each of the different dating techniques has its own

particular set of assumptions which create a margin for

error. A potassium-argon date reflects the number of years

since a rock began accumulating argon gas, not necessarily

the time of fossil deposition. Deposits formed by the

 6

products of volcanic eruptions provide an excellent

reference point for potassium-argon dating. Any

potassium-bearing rock involved will lose its argon gas in

the eruption and wil 1 begin accumulating it anew after

deposition. However, since the half-life of potassium-argon

decay is so long, approximately 1.31 billion years, it

cannot be used to date recent deposits accurately <Metcalf

1982:80). The lower limit of accuracy is 10,000- 75,000

b.p. under exceptionally favorable conditions <Curtis 1981).

There is no upper limit applied to potassium-argon dating

techniques <Dalrymple and Lanphere 1969:45).

Fossil dates that ar~ obtained using potassium-argon

<as well as paleomagnetic reversal or fission-track tests)

can be misleading or in error. Due to a number of factors,

there are no guarantees that the fossil was deposited at the

same time as the surrounding geological material. Nor is it

certain that the organism even lived at the same time of the

geologic deposition or that it died where it was found.

Further, if the argon-bearing rock is contaminated, the

results of testing wi II not be accurate. Sometimes the

contamination wil 1 go undetected and cause dating errors

leading to invalid hypotheses affecting dating and

phy 1 ogenet i c ana 1 yses. :

An example of misconceptions arising from incorrect

dating was the estimation of the geologic age of the KBS

tuff at Lake Turkana, Kenya. The tuff overlaid the famous

KNM-ER 1470 skull and is an important dating horizon. It

 (7\
\ '
~ '

•.ns sampled and dated at least 60 times using K/'Ar and

40Ar/39Ar techniques <Curtis 1975,1981). The contamination

eventually discovered in samples from the KBS tuff ignited a

lengthy controversy over dating discrepancies between Lake

Turkana hominids and those from Hadar, Ethiopia. This

controversy lasted years and is one of the most notable in

East African paleoanthropology <Curtis 1975:205; Curtis

1981:14; Johanson and Edey 1981:170-171).

It is not always possible to use the potassium-argon

techniques to date fossil sites. For example, at

Haritalyangar, India, there are no dateable volcanic ashes

(Johnson et gl. 1983:237). There are very few accurate

potassium-argon dates for the Siwalik Hi! Is of Northern

India and Pakistan <Curtis 1981:12). The few K/Ar dates

aval !able are used to calibrate paleomagnetic estimates.

Using K/Ar techniques, dates as old as 17.2 Myr have

been produced <Leakey and Walker 1985; McDougal I and Watkins

1985) at Buluk. Kenya. These dates provide the earliest

dates for Sivapithecus when Kenyapithecus is submerged

within it <Curtis 1981).

b. Paleomagnetic Reversals

Paleomagnetic reversal methodology, paleomagnetism or

magnetic stratigraphy is a non-radioactive technique of

chronometric dating. As the Earth/s magnetic field

periodically changes, the history of each reversal has been

recorded with considerable accuracy for the last 4-5 Myr.

This time scale, known as the calibrated magnetic polarity

 8

time scale <MPTS), is applied to volcanic and sedimentary

material alike as long as the compounds bear iron <Wolpoff

1980). The MPTS is constantly undergoing revision as it is

recalibrated utilizing recent data and new reversals are

discovered.

Five assumptions must be made to utilize the

paleomagnetic method: (1) "An estimate of the range of

ages that the reaction should encompass" <Tauxe 1979:401);

(2) "For a given lithofacies, when averaged over a large

stratigraphic thickness, is linearly proportional to time"

<Tauxe 1979:401); (3) "The section is densely sampled

enough to assume that it approximates field behaviour"

<Tauxe 1979:401); (4) "The magnetic remanence of the rock

was acquired shortly after deposition and represents a

record of the magnetic field at the time of formation of the

rock" <Tauxe 1979:401); and C5) independent calibration by

radiometric dating of at least one point, but preferably

more, in the local stratigraphy <Curtis 1981:9).

Paleomagnetic dating in the Middle and Upper Siwaliks

is uncertain at earlier than 5 Myr because the time scale is

not as clearly defined as it is after 5 Myr. There are not

as many opportunities to calibrate the magnetic stratigraphy

radiometrically in the Siwaliks as in East Africa. In

addition, the lithics of the Siwaliks have been subjected to

prolonged "diagenetic alterations <such as red pigment

formation) which often obscures the original magnetisation

of the rock" <Tauxe 1979:401).

 9

A narrow stratigraphic band which has produced the

majority of hominoid fossils from the Khaur region of the

Middle Siwalik has been magnetical Iy dated at 8 Myr <Tauxe

1979). Paleomagnetic dates from Haritalyangar revised the

latest appearance of Sivapithecus in Asia to approximately 7

Myr <Johnson~ gl. 1983) and 5.5 Myr <Sankhyan 1985).

c. Fission-track

Fission-track dating is an absolute method for dating

volcanic glass or crystal. It has a range of 100-10,000,000

years b.p. Spontaneous fission of uranium 238 during the

"lifetime" of glassine material leaves measurable tracks.

The li~etime of a crystalline form is defined as the period

of time since it was last heated. The date of the substance

is defined by the number and density of the tracks

<Fleischer 1975; Fleischer and Hart 1972). A major source

of error is track destruction due to excessive heat. Where

this has occurred, the age estimates wil I be low <Fleischer

and Hart 1972).

d. Faunal Correlation

Faunal correlation or biostratigraphy is an extremely

useful method for relative dating. Primate fossils,

specifically those of hominids and pongids, are very rare

compared to other fossi 1 species such as suids (pigs) and

murids <mice).

Recording faunal changes at various locales can link

geologic events with similar stratigraphy throughout the

world <Opdyke et gl. 1979). However, as Opdyke~ gl.

 10

<1979:29) state, "these faunal-stratigraphic divisions are

fairly reliable for intercontinental correlation but are

unreliable for correlation within the Siwaliks". Because

widespread correlation in the Siwaliks is unreliable, faunal

assemblages have been calibrated independently by the

geomagnetic time scale.

Under ideal situations, where faunal correlation can

link two geographically distant sites, one can be used to

relatively date the other. When a site is wei 1 calibrated

by chronometric dating techniques, it can date another

uncalibrated site relatively through a faunal link.

Al 1 of these dating techniques have one thing ~n

common: they areal 1 built upon assumptions that concern

their parameters. These parameters are measurable and

consist of variable values. The accuracy of the dating

estimate is limited by the number and the extent to which

the assumptions are met in the use of the method <Curtis

1981:7).

Since the accuracy of any particular dating method is

dependent upon the fulfillment of its basic assumptions, it

is important to validate the results. Paral lei tests by

another method should be made to minimize and identify any

possible errors. When a date is validated by two or more

techniques, it indicates their parameters have been met. If

a date cannot be confirmed by any of the other dating

methods, its accuracy is low <Curtis 1981:11).

2. Underestimation of Divergence Dates.

 11

It cannot be assumed that the estimated dates available

at any one time of known fossils represent a completely

accurate span of time for a specific fossil species or

genus. This uncertainty leads to a source of error in

estimating phylogenetic divergence: underestimation of

divergence dates. Underestimating the divergence date of a

primate fossil species is the result of two basic problems:

(1) the usual small size of the fossil sample <2) gaps in

the fossil record, and <3) distinguishing two diverging

species in the fossil record.

The sample size refers to the number of fossils found

at any specific location. Related to this is the common

problem of the extent to which a sample of hominoid fossils

is representative of the original species. These

assemblages are rare compared to those of other species such

as suids and bovids. At any one site, such as Kanapoi and

Lothagam in Kenya, there may be only one hominoid fossil.

As an extreme example, there are no hominid-like hominoid or

pongid fossil assemblages at alI between 5.5 and 3.59 Myr

<Boaz 1983). Due to the low frequency ·of fossils between

4-8 Myr, it is very difficult to const~uct any testable or

plausible hominoid phylogenies.

The second problem is the difficulty of distinguishing

between two diverging and developing members of a species in

the fossil record until after they undergo speciation.

After speciation, it is not guaranteed that the species wil 1

be easy to identify. For example, sibling species are

 12

difficult to distiriguish morphologically in the fossil

record. Fossil groupings are not viewed as a continuum but

as separate lineages <Walker 1976). For example, fossils

are designated by genus and species groupings. Group

relationships are regarded as ancestral to and descended

from each other with sharp parameters separating them. The

low frequency of fossils makes this especially true In

hominoid lineages. Pilbeam <1984b:20) also comments upon

this problem.

At the moment, even with the very best of the

Plio-Pleistocene data, we can do no more than
try to view both ancestor and descendant as
blurred biological realities; we must
restrain ourselves from too many flights of
fancy in discussing the transformation of one
into the other.

The dating estimate of a phylogenetic branching point

must be deduced from the dates of fossils preceding and

succeeding the speciation event. As in the case of hominids

and pongids, there is a great deal of debate concerning

their last common ancestor.

The estimated dates for divergence are likely to be

greatly underestimated compared to the actual date of

speciation. Anatomical changes are not immediately seen in

the fossil record after a divergence. Cronin (1983)

estimates a period of 1-2 Myr passed before changes were

recognized between the hominlds and pongids. The following

statement by Walker <1976:70) sums up much of the

underestimation of divergance dating quite well:

When a paleontologist is asked to estimate a

divergence date for hominids and pongids, is
 13

he really being asked to say when, once upon

a time, a pongid mother gave birth to a
hominid infant? If it is true that hominids
developed from pongids, then the first
hominids will be very pongidlike, and how to
categorize intermediate species within higher
categories becomes a matter of resolving the
conceptual conflict between horizontal and
vertical classifications. Al 1 we can hope to
determine is the time of the first record of
the species that show unequivocal hominid
characteristics. This will not be the
splitting time of course, but a much later
event. If it could be demonstrated further
from which pongid species these early
hominids evolved, it would stil 1 remain to
establish from which ancestral species the
living pongids were descended in order to
deduce the last common ancestor and hence a
divergence date for modern hominids and
pongids.

Underestimation must be reflected in the estimated

divergence date of the hominids and pongids. At least 2

million years should be added to the date of the earliest

known hominid fossils. In other words, it the earliest

Sivapithecus is dated at 17.2 Myr <Walker and Leakey 1985;

McDougal 1 and Watkins 1985), then its divergence would be at

least 19.2 Myr. Likewise, it the earliest date for

Australopithecus is 3.6 Myr from Hadar, Ethiopia <Walter and

Aronson 1982) or 5.5 Myr for Lothagam <Patterson ~ gl.

1970) and Sivapithecus is the last common ancestor to the

australopithecines and pongids, then the hominid divergence

could be as late as 5.6 Myr or as early as 7.5 Myr.

 14

C. Phylogenetic Analysis.

Paleoanthropological arguments often center around the

phylogenetic interpretations of fossil forms. Even after

some of the problems associated with the dating techniques

are considered, the dates of specific fossils are not

usually questioned unless they are vitallY important to a

phylogenetic hypothesis. An example of a technical dating

controversy is the potassium-argon dating of the KBS tuff at

Lake Turkana, Kenya, previously introduced <Section B of

this chapter); the date is vital to analyses of the

positions of Homo habilis and Australopithecus afarensis in

hominid phylogeny. ~ afarensis is considered older and

more primitive than~ habilis. While~ afarensis has not

been found at Lake Turkana, finds at Hadar, Ethiopia, have

been dated at 2.6 Myr <Aronson et £1. 1977). These dates

did not agree with the originally published dates of 2.61

Myr <Fitch and Miller 1976) for the KBS Tuff. Revised dates

of 2.9-3.6 Myr for hominid-bearing locales at Hadar have

been published <Walter and Aronson 1982). The KBS Tuff also

has recently been revised to 1.88 ± 0.02 Myr <Brown et ~.

1985). The new dates for Hadar and Lake Turkana make the

hominid material more compatible.

A controversy revolves around the hominid status of

Sivapithecus. Whether Sivapithecus and/or Ramapithecus is

accorded hominid versus pre-hominid status indicates a

"late" or "early" hominid-pongid divergence estimate,

respectively. If Sivapithecus and/or Ramapithecus is

 15

classified as hominid, then the hominid-pongid divergence

may have occurred as early as 17.2 Myr based upon the Buluk,

Kenya, material <Leakey and Walker 1985; McDougall and

Watkins 1985). If it is not a hominid, but only a hominoid,

then the date may be as late as 6-7 Myr <Greenfield

1980:361; Pilbeam 1980:279), 5.5 Myr based on Sankhyan

<1985), or 5-10 Myr <Pilbeam 1986). The phylogenetic status

of Sivapithecus and Ramapithecus is critical to estimation

of a divergence date; also, it is important in determining

whether Sivapithecus and Ramapithecus represent the same

genus: Sivapithecus.

Another important issue is the inconsistent use of key

terms in paleoanthropology. "Sivapithecine" and

"ramaplthecine" refer to those fossil species that exhibit

general Sivapithecus-like and Ramapithecus-like traits,

respective 1 y. "Ramamorph" is a convenient term to genera 1 1 y

summarize the geographical, temporal, and morphological

limits of sivapithecine / ramapithecines <Ward and Pilbeam

1983). Assuming that the slvapithecines include both

lineages, it is apparent that Sivapithecus diversified into

many species. This very complicated issue will be discussed

at length later <Chapters III and IV).

The definition of the term hominid requires

clarification. Technically, a hominid is any genus or

species member of the family Hominidae. In order to analyze

a trend towards hominization in a fossil species, the term

must first be understood. Hominization is the development

 16

of characteristics toward or in the direction of a

"human-like" level. These'£iaits are either exclusively

characteristic of hominids compared to pongids, or show an

evolutionary trend toward a typical human trait. The

features compared between Miocene hominoids and later

hominids are primarily dental and mandibular, because

post-cranial remains are rare. The presence of any one

hominid-like characteristic may not support classifying a

fossil as a hominid, but the greater their number, the more

likely it is that the fossil is a hominid. Recently,

Pilbeam <1986), has questioned the validity of some of these

traits for defining hominid status. For example, thick

enamel may be a primitive characteristic and thin enamel, as

seen in the chimpanzee, is derived. Ramapithecus and

Australopithecus have often been linked together because of

thick tooth enamel. Para! lei development of hominid traits

may also account for difficulties in their determination.

If Australopithecus and the chimpanzee are closer to each

other than to Ramapithecus, then thick enamel is

para! lellsm. The following is a list of typical traits used

to measure the level of hominization in a fossil hominioid:

1. Rare occurance of diastemata <Greenfield

1980; Wolpoff 1980).

2. Little or no inter·locking of the canines

<Greenfield 1980).

3. Small incisors compared to the extant apes

<Wolpoff 1980; Kay and Simons 1983).

4. Lower crowned cheek teeth < Greenfield 1980;

Kay and Simons 1983).
 17

5. Low pe.rcen tage of pr;ogna th ism (Kay and

Simons 1983).

6. Multi-cusped and round third lower premolar,

with a morphology suited for: grinding and
crushing <Greenfield 1980; Kay and Simons
1983).

7. Very thick occlusal enamel on cheek teeth

<Greenfield 1980; Kay and Simons 1983).
This view is not accepted by Pilbeam (1986).

8. Rectangular morphology of the cheek teeth

<Greenfield 1980).

9. Reduced sexual dimorphism of the canines

compared to Dryopithecus, Pan, Pengo, and
Goril Ia <Greenfield 1980; Kay and Simons
1983).

10. Dental arcade curved, sometimes anteriorly

square with the general shape short and
broad <Greenfield 1980).

The nomenclature for Miocene hominoids has changed

several times since the 1920/s and 1930/s. Sivapithecus was

first used by Pilgrim <1927) to designate a late Miocene

hominid-like hominoid from the Siwalik Hil Is. Ramapithecus

is now included in the description of Sivapithecus <Kay and

Simons 1983:601-605; Greenfield 1980:353). Unless otherwise

stated, Ramapithecus will be referred to as an independent

genus, as was done in the literature during the 1960/s when

Sivapithecus and Ramapithecus were considered separate

genera.

,~
 Chapter II

The Sivapithecus Problem

Estimates for the timing of the hominid-pongid

divergence are directly related to interpretations of the

phylogenetic status of Sivapithecus. If Sivapithecus is

considered a hominid ancestral to the australopithecines and

Homo, then, based upon the established dates, the

hominid-pongid divergence is considered to be early. The

dates associated with sivapithecine fossils as a whole are

consistent and reliable. There is a broad range of 8-14 Myr

accepted by several leading researchers <Curtis 1981; Lipson

and Pilbeam 1982; Kay and Simons 1983). Boaz <1983)

supports a time range of 8-10 Myr when Kenyapithecus is

excluded from the sivapithecines. The fossils of certain

locales, such as Fort Ternan, Kenya <Simons 1969), are dated

earlier than Indian forms. A later date of 5.5 Myr for

Asian and African Sivaplthecus is proposed by Sankhyan

<1985) from the Siwaliks, increasing the range to 5.5-17.2

Myr when Kenyapithecus is included. AI lowing for

underestimation, the split could be as early as 17 Myr as

opposed to approximately 19 Myr without underestimation.

However, if these fossils are classified as pre-hominid,

then the estimated dates of divergence may be as late as 5-7

Myr <Cronin 1983:130). Cronin <1983) has a built-in 1-2

mil lion-year underestimation value in his proposal.

The differences between the estimates may be reduced if

only one species of Sivapithecus is postulated as ancestral

18
 19

to both hominids and pongids. However, a particular species

has not yet been identified as this ancestor. If a specific

sivapithecine species is determined to be ancestral to the

hominids and/or pongids, Sivapithecus as a genus would be

classified as a pre-hominid.

The Sivapithecus problem has been discussed

periodically for the last 50-60 years. In the historical

analysis that follows, the classification of Sivapithecus is

shown to have exhibited several trends.

In the first trend, hominoid phylogenies did not

specify the position of Sivapithecus. It was considered

either to be a hominid or pre-hominid. However, given that

its phylogenetic position was generalized, Sivapithecus and

other Miocene hominoids were classified into several

different genera and species. Sivapithecus was later given

a specific position within the phylogeny and its

diversification into several species was reduced and

simplified. Phylogenetic ties were drawn between the

ramapithecines and specific fossil hominoid forms. Current

trends address the overal 1 role of the sivapithecines in

hominoid ancestry, while at the same time Sivapithecus was

again split taxonomically into several distinct species.

These identified species are not necessarily linked with any

other hominoid fossil group either as an ancestor or a

descendant.
 20

A. 1 930 to 1 960 .

There have been two general trends or advances in

hominoid fossil analyses since the 1930?s and 1940/s. (1)

Taxonomic thinking and phylogenies have become more complex

and sophisticated. Inconclusive systematics have evolved

into elaborate phenetic and cladistic constructs. (2)

Researchers began to relate the form and function of extant

primates to fossil groups. The following analysis of

pre-1960 research ls a brief description of paleontological

hypotheses with specific examples.

The indistinct classification of the 192G/s and 1930/s

is reflected in specific studies. Pilgrim <1927) placed

Sivapithecus as central to the ancestry of hominids and

ponglds, not classifying it as either one, but placing it

well before the hominid-pongid divergence. Lewis <1934)

proposed that Ramapithecus was very close to the Hominidae.

He later placed Ramapithecus as an intermediate form between

the Pongldae and Hominidae <Lewis 1937). Hrdlicka <1935)

refuted hominid similarities of Ramapithecus and classified

it as a small ape. He claimed Lewis (1934) studied two

specimens that were too defective and fragmentary to justify

a .new genus.

Between the mid-1940/s and early 1950/s, very little

was done in the area of Miocene primate paleontology. World

War II and the emergence of australopithecine studies were

of greater interest <Fleagle and Jungers 1982:201).

A notable exception was the work of Le Gras Clark and

 21

Leakey <1951). They reported new discoveries of Proconsul

and a then new, unidentified fossil hominoid. This new

fossil was significantly distinct from Proconsul and showed

similarities to~ sivalensis. It was also different from

Dryopithecus. The small maxi! lary fragment was first

classified as.§..:.. africanus <Le Gras Clark and Leakey 1951)

and later revised as Kenyapithecus africanus <Leakey 1967).

In an essay, Heberer (1959) did not accept hominid

status for Ramapithecus. In fact, he labeled Ramapithecus

as a dryopithecine and considered the dental similarities

between Ramapithecus and hominlds to be the result of

parallel evolution <Heberer 1959:225). He estimated the

divergence to be no later than the Middle Miocene.

If the molars of Ramapithecus have an

appearance more like hominids than is the
case in other dryopithecines, this is more an
exmple of parallel evolution than a sign of
direct ancestry. This does not involve an
entire pattern of traits, and thus cannot be
used as a phylogenetic guidepost <Heberer
1959: 225).

These early studies analyzed the dental characteristics

of Ramapithecus and Sivapithecus, because, until at least

the late 1960's, there were few, if any, cranial or

post-cranial remains known <Prasad 1969). At the time of

their studies, the specimens primarily consisted of a few

teeth and partial mandibles. The same data W;ere interpreted

and re-interpreted several ways. Sivapithecus was used as a

-support for the Asian origin for man. The popular hyothesis

then was that man originated in Asia and Sivapithecus was

examined closely as a human ancestor because of its

 22

discoveLy in NortheLn India and China.

 23

B. The 1960/s.

By the early 1960/s, paleontological analyses followed

three general trends: (1) Direct ties were proposed between

specific Miocene hominoids and extant species. The fossil

species were considered prototypes of the modern hominoids

<Pilbeam 1979:340). (2) There were implicit expectations of

what the fossil record should reveal based on ideas derived

from studies on the living primates <Pilbeam 1979:341). (3)

The total sample of fossil hominoids included several

hundred specimens, the bulk of them from Kenya, for which a

large number of generic and specific names had been coined

<Pilbeam 1979:337) and there was a great deal of concern

with their phylogenetic classifications.

These trends resulted in an overdiversification of

fossil phylogenies. By 1965, there were at least

twenty-eight genera <Simons and Pilbeam 1965) classified for

Miocene hominoids. According to Simons <1964:529), there

may have been four dozen taxonomies for potential

dryopithecines. The phylogeny of Miocene hominoids had

become confused and unwieldly. A common belief was that the

dryopithecines had undergone a very extensive adaptive

radiation <Pilbeam 1966, 1968). Simons <1963:886) claimed

that the extensive diversification of Dryopithecus and

Sivapithecus represented more species than there were in

reality. The number of genera should have been reduced to 3

or 4 <Prasad 1969:299), perhaps even fewer <Simons

1963:886).
 24

Due to the proliferation of australopithecine finds in

East Africa, the location of hominid origins was moved

westward. As a result, Asia was no longer considered the

birthplace of man and Sivapithecus was largely ignored as a

hominid or even a hominoid ancestor.

Ramapithecus continued to be classified as a genus

separate f~om the dryopithecines <Simons 1963:887). Prasad

<1964) also separated Sivapithecus from Ramapithecus and

Dryopithecus. Ramapithecus was considered slightly closer

morphologically to the hominids than to the pongids <Simons

1963:887), while Prasad (1964) grouped Ramapithecus,

Sivapithecus, and Dryopithecus in the Pongidae. Simons

<1963) stressed the transitional tendencies of Ramapithecus

that made it difficult to classify. And Simons <1964:535)

concluded that Ramapithecus was definitely an ancestor to

Homo. On the other hand, Leakey (1963:48) described

Ramapithecus, ~ wickeri, Dryopithecus, and Sivapithecus as

incertae sedis. He felt that Ramapithecus and Kenyapithecus

would eventually have to be placed in the Hominidae.

Ramapithecus and sometimes Kenyapithecus <Leakey 1963)

were stilI thought of as being generally related to the

hominids <Simons 1963). When some hominization trends were

considered in combination, Raroapithecus appeared to fulfill

the criteria of a hominid <Simons 1963:886). Singularly,

however, each trait could be found among the pongids.

Indeed, one gets the very strong impression

that· during this period primate
paleontologists -- such as Simons, Leakey,
Pilbeam, and others-- were not really
 25

involved with the issues that had occupied

primate phylogeny during the first half of
the century. Questions about the exact
phyletic relationship between humans and
living ape genera or the extent to which the
hominid lineage had passed through a
brachiating stage were not being posed.
Rather, these researchers/ primary task was
to place the newly found fossils in their
proper phyletic and systematic positions
<Fleagle and Jungers 1982:209).

In 1965, Simons and Pi 1beam pub 1 i shed a 1 andmark P.aper

with the intention of clarifying and reorganizing the

taxonomies and descriptions of the Miocene hominoids. Their

reasons for doing so were: (1) the fossil record was

primarily limited to maxillary and mandibular fragments, as

opposed to post-cranial remains; (2) specimens had been

discovered at many widely separate locales <China to India

to East Africa); (3) descriptions for these specimens were

widely scattered temporally as far back as 1856; and (4)

there was a tendency towards taxonomic splitting of new

fossils <Simons and Pilbeam 1965:81). They felt the new

specimens were not being analyzed thoroughly enough compared

to previously known species to justify new classifications.

Simons and Pilbeam <1965) considered their work as a type of

progress report .that was flexible enough to welcome

progressive change.

Simons and Pilbeam <1965) reduced the number of Miocene

hominoid taxa from fourteen to three genera and from

thirty-one to nine species. The fossils were divided into

two families: the Hominidae and Pongidae. The Pongidae

contained Dryopithecus and Gigantopithecus while the

 26

Hominidae was represented by Ramapithecus only.

Dryopithecus contained seven species <see below), while

Gigantopithecus and Ramapithecus each remained monotypic.

They concluded that African apes and Homo were

phylogenetically closer to each other than either was to the

orangutan. Pengo may have split off earlier from the

dryopithecines, but at that time the evidence was

inconclusive <Simons and Pilbeam 1965). Table I is a

complete list from Simons and Pilbeam <1965:142-143) that

outlines the variety of taxonomic names that were reduced

and reclassified in their proposal.

Only two specimens showed any orangutan-like features.

Simons and Pilbeam <1965) reported a canine described by

Lydekker <1886) as indistinguishable from Pengo. In their

reorganization it was redefined first as~ indicus <Simons

and Pilbeam 1965:87), but was later reclassified as~

indicus <Simons and Pilbeam 1965:114). Pilgrim <1915)

defined the second fossil, a molar, as "Paleosimia

rugosidens" and related it to Pengo. Later, Lewis C1937>

reclassified it as~ sivalensis while Simons and Pilbeam

<1965:142) redefined it as~ sivalensis. Both of these

fossils were at one time classified as Sivapithecus but were

changed when Sivapithecus was submerged into Dryopithecus.

Simons and Pilbeam <1965) proposed a four-part

phylogeny associating specific Miocene hominoids with

subsequent hominid or ape genera <Fig. I>.

1. ~major is ancestral to Gorilla.

 27

2. ~ africanus is ancestral to Pan.

3. ~ sivalensis may be ancestral to Pengo.

4. Ramapithecus is a hominid leading to the

australopithecines.

The trends of the 1960/s were illustrated by Simons and

Pllbeam <1965). They proposed direct phylogenetic ties

between three species of Dryopithecus and the extant Great

Apes, and Ramapithecus was associated with Homo. These

fossil species were chosen because they were morphologically

closer to what the ancestors of hominids and pongids were

expected to be. The paper emphasized the restructuring of

the taxonomic classifications.

From the 1930/s through the 1960/s, a growing consensus

among paleoanthropologists accepted a pre-australopithecine

hominid phase in hominid phylogeny. Simons and Pilbeam

<1965) and Pilbeam <1966) proposed a loose connection

between Ramapithecus and the australopithecines, even though

they accepted Ramapithecus as a hominid. Based upon its

dental development, Le Gras Clark <1964) placed Ramapithecus

in a transitional phase between the Miocene apes and the

hominids.
Simons <1967:35) proposed a tentative phylogeny that

supported a solid tie between Ramapithecus and~ africanus.

This phylogeny indicated a divergence of dryopithecines that

eventually lead to distinct hominid and pongld lineages to

have occurred greater than 25 Myr with Aegyptopithecus

zeuxis as the last common ancestor of the pre-hominid and

pre-pongid lineages <Fig. II). The Pengo lineage is ignored

 28

Table I
Revision of the Dryopithecinae
<Simons and Pilbeam 1965:142-143)

Prior Terms Terminology

of this paper
1856 - Dryopithecus fontani LARTET; .IL.. fontani
mandible
1879 - Palaeopithecus* sivalensis 1L... sivalensis
LYDEKKER; palate.
1894- Anthropodus rouvil lei DE LAPOUGE; Nomen dubium
tooth
1895 - Paidopithex* rhenaus PORLIG; femur .IL.. fontani?
1895- Pliohylobates eppelsheimensis .IL.. fontani?
DUBOIS; above femur.
1901 - Dryopithecus rhenanus SCHLOSSER; .IL.. fontani
teeth
1902- Dryopithecus darwini ABEL; teeth .IL.. fontani
1902 - Neopithecus* brancoi PI i op i thecus?
<SCHLOSSER) ABEL; teeth. brancoi
1902- Griphopithecus* suessi* ABEL; tooth Nomen dubium
1905- Dryopithecus suevicus KOKEN; tooth Nomen dubium
1910 - Sivapithecus indicus PILGRIM; tooth .IL.. indicus
1910 - Dryopithecus punjabicus PILGRIM; R.:.. pun.iabicus
mandibular fragments.
1915- Dryopithecus chinjiensis PILGRIM; .IL.. sivalensis
tooth
1915 - Dryopithecus giganteus PILGRIM; 1h. indicus
tooth
1915 - Palaeosimia rugosidens* PILGRIM; .IL.. sivalensis
tooth
1919 - Dryopithecus germanicus ABEL; teeth .IL.. fontani
1920 - Dryopithecus? mogharensis FOURTAU; Prohylobates
mandible. tandy i <monkey)
1924- Dryopithecus pilgrimi* BROWN, .IL.. sivalensis
GREGORY and HELLMAN; mandible.
1924 - Dryopithecus cautlevi* BROWN, 1L... sivalensis
GREGORY and HELLMAN; mandible.
1924- Dryopithecus? frickae* BROWN, 1L... indlcus
GREGORY and HELLMAN; mandible.
1927 - Slvapithecus himalayensis* PILGRIM; 1L... indlcus
mandible.
1927- Sivapithecus orientalis* PILGRIM; 1L... indicus
maxi 1 I a.
1927- Sivapithecus middlemissi* PILGRIM; 1L... indicus
mandible.
1927- Palaeopithecus? sylvaticus* .IL.. sivalensis
PILGRIM; mandible.
1927- Hylopithecus hysudricus PILGRIM; Nomen dublum
tooth
 29

Table I <continued)

1933 - Xenopithecus koruensis* HOPWOOD; .l2:.. africanus

maxi 11 a
1933 - Proconsul africanus HOPWOOD; .ll:.. ·afrlcanus
maxi 1 1a
1934 - Ramaplthecus brevlrostris LEWIS; R:.. pun.iabicus
maxi 1 1a
1934- Ramaplthecus harlensls* LEWIS; .ll:.. slvalensis
maxi 11 a
1934 - Sugrivapithecus sa1montanus LEWIS; J:L.. sivalensls
mandible.
1934 - Dryopithecus sivalensis* LEWIS; R:.. punJabicus
mandible
1934 - Bramapithecus thorpei LEWIS; R:... pun.iabicus
mandible
1934- Adaetontherium incognitum LEWIS; Nomen dubium
tooth
1935 - Semnopithecus eppelshelmensls HAUPT; Nomen dublum
tooth.
1935 - Gigantoplthecus blackl ~ blackl
von KOENIGSWALD; teeth.
1936 Sugrlvapithecus gregory! LEWIS; .ll:.. sivalensis
teeth
1937 - Proconsuloldes* nalvashae LONNBERG; Pan spec.
teeth.
1938 - Austrlacopithecus welnfurterl .ll:.. fontanl?
EHRENBERG; humerus, ulna.
1944- Sivaplthecus occidentalis VILLALTA IL.. laietanus
and CRUSAFONT; mandible.
1944- Hispanopithecus·laletanus VILLALTA IL.. laietanus
and CRUSAFONT; mandible.
1946 - "Kansuplthecus" BOHLIN; mandible Nomen nudum
1950 - Proconsul nyanzae CLARK and LEAKEY; IL.. nyanzae
maxi I 1 a.
1950 - Proconsul major CLARK and LEAKEY; IL.. maJor
mandible.
1950 - Slvapithecus africanus CLARK and J:L.. sivalensis
LEAKEY; maxilla.
1950 - Indopithecus* giganteus J:L.. indicus
von KOENIGSWALD; teeth.
1950 - Udabnopithecus garedziensis JL..? fontani
BURTSCHAK-ABRAMOVITSCH and
GABACHVILI; teeth.
1956 - Rhenopithecus eppelsheimensis von Nomen dubium
KOENIGSWALD; tooth.
1957 - Ankaraplthecus meteai OZANSOY; IL.. i ndi cus
mandible
1957- Dryopithecus kelyuanensis WOO; IL.. i ndl cus
teeth part im
IL.. slvalensis
partim
 30

Table I <continued)

1961 - DLyopithecus piveteaui CRUSAFONT Nonem nudum

and HURZELER; no type designated.
1961- Rahonapithecus sabadellensis Nomen nudum
CRUSAFONT and HURZELER; no type
designated.
1962 - Sivapithecus aiyengaLi PRASAD; ~ indicus
mandible
1962- Kenyapithecus wickeLi* LEAKEY; & punjabicus
maxi 11 a
 31

Figure I

Phylogeny of Large-Bodied Miocene Hominoids

Simons and Pilbeam <1965)

M 0-
Pleistocene I Gar i 1 1 a Pan Homo
L AustraloQithecus
L 5-
I
0
N 10-
s
0 15- R:.. Qun,j ab i cus
F
JL.. ajor
y 20- JL.. sivalensis I
-II
D. africanus
E
A
R 25-
s \
 32

Figure II
Phylogeny that Supports a Relationship
Between Ramapithecus and Australooithecus
Simons (1967)

Pleistocene
M

L
L
r
o-

5-
-,
Homo
lb.. africanus
Gor i l l a

PI i ocene I
0
N 10-
Miocene S R.:.. pun.iabicus
0 15-
F

y 20- undetermined ~ africanus ~rna or

E dryopith~
A
R 25-
S
Aegyptopithecus zeuxis
 33

in the following schematic (Simons 1967:35).

1. Aegyptopithecus zeuxis is ancestral to:

a. An as yet unidentified dryopithecine.

b. The common ancestor to Dryopithecus

africanus and 12.:.. ma.ior.

2. The unidentified dryopithecine is ancestral

to & pun.iabicus.

3. 12.:.. africanus is ancestral to Pan.

4. 12.:.. ma.ior is ancestral to Goril Ia.
5. & pun.iabicus is ancestral to ~ africanus.

6. A.:.. africanus is ancestral to Homo.

Reynolds (1966:442) also proposed an early separation

date for hominids and pongids during the late Oligocene and

early Miocene based on other taxonomic studies. This

speciation event would have occurred before the three great

apes became distinct. Reynolds (1966) did not address

Sivapithecus in his hypothesis. Ramapithecus was introduced

as a proto-hominid that spread out from Africa to Europe and

Asia. This radiation was the result of the transition

period between the hominid and pongid lineages during the

Miocene <Reynolds 1966:447). In a similar view, Pilbeam

(1966, 1968:1337) regarded Kenyaplthecus as synonymous with

Rampithecus.

On the other hand, Leakey (1963) separated

Kenyapithecus and Ramaoithecus, and tentatively placed them

both in the Hominidea. By 1967, Leakey had concluded that

they definitely were hominids, but still retained their

independence from each other <Leakey 1967). Leakey (1967)

 34

revised the classification of ~ africanus to~ africanus

and stated that the range of variability between

Kenyapithecus and Ramapithecus would result in a very

liberal grouping if they were combined. Zwell <1972)

refuted Leakey/s claim that Kenyapithecus was a hominid.

Due to the shape of the tooth row, he claimed that the ~

africanus specimen <KNM-RU 2087) should have been classified

as lL., nyanzae. In Zwell "s <1972) opinion, ~ africanus did

not possess any of the distinguishing morphological

characteristics of hominids.

Pilbeam <1968) viewed both Dryopithecus and

Sivapithecus as dryopithecines separated by locality. He

defined the European and Asian forms as Dryopithecus and

Sivapithecus, respectively. Sivapithecus was hypothesized

as ancestral to Pengo while Ramapithecus, separate from

Sivapithecus, was linked to the hominid lineage. As stated

by Pilbeam (1968:1336):

Some of the dryopithecines from deposits in

the Siwalik Hills, north-west India, are, in
alI probability, ancestral to the only living
Asiatic pongid, Pengo pygmaeus, the
orang:-utan.

In short, most of the features.of hominid

dentitions are found in Ramapithecus and
there are enough simi 1 ar it i es between
Ramap i thecus and PIe i stocene .homi n ids to
Justify the view that they ar~ part of one
lineage or of closely related :lineages.

In opposition to Pilbeam (1968), Prasad <1969)

described three genera of dryopithecines; Dryopithecus,

Proconsul, and Sivapithecus. He defined eight species among

three genera: two in Dryopithecus (~ fontani and~

 35

laletanus), three in Proconsul <£.:.. africanus, f.:_ nyanzae,

and ~ ma.i or), and three in S i vap i thecus <~ i ndi cus, ~

si•Jalensis, and~ chinj iensis).

Prasad <1~69) retained Proconsul as a genus because the

size ratios of several traits did not conform to the

acceptable range of variation for either Dryopithecus or

Sivapithecus. The large size of the third molar, the

emplacement of the canine, and relative straightness in the

profile of the symphysis compared to Dryopithecus and

Si vap i thecus are used to support Proconsu 1 ma.i or as a genus.

By 1969, Ramapithecus continued to be defined as a

hominid <Simons 1969; Prasad 1969). It was dated at 14 Myr

from Fort Ternan, Kenya, by potassium-argon techniques

leading Simons <1969) to estimate the hominid-pongid

divergence to be prior to 20 Myr. This 5 million-year

revision from his 1967 estimate still places the split

~.Jithin the dryopithecine time span. Pilbeam <1968:1338)

concluded that the hominids and pongids were recognizably

distinct by 14 Myr.

Simons <1969) believed that a divergence later than 20

Myr would eliminate Ramapithecus from the hominid phylogeny.

A late divergence would require any shared characteristics

with the australopithecines to be a result of para! lei

evolution, not shared derived characteristics resulting from

an ancestor-descendant relationship. Simons (1969:328)

outlined six shared characteristics between Ramaplthecus and

Australopithecus: <1) relative size of canines, crown

 36

height and root length more reduced compared to cheek-teeth

than is typical of even the smallest of female African apes;

<2) crown height of unworn incisors and root length reduced

relative to the area of the occlusal surface <and length) of

molar teeth; <3) dental arcades curved <parabolic), not

straight-sided or U-shaped; (4) low-cusped cheek-teeth with

1 a tera I 1 y shifted cusps; <5) c I ose I y comparab I e cheek-tooth

morphology; and (6) shallow and thick mandibular horizontal

rami.

During the 1960/s. there was increasing interest in the

relationship between the earliest hominid and Ramapithecus.

Sivapithecus was treated as one of many dryopithecines. The

similarities between Sivapithecus and Ramapithecus were not

discussed. Leakey <1963, 1967) proposed Kenyapithecus as

the earliest hominid. This judgement placed the emergence

of the Hominidea back to the Lower Miocene. Le Gras Clark

<1964:188) proposed Ramapithecus as a transitional figure

between Miocene hominoids and hominids. As a result, he

felt the hominid-pongid divergence may have occurred in the

early part of the Pliocene.

By the middle to late 1960/s, a general consensus

developed that there was an ancestor-descendant relationship

between Ramapithecus and hominids, specifically

Australopithecus <Simons and Pilbeam 1965; Reynolds 1966;

Simons 1967; Pilbeam 1968). Some investigators proposed

Ramapithecus as a hominid in its own right <Simons and

Pilbeam 1965; Simons 1967; Simons 1969; Prasad 1969).

 37

Two sources of data were used to study Miocene hominoid

evolution during the 1970/s. Fossils and other

paleontological evidence were the primary source of

information. Of a secondary nature, results from molecular

evolutionary studies were slowly incorporated into

phylogenetic analyses along with paleontological data.

1. Analyses Based on Paleontological Evidence.

In the early 1970/s. the strict Miocene phylogenies of

the 1960/s were not accepted as solidly as when Simons and

Pilbeam <1966) and Simons <1967) first introduced them.

Uzzel I and Pilbeam <1971) and Pilbeam <1968) basically

accepted the earlier consensus that Ramapithecus, even if it

was not a hominid, was ancestral to Australopithecus. The

following phylogeny represents their reservations about the

earlier view of a firm relationship between Miocene and

recent hominoids.

1. 12.:_ maJor is probably ancestral to Goril Ia.

2. ~ africanus may be ancestral to Pan.

3. 12.:_ sivalensis is tentatively ancestral to

Pengo.

4. R.:_ pun.iabicus is probably ancestral to

Australopithecus.

These pairings illustrate a more tenuous belief as

opposed to the definitive view of ancestor-descendant

relationships for these lineages proposed earlier by Simons

and Piibeam <1966). According to Andrews and Van Couvering

<1976:98), a direct relationship cannot be determined

 38

between any of the Miocene dryopithecines and the extant

African apes. It is reasonable to assume the modern pongids

only represent a small segment of the Miocene hominoids.

Because climatic changes and geographic uplifting and

rifting in Africa were occurring in the Miocene, the

environment was much different than it is today. The

Miocene hominoids were adapted to a different habitat and

those leading to the modern pongids would be difficult to

distinguish as their direct ancestors. These doubts raised

some new questions. According to the former phylogenetic

divergences hypothesized by Simons and Pilbeam <1965), Pan

and Gorilla diverged in the Miocene. But they are too close

morphologically and biochemically to have separated as early

as 20 Myr. The solution which postulated~ ma.ior as a

possible common ancestor to Gorilla and Pan preserved the

basic hypothesis involving hominid status for Ramapithecus

and an early hominid-pongid divergence.

The hominid status of Ramapithecus was not accepted

without question. According to Uzzel 1 and Pilbeam <1971),

Ramapithecus enjoyed its status through the process of

elimination. It was the only Tertiary hominoid older than 5

Myr that resembled the Hominidae significantly, a

resemblance based solely upon dental remains. It was the

dental features of Ramapithecus which led Howe! 1 <1972) to

place~ punjabicus in the Hominidae.

Uzzel I and Pilbeam <1971:620) proposed another set of

problems to the acceptance of Ramapithecus as a hominid.

 39

(1) If Ramapithecus is a hominid then it is probably

ancestral to Australopithecus and Homo. But this hypothesis

is based on the lack of evidence for a great hominid-like

radiation. And Ramapithecus is the only hominoid exhibiting

traits that are broadly ancestral to Australopithecus. <2)

If Ramapithecus is considered ancestral to Australopithecus,

then it does not matter where the taxonomic boundary between

the Hominidae and Pongidae is placed and it would not be

important whether Ramapithecus is a hominid or pongid. In

either case, the divergence would have been greater than 14

Myr.

The landmark paper by Uzzel I and Pilbeam C1971) was one

of the first which seriously questioned the simplistic

hypotheses of the previous 10 years. It also incorporated

some of the findings of the molecular evolutionary studies.

Uzzel 1 and Pilbeam <1971) did not accept the late divergence

dates of any of the molecular studies. particularly the

"molecular clock" of Sarich and Wilson <1967). They did

utilize the data gained from immunological studies

determining the degree of relatedness between Gorilla and

Pan to question whether these two taxa diverged in the

Miocene.

In an essay on paleoanthropological attitudes, Washburn

(1973) proposed a revision of the hominid-pongid divergence

dates to 8 ± 2 Myr. This is suggested as a compromise

between early and late divergence estimates. He did not

offer any support for this date, but asked for new analyses
 40

and definitions based upon the revision of Pliocene and

Pleistocene dating.

For example, everyone regarded the Pliocene

as several times as long as the Pleistocene,
begining some 10 or 12 mil lion years ago.
But the evidence has recently been reviewed
by Van Couvering </72) and the Pliocene
probably began considerably less than 6
mil lion years ago. It the Pleistocene is
regarded as beginning 3 mil lion years ago,
then Pleistocene and Pliocene are of
comparable length <as Keith thought more than
40 years ago), and the combined epochs amount
to only a third of the Miocene. Obviously,
when more radiometric dates are available, it
wil I not be necessary to use the old terms at
a! I. But the point at the moment is simply
that the argument that we "know" that the
human lineage separated from that of the apes
in the Miocene and, therefore, we "know" that
the separation "must" have been more than 12
million years ago does not necessarily hold.
One game plan might be a separation of 8 + 2
mil lion years, a late Miocene separation
played against twice that, an early Miocene
divergence <Washburn 1973:68).

By the mid-1970/s, many paleontologists realized that

what are now cal led ramapithecines were less hominid-like

than previously believed. The following illustrates some of

the changes in ideas in hominoid paleontology: (1)

ramapithecines were considered distinct from other Miocene

hominoids; <2) living populations were no longer generally

used as prototypes for fossil groups. Classifications of

fossil forms that were based upon morphological comparisons

to extant apes were not viewed as being very accurate

<Pilbeam ~ gl. 1977); and C3) multidisciplinary studies of

fossils became more widespread by this time, yielding more

comprehensive analyses of sites. Paleoenvironmental

reconstructions could be deduced from studies of geology,

 41

fauna, and flora, while intensive study and testing of the

surrounding geology could estimate the era and type of

deposition of the fossil material.

Utilizing multidisciplinary studies, Conroy and Pilbeam

<1975) presented the following hypothesis concerning

Ramapithecus. They did not firmly commit themselves to

hominid status for Ramapithecus. In comparison to some

earlier views <Simons 1969; Prasad 1969), Conroy and Pilbeam

<1975) continued along the same trend that Uzzel 1 and

Pilbeam <1971) initiated in proposing a phylogenetic tie

between Ramapithecus and Australopithecus but that position

was viewed as being even more tenuous than the one given by

Uzzel I and Pilbeam <1971). As the most plausible hominid

phylogeny, Conroy and Pilbeam <1975) accepted Ramapithecus

as the genus most likely to be directly ancestral to

Australopithecus.

Conroy and Pilbeam <1975) expanded upon their ovn

hypothesis to explain their reluctance to fully support

Ramapithecus' tie to Australopithecus. Al 1 of the

ramapithecines could not be considered ancestral to the

australopithecines. Late ramapithecines and early

australopithecines may have been contemporary if not

sympatric. While Ramapithecus was considered monotypic by

Simons and Pilbeam (1965), Conroy and Pilbeam <1975)

diversified the genus into several species. Ramapithecus

from India and Pakistan and Kenyapithecus from Kenya had

been combined into~ pun.iabicus by Simons and Pilbeam

 42

<1965). Conroy and Pilbeam <1975) split Ramapithecus at the

species level so that the Indian and Kenyan specimens were

included in~ pun.iabicus and~ wicker!. respectively.

Conroy and Pilbeam <1975) estimated the hominid-pangid

divergence at 15 Myr when Ramapithecus was considered the

earliest hominid. This is based upon the 14 Myr date

assigned to~ wickeri at Fort Ternan, Kenya <Simons 1969).

In a more decisive viewpoint, Vogel <1975) claimed that

Ramapithecus could not be classified as a hominid based

solely upon dental evidence. In his opinion, Ramapithecus

had a pongld-like U-shaped dental arcade, unlike the earlier

attribution of a parabolic arcade to Ramapithecus by Simons

<1963) that resulted from distortions in reconstruction.

Specimens of the arcade were fragmentary and descriptions

were the result of subjective reconstruction. Vogel <1975)

wanted pastcranial evidence to classify Ramapithecus as a

hominid.

Krantz <1975) agreed with Vogel/s <1975) summation of

the problem~ in reconstructing Ramapithecus/ dental arcade.

In addition, he claimed that a pre-canine diastema was

ignored in the reconstructions resulting in

misinterpretation of Ramapithecus as a hominid. Since

chimpanzees show as much interstitial wear as does man,

Krantz <1975) concluded the wear observed in Ramapithecus is

not a uniquely hominid trait. He drew his conclusions from

previous studies of ather workers and his own observations

of photographs of two Indian Ramapithecus specimens. Krantz

 43 ~ .

<1975) offered three alternative hypotheses for Ramapithecus

but none of them proposed Ramapithecus as a hominid:

1. Ramapithecus represents an extinct dead-end

group.

2. Ramapithecus is ancestral to all apes and

man while other forms <dryopithecines)
became extinct.

3. The ramapithecines are the females of a

species in which Proconsul major and~
indicus are the males.

Krantz <1975:147-149) preferred the third hypothesis.

He proposed that sexual dimorphism occurred within species

of Miocene hominoids. He suggested that the sexual

dimorphic differences between Ramapithecus and~ indicus,

such as in canine length, are the same as between male and

female baboons. The body weight ratio of the fossil species

could have been 3:1 <male to female) based on modern baboon

studies. In addition, specimens of both Ramapithecus and~

indicus have been found in the same deposits. They are also

the latest in time of the ground "apes" and presumably the

most evolved. A species with such dimorphism would have

contained the genetic potential to give rise to later

populations of either large or small body sizes as we! 1 as

to populations retaining this dimorphism.

Kretzoi <1975) analyzed the Rudapithecus specimens from

Hungary and maintained its generic independence from

Ramapithecus and Kenyapithecus because, in his opinion,

Ramapithecus was more primitive than Rudapithecus.

Ramapithecus "represents a different line of hominid

evolution" <Kretzoi 1975:579). Based on facial structure,

 44

Rudapithecus represented a direct lineoge towards Homo

<Kretzoi 1975:580). The australopithecines were members of

a side branch that developed independent of the Homo sapiens

phylogeny.

Contrary to growing scepticism, Simons <1976a)

continued to support Ramapithecus as a hominid. In his

opinion, each discovery reinforced the similarities between

Ramapithecus and Australopithecus.

By the late-1970/s, hypotheses concerning hominoid

evolution had become more complex and flexible, while the

fossil evidence was stil 1 primarily restricted to dental

remains with few cranial or post-crani~l specimens of

Ramapithecus available for analysis. Hypotheses concerned

with specific ancestral-descendant relationships resulting

in the hominoid or hominid status of Ramapithecus were no

longer accepted. Multidisciplinary studies resulted in

complex ideas and concepts where behavior patterns and

ecological adaptations became more and more important for a

complete understanding of the relationships between fossil

species. The analysis of Pilbeam et gL. <1977) illustrates

this approach. They revised the earlier and simpler

proposals of Simons and Pilbeam <1965), Pilbeam <1968),

Prasad (1969), and Simons (1969) to include

multidisciplinary information. These revisions included the

tal lowing interpretations of the fossil record. (1) The

dental differences between modern homlnids and extant

pongids are in relative tooth size and occlusal enamel

 45

thickness of the cheek teeth, not in arcade shape and

incisor size. (2) Miocene apes were more diverse than

originally proposed by Simons and Pilbeam <1965). At least

three genera in East Africa <Proconsul, Rangwapithecus, and

LimnoPithecus) should be distinguished from Dryopithecus in

Europe. (3) Postcranial remains of the Miocene apes appear

to have been more primitive, arboreal, and monkey-like that

modern extant apes. They were often compared to the

structure and function of living apes. Pilbeam ~ gl.

<1977) did not approve of these studies and believed that

fossil species are easier to interpret when analyzed as

individual groups rather than as prototypes for extant

hominoids. (4) The paleoecological information indicates

that the Miocene apes were associated with predominately

forest flora and fauna pointing more to a resemblance to

modern ceboids or cercopithecoids than to the less diverse

modern hominoids. <5) There was a revision of the

Superfamily Hominoidea to include the Families Pongidae,

Dryopithecidae, Ramapithecidae, and Hominidae. The

Ramapithecidae consist of two species clusters,

Sivapithecinae and Ramapithecinae, while the Hominidae

includes two additional clusters, Australopithecinae and

Homininae. <6) In Africa and possibly Asia, several hominid

lineages coexisted between 3.75-1 Myr.

Some conclusions based upon these revisions are as

follows <Pilbeam ~ £1. 1977:694-695): <1) Hominoids were a

relatively diverse superfamily during their evolution. They

 46

should not be subjected to a rigid classification scheme.

<2) Fossil hominoids were not necessarily like modern

hominoids and should not be compared with them in order to

avoid any misconceptions concerning their morphology and

behavior. (3) Because of the complexity of an entire

phylogeny at any particular point, apparent differences

between ancestor and descendant groups, and gaps in the

fossil record, it is difficult to draw specific and exact

ancestor and descendant relationships. Each group should be

studied in its entirety in order to understand it as a

viable species. (4) The Pongidae and Hominidae are two

taxonomic labels that should not be used to describe Miocene

species. The use of these categories could attach

preconceived ideas of the fossil species and result in

overlooking important data and obscuring evolutionary

concepts.

Pilbeam et s1. <1977:692, 695) and Pilbeam <1980:273)

presented three alternate phylogenies based upon their

revisions <Fig. III):

1. a. The Dryopithecidae were ancestral to the

Pongidae and Ramapithecidae.

b. The Ramapithecidae were ancestral to

Australopithecus.

c. Australopithecus was ancestral to Homo.

2. a. The Dryopithecidae were ancestral to the

Ramapithecidae.

b. The Ramapithecidae were ancestral to the

Pongidae and Australopithecus. The
Ramapithecidae represented the last
common ancestor of hominids and pongids.
 47

3. a. The Dryopithecidae were ancestral to the

last common ancestor of hominids and
pongids and the Ramapithecidae.

b. The Ramapithecidae were ancestral to an

undetermined form.

c. The Pongidae and Australopithecus

diverged from the same ancestor.

These more recent hypotheses are much more complex and

detailed than the previous ones based on the simplistic

assumption that Ramapithecus was most likely to be ancestral

to Australopithecus. Of these three phylogenies, Pilbeam

and his co-authors prefered the first. Pilbeam et ~-

<1977) did not disagree with the basic premise of Conroy and

Pilbeam (1975) and Uzzel 1 and Pilbeam <1971) that

phylogenetical ly ties Ramapithecus and Australopithecus

together.

Addressing the problem of divergence dating in general,

rather than the status of Ramapithecus specifically,

Washburn <1978), along with Pilbeam <1979), suggested that

paleoanthropologists consider results from molecular studies

in their analyses. Previously, as students of the fossil

record began to incorporate the results of multidisciplinary

analyses in their studies, data from molecular evolutionary

techniques were not utilized, except to observe the

molecular relatedness between individual primate species.

Because Gorilla and Pan have been shown to be closely

related <Sarich and Wilson 1967; Goodman 1974; Goodman~

~. 1983), Uzzel 1 and Pilbeam <1971) began to doubt an early

Miocene divergence between them.

 48

Figure III

Three Alternative Hypotheses of Miocene Hominoids

Pilbeam et ~- <1977)

M .o- Modern pongids Homo

Australo~s
Pleistocene I
L
L 5- Ponjidae
PI iocene I
0
N 10-
_r,Ji ocene s
0 15-
F
y 20-
E Dryopithecidae
A
R 25-
s

Fig IIIa. Hypothesis One.

M 0- Modern Pongids Homo

Pleistocene I ;r-
L Australo ithecus
L 5- PonLdae

~apithecidae
Pliocene I
0
N 10-
Miocene s
0 15-
F
~7
J. 20-
E Dryopithecidae
A
R 25-
s
 I •

49

Figure III (continued)

M 0- Pongidae Homo

Austr~IoQithecus
Pleistocene I

PI i ocene
L
L
I
0
N 10-
5-
\ 7
last common
ancestor
undetermined
species
I
Ramapithecidae
Miocene s
0 15-
F
y 20-
E Dryopithecidae
A
R 25-
s

Fig. IIIc. Hypothesis Three.

 50

2. Incorporation of Molecular Evidence.

Molecular evolutionary studies are based on comparative

protein analyses of living species. Several proteins can be

analyzed, but blood serum components in general and albumin

specifically have been the most commonly used.

The molecular clock technique, from which 4-6 Myr

<Cronin 1983; Sarich 1973; Sarich and Wilson 1967)

divergence dates have been derived for the Hominidea,

estimates an approximate date based upon the degrees of

difference between immunological responses of the same

protein for two species. This technique is not universally

accepted for producing divergence dates and some alternate

methods are used by molecular evolutionists. Goodman <1974)

suggested that the results from the clock model were not

consistent enough to be used accurately. Precise divergence

dates obtained from the molecular clock are unreliable due

to clearly marked flucuations in rates of molecular change

in evolution <Goodman et s1. 1983). It may be useful if

several different proteins are tested and their divergence

dates averaged together <Goodman 1974:220). Goodman et gl.

<1983:85) proposed another test for the molecular clock.

However, it may be noted that the vast

majority of silent mutations are not revealed
in amino acid sequence data. Detection of
these silent nucleotide replacements, which
is now possible by nucleotide sequencing,
should eventually produce sufficient data for
a more thorough test of the clock concept.

In a recent paper, Li and Tanimura (1987) suggest that

the molecular clock is slower when applied to hominids

 51

compared to the pongids. This proposal is based primarily

upon generation length in hominids. The longer the

generation length is in a species, the slower the molecular

clock should run; When generation time is used as a

variable, Li and Tanimura <1987:95) estimate the hominid and

pongid divergence was 7.7-9.9 Myr.

Goodman <1974) supported the maximum parsimony model

which constructs phylogenetic trees based on amino acid

sequences that have lrr toto the fewest number of mutations

in ancestor and descendant sequences. He argues that

molecular evolution decelerated dramatically in the

superfamily Hominoidea. Longer gestational and generation

lengths and fewer neutral mutations account for an

evolutionary slow-down. The hemochorial placenta favors the

exchange of proteins between the mother to fetal circulation

systems. The expression of mutant genes in the fetus are

more likely to be subjected to maternal immunization due to

longer gestation periods and the exchange of blood through

the placenta <Goodman 1974:222). Therefore the expression

of mutations is likely to be low in hominoids. Goodman

<1974) modified the maximum parsimony model with mutational

deceleration, and he estimated a hominid and pongid

divergence of <15 Myr.

3. Use of Molecular Data in Paleoanthropology.

Hypotheses regarding the divergence time of hominids

which were based on interpretations of the ~aleontological

evidence all proposed an early divergence of >25 Myr <Simons

 52

1967), >20 Myr <Simons 1969), 15 Myr <Conroy and Pilbeam

1975) to the 8 + 2 Myr suggestion by Washburn <1973). Since

Ramapithecus was still loosely considered a hominid by the

late 1970/s <Simons 1976a), or, at the very least, directly

related to the hominids, the 4-6 Myr divergence estimate of

Sarich and Wilson <1967), Sarich <1973), and Sarich and

Cronin <1976) did not conform to hypotheses based on fossil

evidence.

However, it is plausible that the molecular clock can

harmonize with the fossil record it generation time is not a

significant variable and neutral mutations constitute the

majority of change in a molecule <Goodman et gl. 1983:85).

But according to Fitch and Langley (1976), the neutral

mutations would have to be the most important forces of

evolution at the molecular level.

Simons (1976b) did not accept phylogenies or divergence

estimates based upon molecular evidence alone. He argued

that phylogenies should not be derived from one source of

data, but instead should be based on information from fossil

forms, living species, dating, anatomy, and chemistry.

Thus, Simons C1976b) did not think it was possible to

develop a reliable clock model using molecular data. In his

opinion, with more and more accurate dating methods in

paleontology, molecular biologists need not concern

themselves with estimating divergence dates, but only with

determining the relatedness among living species. Washburn

<1978:204) illustrates the conflict between paleontologists

 53

and molecular evolutionists in the following:

when it comes to the primates, the

molecular distance between the New World and
Old World monkeys is much too small to fit in
with conventional phylogeny. What is worse,
the distance between man and the African apes
is startlingly less than convention demands.
I suspect that if molecular anthropology had
shown man and apes to be very far apart, the
concept of a correlation between genetic
difference and time would have been accepted
without debate.

Meanwhile, the fossil evidence makes it

highly unlikely that the ape and human lines
separated less than five million years ago
and the molecular evidence makes it highly
unlikely that they separated more than 10
million years ago.

By the late-1970's, molecular anthropology began to

gain wider acceptance among paleontologists. Pilbeam <1979)

observed that the results of biochemical comparisons and the

fossil record became more compatible due to a more liberal

interpretation of the latter. He did not support using one

method over the other because the low quantity and quality

of the fossil record cannot be used to reliably test either

technique. But Pilbeam <1979:341) also cautioned against

using living primates to construct prototypes of a fossil

species, Just as fossils should no longer be used to

exemplify modern forms:

In my opinion, the hominoid fossil record is

still too poor for it to be used reliably in
evaluating the various hypothetical
evolutionary schemes based on comparative
studies of living hominoids, either
anatomical or biochemical. Until recently,
implicit expectations of what the fossil
record should tell us, based on ideas derived
from studies on the living have biased many
attempts at analysis.
 54

Thus, to Pilbeam (1979), Ramapithecus no longer

resembled the hominid lineage as clearly as it once did. A

few postcranial fragments were discovered in Pakistan that

showed ape-like characteristics. Based on size, these

postcranial remains were attributed to Ramapithecus,

Sivapithecus, and~ bilaspurensis (Pilbeam 1979:347).

However, none of these fossils was found in direct

association with any of the cranial or dental material. The

few postcranial specimens tentatively classified as

Ramapithecus, were from a small arboreal hominoid <Pilbeam

1979:349). As a result, he concluded Ramapithecus was not

necessarily a biped. Yet the fossils had enough

hominid-like traits that they could easily have been

interpreted as hominid by other researchers. While Pilbeam

(1979) did not actually propose Ramapithecus as the last

common ancestor of the hominids and pongids, he did consider

the idea to be worthy of study. His conclusions represented

a compromise between the paleontological divergence

estimates of 15 Myr <Conroy and Pilbeam 1975) and that of

the molecular clock of 4-6 Myr <Sarich and Wilson 1967)

based upon the phylogenetic status of Ramaplthecus.

Compared to the situation in the 1960's, when there

were fewer fossils and simpler assumptions regarding

Ramapithecus/ hominid status, Ramapithecus was studied in

greater detail by the 1970's, due to the steadily increasing

number of finds of fossil specimens <Simons and Pilbeam

1965; Simons 1967). One result of the later studies was

 55

that Ramapithecus was given only tentative status as a

hominid <Uzzel I and Pilbeam 1971; Conroy and Pilbeam 1975).

During this period, dating estimates from multidisciplinary

studies, including molecular comparisons, were incorporated

into hypotheses of the hominid-pongid divergence.

By the mid-1970/s, Ramapithecus was treated as a fossil

hominoid in its own right rather than how it could be made

to fit as an ancestor to Homo. Ramapithecus was not

classified based upon which extant living species it most

resembled <Pilbeam et £1. 1977). In the previous decade,

Ramapithecus was referred to as a monotypic genus <Simons

and Pilbeam 1965). By the mid-1970/s, a number of

investigators rejected the hominid status of Ramapithecus

<Conroy and Pilbeam 1975; Kretzoi 1975; Krantz 1975; Vogel

1975).

There was even more skepticism in the late-1970/s

concerning Ramapithecus than in the early portion of the

decade. Paleoanthropologists did not make broad assumptions

about Ramapithecus based upon dental evidence alone <Vogel

1975; Pilbeam 1979; Greenfield 1978). Biomolecular

comparisons were being utilized in the study of hominid and

pongid origins by paleontologists <Pllbeam 1979, 1980).

These more detailed studies led to hypotheses that only

considered an evolutionary tie between Ramapithecus and

Australopithecus <Pilbeam 1979, 1980). While Ramapithecus

was no longer considered a hominid by many researchers,

neither was it labeled a pongid or dryopithecine, but rather

 56 ~ .

retained its individuality as a genus and as a hominoid.

 Chapter III

The 1980"s

Recently, the hominid status of Ramapithecus has been

questioned anew. Ramapithecus was even questioned as a

possible hominid ancestor <Andrews 1982:186). In 1980, at

the symposium titled Miocene Homlnoids and New

Interpretations of Ape and Human Ancestry held in Florence,

Italy, the phylogenetic status of Ramapithecus was

criticized by a significant number of scientists <Andrews

1983; Boaz 1983; Ciochon 1983; Corruccini and Ciochon 1983;

DeBonis 1983; Greenfield 1983; Ward and Pilbeam 1983;

Wolpoff 1983). In the latter 1970"s, more ramapitheclne

specimens were recovered from Pasalar, Candir, and the

Middle Sinap in Turkey, Lufeng County in China, and the

Potwar Plateau in Pakistan, to name a few localities. These

fossils led to many re-evaluations of the placement of

Ramapithecus. Even the~ afarensis finds at Hadar

<Johanson and White 1977) contributed to the reassessment of

Ramapithecus <Greenfield 1980, 1983:701; Corruccini and

Ciochon 1983:13).

Corruccini and Ciochon <1983:7), along with Lipson and

Pilbeam (1982), have condensed some of the new controversies

into three categories:

1. Ramapithecus is a sister group to

Slvapithecus and shares a functional complex
with early hominids, indicating a tendency
towards heavy chewing.

2. Ramapithecus and Sivapithecus are no longer

distinct genera.

57
 ----~·-----·- --~--· --~~---.....--- --~--

58

3. The general consensus among

paleoanthropologists has moved away from the
idea that Ramapithecus was the earliest
hominid and therefore directly ancestral to
the australopitheclnes.

Since 1980, questioning of previously accepted

hypotheses regarding the hominid status of Ramapithecus have

come about due to the re-evaluation of some basic

ramapithecine characteristics. For example, the

microstructure of dental enamel in Ramapithecus was once

considered to be hominid-like, but more recently the same

characteristic is being re-evaluated and may not be

considered hominid-like after all CCorruccini and Ciochon

1983:7).

Pilbeam (1986:295-296) now believes that thick enamel

such as seen in Ramapithecus and Australopithecus is a

primitive trait and thin enamel, as exhibited by Pan, is a

derived characteristic. This would indicate the

relationship between Ramapithecus and Australoplthecus that

is based upon enamel thickness is one of convergence or

parallel development rather than inheritance from ancestor

to descendant.

Another change that has complicated current studies is

a redefinition of the taxonomic nomenclature. Most, but not

all paleoanthropologists (Andrews 1982; Andrews and Tekkaya

1980; Greenfield 1980; Kay and Simons 1983; Ward and Pilbeam

1983) have lumped the genus Ramapithecus into the genus

Sivapithecus. This trend started when Greenfield (1978:345)

refused to separate Ramapithecus and Sivaplthecus based upon

 ---------- __ . . ~...-=-- . . - .. -~- --
59

dental and gnathic evidence. This combination at the genus

level is counteLed with a diveLsification at the species

level that is Leminiscent of pLe-Simons and Pilbeam <1965)

classification systems. AndLews <1982) incorporated

Ramapithecus into a Sivapithecus complex wlth four species

Langing in age from 8-12 Myr. However, in a lateL study,

Andrews <1983) did not necessar"ily combine the two genera.

Instead, he proposed Sivapithecus and Ramapithecus as sister

groups; that is, they represent two fossil groups which are

closer to each other than to any other group. In agreement

with Andrews <1983), Pilbeam (1985, 1986) re-evaluated the

taxonomic classifications of Slvapithecus and Ramaplthecus,

and concluded that they should be separate genera.

Kay and Simons <1983:581) also revised the

classification of Ramapithecus and placed lt in the genus

Sivapithecus. They recommended that the range of

variability within extant monkey and ape species can and

should be used to assess variation in a fossil hominoid

species. This proposal is based upon 11

improved scientific

understanding of variability within living species .. <Kay and

Simons 1983:581). If this statement ls valid, then the

range of variation in Sivapithecys is more than enough to

accomodate Ramapithecus. GLeenfield (1980:353) indicates

that the differences between Sivapithecus and Ramapithecus

were no greater than those between Pan troglodytes and Pan

panlscus.

The merging of Sivaplthecus and Kenyaplthecus by some

 60

investigators <Andrews and Tekkaya 1980; Greenfield 1980;

Kay and Simons 1983; Wolpoff 1982) has added new

significance to the status of Sivaplthecus. If

Kenyapithecus is accepted as a sivapithecine, then

Sivaplthecus could be as old as 17.2 Myr <Leakey and Walker

1985:173; McDougall and Watkins 1985). The combination

would also expand the slvapithecine radiation to East Africa

based upon fossils discovered at Fort Ternan, Rusinga

Island, Buluk, and Maboko, Kenya (Leakey 1962, 1963;

Pickford 1985; Leakey and Walker 1985). Analyses of

Sivapithecus and Kenyaplthecus range from combination into

one genus <Kay and Simons 1983) to a total separation

<Pickford 1985a, 1985b; Boaz 1983). Pickford C1985a,

1985b), Boaz (1983>, and Pllbeam <1986:305) do not accept

inclusion of Kenyapithecus Into Sivapithecus. Pickford

(1985a) however, does allow for some kind of relationship

between them.

I currently accept the position that~

afrlcanus is generically distinct from
Sivapithecus and Ramapithecus. However, both
may be considered to be parts of a single
adaptive radiation <Pickford 1985:134).

Boaz (1983) also proposes an ancestor - descendant

relationship between Kenyaplthecus and Sivapithecus.

Sivapithecus is concluded to be ancestral to Gigantopithecus

and Pongo.

While the relationships between Slvapithecus,

Ramapithecus, and Kenyapithecus are being discussed, there

have been some new Miocene homlnoids introduced. Kretzol

 61

<1975) descLibed a hominoid fLom HungaLy, Rudapithecus

hungaLicus, as closeL to Kenyapithecus than to Ramapithecus.

Rudapithecus hungaLicus is also descLibed as less pLimitive

and ape-like than Ramapithecus. Kay and Simons <1983:592)

now classify Rudaplthecus as conspecific with~ fontani.

They also classify GLiphopithecus suessi fLom the Vienna

Basin as a dLyopithecine. OtheL Lecent Miocene hominoids,

specifically, OuLanopithecus and GLacecopithecus fLom

GLeece, AnkaLapithecus fLom TuLkey, and Bodvapithecus fLom

HungaLy have all been lumped into Sivapithecus by Kay and

Simons <1983:604). Pilbeam <1986:305) does not agLee with

Kay and Simons <1983). He excludes OuLanopithecus and

Rudapithecus fLom Sivaplthecus. Recently Leakey and Leakey

<1986) have intLoduced anotheL laLge-bodied Miocene

hominoid, AfLopithecus tuLkanensis, fLom KalodlLL, Kenya.

They have also deteLmlned that ~ turkanensis is distinct

from Dryopithecus, Ouranoplthecus, Rudapithecus, and

Bodvaplthecus. Lea~ey and Leakey <1986:144-145)

reclassified the Sivapithecus material found at Buluk,

Kenya, as~ turkanensls, and confined Sivaplthecus to Asia.

FouL majoL hypotheses will be discussed in the

LemaindeL of this chapteL concerning the status of

Sivapithecus and Ramapithecus. <1) The eaLly diveLgence

hypothesis in which Sivapithecus is a hominid descendant of

the last common ancestor of the hominid and pongid lineages.

It ls also ancestral to the australopithecines and Homo <Kay

and Simons 1983). (2) The late divergence hypothesis in

 62

which Sivaplthecus is the last common ancestor of the

hominids and pongids and not a hominid. It is also not

considered to be a dryopithecine <Greenfield 1980, 1983).

(3) Slvapithecus is ancestral to Pengo but not the African

Great Apes <Andrews 1982; Andrews and Tekkaya 1980). It is

therefore older than the last common ancestor of hominids

and the African Great Apes. And <4), The establishment of

two sympatric populations of Late Miocene hominoids at

Lufeng, China. The Chinese ramapitheclnes are not

integrated into one sexually dimorphic species of

Slvapithecus but represent distince genera: Ramaplthecus

and Slvapithecus.

Each of these hypotheses will be discussed with a focus

on the evolutionary status of Sivapithecus and Ramapithecus

and its effect upon estimating a divergence date between the

hominids and ponglds. Also. unless specifically named,

Ramaplthecus is included in the genus Sivaplthecus.

 _, ----- ---- ---

63

A. Early Divergence Hypothesis.

The early divergence hypothesis as described by Kay and

Simons <1983) proposed a hominid-pongid divergence occurring

during the Middle Miocene. This hypothesis would establish

the hominid and pongid divergence prior to or near the

emergence of Sivapithecus.

Kay and Simons (1983) are the staunchest supporters of

the sivapithecines as either the earliest hominids or

descendants from the last common ancestor of hominids and

pongids. They recognize only two genera of ramapithecines,

Sivapithecus and Gigantopithecus, and Dryopithecus is the

only acknowledged dryopithecine. Kay and Simons <1983:604)

do not support Kenyapithecus as an independent genus. They

think that the specimens attributed to Kenyapithecus are

very similar to and easily incorporated into Sivapithecus.

Kay and Simons (1983:620) "believe available evidence is

strong enough to indicate that ramapitheclnes are broadly

ancestral to Pliocene-Recent Australopithecus and Homo, but

not to any living ape".

The "available evidence" Kay and Simons <1983) refer to

is the result of comparative anatomical studies among

Sivapithecus, Australopithecus, and the extant great apes.

Kay and Simons <1983) have divided the traits into those

that are shared derived characteristics and those that are

the result of parallelism. Those traits defined as shared

derived are used to support close relationships between

phylogenetic groups.
 64

In assessing the phylogenetic position

of Ramapithecinae <Sivapithecus and
Gigantopithecus), we believe that only shared
derived characters can indicate a close
relationship between taxa. By this
analytical strategy, first we identified
preserved anatomical characters which show
some variation <exhibit several different
/states/> among Miocene to Recent Great Apes.
Once this was done, we determined which
character states were likely primitive for
the group and which represent derived
conditions. Ideally, any time two taxa share
a derived state, they should be more closely
related to one another than either is to the
other taxa being considered. But what has
become apparent to us is that considerable
parallel evolution among Miocene to Recent
apes frequently obscures the true
phylogenetic picture <Kay and Simons
1983:618).

The following traits are those which Kay and Simons

<1983:619) have defined as uniquely shared derived

characteristics that link the slvapithecines and early

Australoplthecus:

<1> shallow, broad mandibular corpora, robust

mandibular symphyses, (2) low-cusped molars
with extremely thick enamel, <3> somewhat
reduced canine size, (4) somewhat reduced
canine sexual dimorphism in Slvaplthecus
<both traits 3 and 4 foreshadowing the
condition of Australopithecus>, <5>
buccal-lingually broad, mesial-distally short
upper canines, and <6> a tendency toward
great enlargement of the P3 metaconid.

Those traits that are the result of parallelism are

explained in the following definition:

Given that a certain amount of

parallelism is inevitable in all comparisons
of this kind, we analyzed as many characters
as possible and assumed that the phylogeny
which is most likely has the fewest
parallelisms and, where the parallelisms are
found, they occur in characters showing a
high frequency of similar parallelisms in
other primate groups <e.g. the characters in
 65

question are assumed to have a low 'phyletic

valence') Kay and Simons (1983:618).

Any characteristics that appear to be unique shared derived

traits <synapomorphies) linking Sivapithecus with the

pongids are defined by Kay and Simons <1983> as parallelisms

or retentions from a Proconsul-like ancestor and the last

common ancestor. Some of these traits Indicate advancement

over Proconsul towards a pongid grade of development but not

advancement toward Australopithecus. As a result, they

believe Sivaplthecus could not have been the last common

ancestor of extant apes and hominids. The following are

traits believed by Kay and Simons <1983:613-614> to be

parallelisms between Sivapithecus and the extant apes.

1. Zygomatic arch is raised above the level of

the occlusal plane.

2. An arched palate (primitive in

Australopithecus>.

3. Zygomatic deep and flaring (primitive for

great apes>.

4. Greatly reduced molar cingula and limited

accusal relief.

Kay and Simons <1983:619) define the genus Sivapithecus

as a dental hominid exhibiting canine-size reduction and

reduced sexual dimorphism. But it also shows cranial and

post-cranial features reminiscent of the extant great apes

with evidence of arboreal locomotion. Kay and Simons <1983)

relied upon fossil evidence as a primary source for their

analyses but they also compared anatomical characteristics

between fossil hominoids and modern primates.

The evidence presented by Kay and Simons (1983:616)

 66

leads them to estimate a divergence between the last common

ancestor of hominids and Pengo during the Middle Miocene at

15 Myr. Since the earliest date known at the time of their

analyses for Sivaplthecus <at Fort Ternan) Is 14 Myr, a 15

Myr divergence is consistent when Sivaplthecus is granted

hominid status. If Kay and Simons <1983) accept the Buluk,

Kenya, material as Sivapithecus and/or revise the status of

Kenyapithecus, a more recent date of 17.2 Myr <McDougall and

Watkins 1985) from this locale will force the early

divergence estimate to approximately 18-19 Myr.

Prasad <1983) is not quite as decisive about the

evolutionary standing of Ramapithecus as Kay and Simons

<1983). He lists a series of traits that only "suggest" a

potential phylogenetic relationship between Ramaplthecus and

the African Plio-Pleistocene hominids <Prasad 1983:560):

1. A modification of the dryopitheclne Y-5 cusp

morphology that resulted in broader grinding
surfaces for a graminivorous diet.

2. A shortening of the mandible resulting in a

reduction of the diastema space.

3. An absence of cingular extension of the

premolars.

4. Molarization of the premolars.

5. Vertical placement of the incisors.

6. A possible parabolic contour of the maxilla.

The ramapithecine that would be the most likely

candidate for the earliest hominid is~ punjabicus. Prasad

<1983) does not include~ pubjabicus as a slvapitheclne nor

does he include Kenyapithecus wicker!, as described by

 67

Leakey <1962), as a ramapithecine or a sivapithecine. This

separation of Ramapithecus, Kenyapithecus, and Sivapithecus

from each other blends well with Kay and Simons; <1983) idea

of hominid status for ramapithecines. Prasad <1983:568)

suggests the following terminology for the Miocene

hominoids, which promotes taxonomic diversification:

1. ~ sivalensis

2. ~ indicus

3. ~ pun.iablcus

4. ~ chin.iiensis

5. ~ giganteus

6. L. afrlcanus
Prasad <1983:569) also argues for the separation of

Sugrivapithecus from Sivapithecus at the genus level or, at

the very least, establish a new species for the the former

genus within Sivapithecus.

Although Kay and Simons <1983) and Prasad <1983)

hypothesize Ramaplthecus as the earliest hominid, they refer

to different fossils. Kay and Simons <1983) refer to

Sivapithecus as a genus, and specifically regard the East

African Sivapithecus africanus of Fort Ternan as a hominid.

Prasad (1983), on the other hand, proposes~ pun.iablcus

from the Siwaliks as a potential hominid.

Prasad <1983:560) dates the Siwalik Ramapithecus at

8-10 Myr. His hypothesis of a possible hominid status for

the later ~ pun.iabicus, as opposed to the older African

Sivapithecus, would imply a divergence date of 10-12 Myr for

 68

the hominids and pongids.

 69

Figure IV

Cladogram of the Early Divergence Hypothesis

Ciochon <1983:812)

Gi~ons Dryopiths OranjUtans African

A es
\
\
\
\
/
\
\
\
\
\
\
\
\
\
 70 ,.

B. Late Divergence Hypothesis.

The late divergence hypothesis as described by

Greenfield <1980:351, 1983), Shea (1985), Pilbeam <1980),

and Wolpoff <1982), proposed a hominld-pongid divergence

occurring in the Late Miocene. This hypothesis would

establish the hominid and pongid radiation near the end, or

possibly after, the dates established for Sivapithecus.

According to Greenfield <1980:351-352), the late

divergence hypothesis postulates the following: (1) None of

the Middle Miocene hominoids can be singled out exclusively

as the phyletic ancestor of Australopithecus and Homo. The

ramapithecines have too many dryoplthecine-like traits to be

ruled out as a potential ancestor to both the extant apes

and Australopithecus. (2) ~ afarensis shows several

dryopitheclne-like traits that suggest a post-Miocene

development of most hominid dental features. The hypothesis

may call for a redefinition of what is a hominid versus a

pongid trait. (3) Proconsul, assumed to be the last common

ancestor of hominids and pongids by Kay and Simons <1983),

predates the last common ancestor. Proconsul Is dated at L

16 Myr <Kay and Simons 1983) and Sivapithecus/ latest date

is about 6-7 Myr. Sankhyan <1985), however, describes a

date of 5.5 Myr for a Sivapithecus right mandibular molar

<HT.A10) from Haritalyangar, India, in the Siwalik Region.

There are no special similarities between Proconsul and Pan

or Gorilla to attribute a phyletic relationship between

them. Proposals for such a relationship have been based

 71

upon the assumption that a tie exists between Ramapithecus

and Australopithecus. The radiation of the pongids occurred

first in the split leading to Pongo and was followed at a

later date by the divergence of the African apes and

hominids. This two-state divergence is supported both

paleontological ly and biochemically. <4) Sivapithecus may

well be the last common ancestor of hominids and African

Apes even though there would be some dental reversals

experienced by the extant apes. Greenfield <1983) proposes

only a general relationship of Sivapithecus to hominids and

extant great apes as their last common ancestor.

Greenfield (1980, 1983) utilizes two primary dating

sources for analysis including: (1) The fossil record and

(2) comparative studies of extant apes and man on two

levels. The first level of comparison involves embryology,

ontogeny, and gross anatomy and the second method analyzes

immunological differences of serum proteins and DNA.

Comparisons of serum proteins and DNA provide more

convincing data bearing on phylogenies than other studies

because genetic similarities and differences are clearly

defined <Greenfield 1980:353). Theoretically, the fossil

evidence should provide a test for the conclusions of

phylogenetic relationships based upon biochemical analyses.

But the fossil record of Miocene and Pliocene hominoids is

too poor to provide a reliable test of these results.

Greenfield <1980:353) states the following concerning

paleoanthropologlcal attitudes towards molecular

 72

evolutionary studies:

However, because the relevant fossils

necessary to set up a test are presently
lacking, most paleoanthropologlsts,
implicitly or explicitly, have considered the
phenetic evidence of human/ape phylogeny
(just the suggested general relationships and
not the dating of the branching events) a
relatively more important, or a more
convincing source of phylogenetic information
than the fossil record.

Immunological testing indicates that Homo, Pan, and

Gorilla are closer to each other phylogentically than any

one of them is to Pengo <Goodman 1983; Cronin 1983). It is

generally accepted from these analyses that Pengo diverged

from the main hominoid stock earlier than did the African

apes and man. The best test for these results is the fossil

record itself but, unfortunately, the fossil record for the

Late Miocene is too fragmentary to support specific

relationships between lineages.

Greenfield <1980) also incorporated Ramapithecus into

Sivaplthecus, emphasizing dental evidence. The differences

between Ramapithecus and Sivapithecus are not as great as

those between Pan and Gorilla and are equal to those between

Pan panlscus and Pan troglodytes. He rejects any suggestion

that Sivapithecus and Ramapithecus are really sexually

dimorphic representatives of the same species. Greenfield

<1980:353) reclassified~ pun.iabicus and~ wickeri as~

brevlrostris and~ africanus, respectively. Greenfield

<1980:354) reduces the role of Slvapithecus in the late

divergence hypothesis in the following argument:

IF: Ramapithecus alone was perceived to be

 73

uniquely on the way to being human and

not at all related to the extant apes;
AND: Sivapithecus alone predates important
11
hominization 11 trends and possesses
enough shared morphology to make it a
potential ancestor to the hominids and
pong ids;
THEN: Slvapithecus / Ramaplthecus does not
possess the traits to make itself
uniquely a hominid. The human
characteristics seen in Ramapithecus
are the dryopithecine-like traits
possessed by ~ afarensls.
According to this interpretation of
/Ramapithecus/ and /Sivapithecus/, there is no
necessary expectation that /phyletic/
ancestors of each of the extant great apes and
Australopithecus existed by the beginning of
the Middle Miocene, or if they did exist, that
they were clearly distinguishable <Greenfield
1980 :354).
The late divergence hypothesis is largely supported by:
<1> a continuation of many dryopitheclne dental
characteristics through Aystraloplthecus and leading to
Homo, <2) the establishment of~ afarensis as a
transitional form between the dryopitheclnes and Homo, and
<3> the presence of canine sexual dimorphism in Sivapithecus
indicating a relationship with the extant great apes. But
the presence assumes the reduction of sexual dimorphism did
not occur until the Late Miocene.
Assuming that~ afarensis 1s a transitional form,
Greenfield <1980) claimed its ancestors would show reduced
canine sexual dimorphism compared to the extant great apes.
Given this assumption, he proposed three possibilities for
the phyletic ancestor of ~ afarensis:
 --- -- - -- - ~~·.~~"

74

Figure V
Late Divergence Phylogeny
Greenfield <1980:361)

M 0- Pengo Pan
-,- Gor- i I 1a Homo
I
Pleistocene I
L
L 5-
~ blacki
I
I
I I
I
I
' '\.
'
Australooithecus
: I
Pliocene I I
I - .
I
I ' '-1?I ...... ../ - ..--
0 ~ bilasourensls
<. .......
......
...... ......
I
N 10- .....
...... ,,j ........
Miocene s ....... ...... /

',?. . . . . . . .
0 15- Sivaeithecus
I
F I
D. <Dr:toQlthecus)
y 20- I
I

E <Proconsul)
A
R 25-
s
 75

1) a Middle Miocene phyletic ancestor ls

unknown; or 2) that it is known but is a
dryopithecine and indistinguishable from
contemporary species; or 3) that a phyletic
ancestor did not exist until the Late Miocene
<Greenfield 1980:355).

Greenfield <1980:363) presents Sivapithecus as the last

common ancestor to the homlnids and pongids. While it does

not qualify as either a form uniquely ancestral to the

extant great apes or to Australopithecus, there are many

characteristics shared between Sivapithecus and the extant

great apes and Australopithecus that are not shared between

Australopithecus and the extant apes. Sivapithecus also

exhibits characteristics that are shared between

Australopithecus and the extant apes. Greenfield <1980:352>

first estimated the divergence at 6-14 Myr, then later

<Greenfield 1983:702) revised lt to 5-10 Myr.

Shea <1985> supports a stronger tie between the

sivapithecines and hominids and pongids based on comparisons

of cranial morphology between the African Apes and Pengo.

He has found that <1> the exterior shape of the skull is

essentially the same between Gorilla and Pengo after basic

size differences are taken into account. <2> The

fundamental differences of the internal morphology center

around the position of the endocranial base. This results

in different angles of the position of the face. His

proposal endorses an undetermined species of SlvaPlthecus as

the ancestor of al 1 of the 11

extant large bodied hominioids 11 ,

including Pengo, Pan, Gorilla, and Homo <Shea 1985:329).

In 1980, Pilbeam outlined three different phylogenies

 76
' '

that elaborated upon his ideas from 1979 and incorporated

multidisciplinary data into his information base, including

molecular and anatomical comparisons, and fossils:

1. The hominids are descended from

ramapithecids and were evolved by 6 Myr.

2. Ramapithecids were ancestral to both

hominlds and pongids with hominids distinct
by 6 Myr.

3. Ramapithecids became extinct and were

replaced 6-10 Myr. Hominids were descended
from a late dryopltheclne or early pongid.

Phylogeny One is Pilbeam~s C1980) prefered hypothesis.

This hypothesis alone postulates an evolutionary tie between

Ramapithecus and the hominids. Pilbeam C1980:279) did not

accept hominid status for Ramapithecus and dated the

establishment of the hominids to be at least 6 Myr which is

more compatible with estimates based on comparative

biochemistry rather than those founded on paleontological

evidence. But Pllbeam C1980) preferred to use results from

molecular and anatomical comparisons along with

paleontological evidence to offset the ambiguity of the

fossil record.

Wolpoff (1982, 1983) also does not name Sivapithecus as

the last common ancestor. He is consistent in his usage of

the term Ramapithecus and ramapithecine for the same fossils

that have been referred to as Sivapithecus and

sivaplthecine. To retain continuity in this paper, the

terms Sivapithecus and slvapltheclne will be subsitltuted

for Wolpoff~s use of the term ramapithecine. He suggests

that the divergence of the homlnids, African apes, and Pengo

 77

from the basic hominoid stock occurred as part of the

radiation and diversification of the sivapithecines. This

hypothesis supports a phylogenetic tie between Asian

sivapitheclnes and Pengo~ and at the same time does not

include any hominoids formerly referred to as Ramapithecus

in the Pengo lineage.

Wolpoff (1982:507) does not define the sivapithecines

as hominids. However~ he does propose that the earliest

hominids were a specialized form of slvapithecine. While

this statement seems contradictory~ the statements are not

mutually exclusive. The earliest hominids were

sivapithecines that diversified and underwent hominization

specializations until they were distinctive enough to be

labeled 11
hominid". Wolpoff <1982~ 1983) claims~ however~

that the hominization trends did not originate in the

sivaplthecine that eventually developed as a hominid. Many

of them are seen in all of the slvapithecines and some of

the pongids~ particularly the African apes. Some of these

shared characteristics are: (1) rudimentary tool-use, <2)

omnivorous dietary adaptations~ (3) incipient bipedalism,

and (4) a limited, symbolic-based open communication system

<Wolpoff 1982:508).

Unfortunately, the inadequacy of the fossil record,

coupled with underestimation of divergence dates, prevents

precise determination of a divergence point. Sankhyan

<1985) has recently published a late estimate of 5.5 Myr for

Sivapithecus from the Indian Siwaliks. Sankhyan <1985)

•, - . . ~ '···-~.., ...
 78

claims this date to be firmly supported by paleomagnetic,

stratigraphic, and biostratigraphic dating techniques.

Using a non-linear modification of the molecular clock to

incorporate fossil evidence, Gingerich <1985) estimates an

8.9 Myr Homo-Pan divergence. According to Pilbeam

<1980:279), the divergence could neither be later than 6

Myr, nor earlier than 10 Myr <Greenfield 1983), while

Wolpoff <1982, 1983) only estimates the Late Miocene as a

likely time period for the divergence. A more recent

proposal by Pilbeam <1985, 1986) estimates the divergence to

have been 5-10 Myr.

!
 79

Figure VI

Cladogram of the Late Divergence Hypothesis

Ciochon (1983:813)

Dryopi ths Gi~ons Ramapiths Humans

\
\
\
\
\
\
\
\
\
 80

C. Sivapithecus is an Ancestor to Pengo.

The third major hypothesis regarding the phylogenetic

position of Sivapithecus concerns its morphological

affinities to Pengo. Several scholars have observed

similarities between Sivapithecus and Pengo using different

approaches. Some workers <Andrews 1982, 1983; Lipson and

Pilbeam 1982; Schwartz 1984; Pilbeam 1985) have even

hypothesized an ancestor - descendant relationship between

Sivapithecus and Pengo. The following are some of the

postulated premises for this hypothesis <Andrews 1982;

Andrews and Tekkaya 1980; Schwartz 1984):

1. Sivapithecus and Ramapithecus may be

incorporated into one genus.

2. Sivapithecus and Kenyapithecus may be

incorporated into one genus.

3. Sivapithecus and/or Ramapithecus is part of

the orangutan clade.

4. Sivaplthecus cannot be classified as a

hominid.

If Sivapithecus is ancestral to Pengo, then

Sivapithecus and/or Ramapithecus must be included into the

orangutan clade.

The similarities <of Sivapithecus) with man

are fewer in number and more strongly
inter-related with each other than are the
similarities with the orang-utan, and it is
my belief that Slvapithecus must be seen as a
relative of the orang-utan <Andrews
1982: 186).

It is also generally accepted from molecular studies

<Goodman 1983; Cronin 1983) that the African apes and Homo

are more closely related to each other than either is to

 81

Pengo. Andrews <1982:186) analyzed these relationships with

respect to Sivapithecus and found that Sivapithecus and

Pengo are more closely related to each other than either is

to the African apes and Homo.

Current ideas reviewed by Pickford <1985) suggest a

reclassification that places Pan and Gorilla into a sister

group with Australopithecus and Homo, either at the family

or sub-family level <Pickford 1985:113). This would put Pan

and Gorilla into Paninae or Panidae and Australopithecus and

Homo into the Hominiae or Hominidae, as discussed by Boaz

and Cronin <1985). Slvapithecus would be placed in Pongidae

sensu stricto.

Support for a direct relationship between Sivapithecus

and Pengo lies mainly within two areas; (1) gross

morphologlcal comparisons and <2> statistical analyses of

patterns of sexual dimorphism in both living and fossil

populations.

Most of the gross anatomical comparisons between

Sivapithecus and Pengo are concerned with facial and dental

characteristics including the following:

1. Narrow interorbital septum <Andrews 1983;

Andrews and Tekkaya 1980).

2. Deep zygomatic region with strong lateral

flare (Andrews 1983; Andrews and Tekkaya
1980).

3. Broad nose <Andrews 1983).

4. Broad ! 1 significantly larger than ! 1

(Andrews 1983) .

5. Marked alveolar prognathism <Andrews and

Tekkaya 1980).
 82

6. Short upper face <Andrews and Tekkaya 1980).

Much of Andrews <1982, 1983) support lies in what is

known as the Pakistan face <GSP 15000). A~ indicus face

was found in the Siwaliks in 1979-80. It consists of most

of the left face and was described by Pilbeam (1982> as

possessing several similarities to Pengo in general

morphology. He does not state that ~ indicus was ancestral

to Pengo, but rather that an Asian form similar to~

indicus, was. Regardless of Pilbeam/s <1982) analysis,

Andrews <1982, 1983) uses the Pakistan face as a major

support for Pengo ancestry.

On a larger scale, Andrews <1983) uses~ meteai as a

basis for a linkage between Sivapithecus and Pengo. Andrews

and Tekkaya (1980) described a fossil <MTA 2125> consisting

of a complete palate and lower face that they later assigned

to ~ meteai from the Middle Sinap series from Turkey. In

their analysis, Andrews and Tekkaya <1980:94) report that

the orangutan dentition is closer to the maxilla and

dentition of~ meteai than to any of the other extant apes.

Andrews and Tekkaya <1980:94) support a relationship between

the two genera, but qualify the hypothesis in the following

cautionary statement:

Some of these characters (those listed above)

are also present in the maxillae of~ meteai
and Ramapithecus, but whether all of them are
therefore related to the orang-utan or
whether the similarity is due to a functional
convergence resulting, perhaps, from similar
dietary adaptations to similar habitats, is
not known at present. Sivapithecus species
have been considered by many authors in the
 83

past <see Simons and Pilbeam 1965 for

discussion and references> to represent
ancestral orang-utans mainly because the
fossils are known in the right place <Asia)
and the right time <middle to late Miocene>.
Such arguments carry no weight in the absence
of morphological similarity, but with the
evidence of similarity presented here between
~ meteai and the orang-utan the likelihood
must be increased that they are related.

According to Lipson and Pilbeam <1982:546>, the

following general traits may be shared derived

characteristics between Sivapithecus and Pongo: <1> sub

nasal configuration, <2> orbital morphology, <3> facial

foramina, and <4> small incisive foramina. If so, then

Sivapithecus and some or all of the other ramamorphs would

be more closely related to Pongo than to the African apes or

hominids.

The alternative, that ramamorphs are

ancestral to all great apes and humans, is
possible, but improbable, because it seems
likely that at least some of the ramamorph
features are derived traits which are shared
only with orangs among the living hominoids
<Lipson and Pllbeam 1982:546>.

While Lipson and Pilbeam <1982> endorse a Pongo

relationship with Sivapithecus, they refuse to identify a

particular sivapithecine, nor do they assign the ancestry of

Pongo to any specific, known ramamorph. But they do

postulate an unidentified ancestor to a similar late Miocene

species that was most likely located in Asia. This

viewpoint follows the ideas of Wolpoff <1982, 1983)

concerning an adaptive radiation of a Late Miocene hominoid

species, probably a sivapithecine leading to Pongo.

Sankhyan <1985> uses the analysis of a single fossil

 84

CHT. A10> from Haritalyangar, India, to support an argument

for affinity between Sivaplthecus and Pengo. This single

molar is dated at 5.5 Myr which is significantly later than

most estimates for the divergence of Pengo but much less

than that for the divergence of Homo and the African apes.

It also suggests a long temporal continuity

and a great viability of this hominoid taxon
and may have important implications in the
wake of the current debate over the taxonomic
status of Sivapithecus and human-great-ape
relationships <see Ciochon and Corruccini,
1983). However, the present author views
Sivapithecus as the last common ancestor of
the Plio-Pleistocene Australopithecus-Homo
and the Asian Pengo <Sankhyan 1985:577).

But one tooth is not enough on which to base an entire

hypothesis. 5.5 Myr is the latest published date for a

Sivapithecus find, and in order for this date to be

representative of a later population, a larger sample needs

to be provided.

Dating the homlnld-pongld divergence from Sankhyan~s

(1985> hypothesis places most dating estimates between those

derived from the early and late divergence hypotheses. The

early divergence hypothesis has an estimate of 15 Myr <Kay

and Simons 1983) and the late divergence is 5-10 Myr

(Greenfield 1983; Pilbeam 1985, 1986>. Andrews <1982)

considers paleontological dates of 8-14 Myr for Sivapithecus

as quite compatible with molecular estimates of 10 ± 3 Myr

for the divergence of Pengo. This dating range for

Sivapithecus does not, of course, incorporate the new dates

of 5.5 Myr from the Indian Siwaliks <Sankhyan 1985).

Based upon the molecular estimates and the currently

 85

accepted geologic time ranges for Sivapithecus <8-14 Myr if

Kenyapithecus is not included), Lewin <1983) estimates a

hominid-pongid divergence of > 8 Myr. While he has not

analyzed the fossils himself, Lewin <1983) draws his

conclusions based upon the findings of several

paleontologists including Walker, Andrews, and Pilbeam. If

Kenyapithecus is included in Sivapithecus, then the Pengo

divergence is pushed back to 17-18 Myr, and the

hominid-pongid divergence is increased to 10 Myr <Lewin

1983).
 86

Figure VII

Cladogram of a Pongo Linkage to Sivapithecus

Ciochon <1983:812)

Gibbons Ramapiths Or an tans African , a n s

~
\ Ap s /
\
\
\
\
\
\
\
\

/
 87

D. Ramapithecus and Sivapithecus as Sympatric Populations.

The fourth maJor hypothesis of Late Miocene fossil

hominoids proposes Ramapithecus and Sivapithecus as two

independent, co-existing hominoid groups, separable at

either the genus or species level. The coalfields of

Lufeng, Yunnan Province, China have yielded over 2,000

teeth, crania, and mandibles representing Ramapithecus,

Sivapithecus, and Gigantopithecus <Oxnard 1985:11). They

are dated at 8 Myr <Wu 1984; Oxnard 1985; Lieberman et gl.

1985). Of these fossils, measurements of over 900 specimens

have been statistically analyzed <Oxnard 1985; Wu and Oxnard

1983a, 1983b; Lieberman et £1. 1985; Oxnard et £1. 1985; Wu

and Xu 1985).

Multivariate analyses of the teeth result in defining

two distinct fossil groups. Pilbe~~ (1986:325> views this

distinction as a dimorphic expression of one population. Wu

and Oxnard <1983a, 1983b>, Oxnard <1985>, Lieberman et gl.

<1985), and Oxnard et gl. <1985> recognize more variation

between the two groups to allow for sexual dimorphism. In

fact, if they are accepted as the gender groups of one

species, the sexual dimorphism is larger than in any of the

extant apes <Oxnard 1985:24>.

The marked difference between

Ramapithecus <which is always smaller) and
Sivaplthecus is clear. These differences are
far greater than those generally found
between the sexes of such markedly dimorphic
species as gorillas and orangutans <Wu and
Oxnard 1983a:308>.

In this hypothesis. the labels Ramapithecus and

 88

Sivapithecus refer only to the Chinese fossils. The earlier

univariate statistical studies also indicate bimodal

distributions within both Ramapithecus and Sivapithecus.

This disclosure confirms the existence of two sympatric

hominoid species at Lufeng. The bimodal distributions

within each of the two groups, Ramapithecus and

Sivapithecus, can be interpreted in two ways: <1) as

species groups within the genera, or <2) as a result of

sexual dimorphism <Wu and Oxnard 1983b:259).

The degree of sexual dimorphism displayed in

Ramapithecus and Sivapithecus varies within each group.

Ramapithecus has a lower degree of sexual dimorphism than

Sivapithecus <Wu and Oxnard 1983a; Oxnard 1985; Oxnard et

~- 1985). Not only do they differ in degree, but

Raroapithecus and Sivapithecus do not show the same patterns

of sexual dimorphism. Sexual dimorphism is not measured by

size alone <Oxnard 1985; Lieberman ~ gl. 1985; Oxnard et

gl. 1985; Wu and Oxnard 1983a). Bimodal distributions occur

at 11 tooth positions in Ramapithecus and at 20 in

Sivaplthecus <Wu and Oxnard 1983a:318).

Ramapithecus/ bimodal patterns resemble those of Homo,

especially~ habilis and~ erectus <Oxnard 1985). The

bimodal distribution also indicates an equal sex ratio which

is comparible to humans. But Ramapithecus shows a lower

degree of dimorphism overall than Homo. The dimorphisms are

also in different tooth positions <Lieberman et gl. 1985).

Sivapithecus generally resembles the dimorphic patterns

 89

and sex ratios of the extant apes <Wu and Oxnard 1983a).

But each of the extant apes show individual patterns that

are unique <Oxnard 1985:14). The closest similarity is

between Sivapithecus and Pengo <Oxnard 1985).

The complexity of sexual dimorphism becomes apparent

when each of the hominoids displays different patterns. As

a result of these differences in species that are so closely

related, Lieberman et ~. <1985) suggest low sexual

dimorphism may be more primitive. The extreme dimorphisms

in the gorilla would be recently derived characteristics

<Oxnard et ~. 1985).

It does appear likely that smal 1 sexual

dimorphisms of means, dispersions, and
interjaw differences, together with equal
ratios between the sexes, are the generalized
situation or template, from which increased
selection pressures can mold increased sexual
dimorphisms of various kinds and increased
female to male rations. Other recent work
also suggests that sexual dimorphism is
complex and that greater sexual dimorphisms
are of more recent origin in homlnoids
<Lieberman et ~- 1985:321>.

Even though Ramapithecus bears similarities to Homo, it

is not classified as a hominid <Oxnard 1985:31>. If

Ramaplthecus were classified as a hominid, there would have

been a number of evolutionary reversals. There would be at

least two reversals starting with a change from ape-like

dimorphism patterns and 2:1 female to male sex ratios to

Ramaplthecus with human-like patterns and a sex ratio of

1:1. The first reversal would be evident in a 2:1 ratio and

ape-like patterns of Australopithecys. The second reversal

wouid be an adaptive change back to a 1:1 ratio and human

 90 ~ .

patterns for Homo. For the same reason, Ramapithecus cannot

be the last common ancestor of hominids and pongids

<Lieberman et sl· 1985:322). Nor can Sivapithecus be

ancestral to all of the pongids because of the uniqueness of

each hominoid/s fossil or extant, sexual dimorphic pattern.

Ramapithecus best flts into a radiation system leading

through~ habilis and~ erectus to~ sapiens with

Australopithecus as a related radiation not involved with

that of Homo. Wu and Oxnard <1983a:342) also support

~amapithecus as simply ancestral to humans.

In addition to the Homo radiation, there are others

leading to sister groups. Sivapithecus may have been an

early branch leading to Pongo <Oxnard 1985; Wu and Oxnard

1983a:342). Another may have led to the australopitheclnes.

The African apes would be the result of another radiation

that split again into lineages with specific sexual

dimorphic patterns. The initial radiation of Miocene

hominoids into Homo and pongid radiations is estimated at

close to 10 Myr.

Since Ramaplthecus and Sivapithecus from

China <not, it should be noted, the meager
remains of much earlier ramapithecines from
the rest of the world) were sympatric
approximately 8 million years ago, a date for
the human-African ape split at 10 million
years or even earlier is consistent with the
findings of this study: that Ramapithecus is
closely similar to hominlds but not to the
African pongids. It is possible that the
cladogenesls that led to Ramaplthecus was
preceded by the human-African ape divergence
<Lieberman et aL. 1985:322>.

The advantages of this hypothesis include few

 91

parallelisms and no multiple reversals of dimorphic

patterns. An australopithecine radiation does not

categorize Australopithecus into those species that are

dated prior to the establishment of Homo and those that are

sympatric. This proposal allows enough flexibility to

include older and younger australopithecines. It also has

the ability to assimilate any yet-to-be discovered older

Homo or other hominoids that are dated 8-3.5 Myr.

The disadvantages in this hypothesis are two-fold. The

divergence estimate of 10 Myr conflicts with the date of 4-5

Myr supported by the molecular clock <Sarich and Wilson

1977) and 4-8 Myr proposed by Cronin <1983:134). But if the

molecular dates can be modified to 10 Myr, they would be

compatible with the 8 Myr dates of the Chinese

ramapithecines <Lieberman et gl. 1985). This hypothesis

would also change the current ideas on the location of human

origins from Africa to China.

 Chapter IV

Criticisms of Current Ramapithecine Hypotheses

A. Early Divergence Hypothesis.

The e~t·Iy divergence hypothesis of hominid origins is

the proposal which has drawn the most criticism. The most

common controversy centers on the definition of what are

hominization trends and what are actually hominid traits.

Before the inclusion of Ramapithecus into Sivapithecus, it

was easy to attribute hominid affinities to Ramapithecus.

But after many scholars combined the two genera, the

ape-like characteristics of Sivapithecus did not fit the

hominid package.

The conclusions of Lipson and Pilbeam <1982), Pilbeam

<1986), and Wolpoff (1982, 1983) are not the same, but their

basic premises are. Lipson and Pilbeam <1982) emphasize the

affinities between Sivapithecus and Pongo, while Wolpoff

<1982,1983) and Pllbeam <1986) introduce the earliest

hominid as a sivapithecine. But they all agree that

Sivapithecus was not a hominid. Their basic premise lies in

the assumption that Sivapithecus underwent an adaptive

radiation in which an Asian branch led to Pongo. Wolpoff

<1982,1983) and Pilbeam <1986) extend this hypothesis to

include a second, African radiation leading to the African

apes and~ afarensis. Lipson and Pilbeam (1982) and

Wolpoff <1982, 1983:663) agree that at least four of the

Sivapithecus traits considered to be shared and derived with

Homo by the early divergence hypothesis are in reality

92
 93

primitive. These traits are thick molar enamel, the degree

of maxillary prognathism, basic molar morphology, and

maxillary incisor heteromorphy. Many of the traits that are

considered hominid-like in Sivapithecus and used as evidence

by supporters of an early divergence are viewed as ape-like

in ~ afarensis

Greenfield <1980) also questions the validity of

characteristics defined as hominid traits in the early

divergence hypothesis. It is his opinion that "many unique

and likely irreversible human dental adaptations did not

evolve until after the Miocene, and that many

dryopitheclne-llke characteristics persisted in the human

lineage in the Pliocene" <Greenfield 1980:354).

Ward and Pilbeam <1983) and Pilbeam <1984a) consider

such traits as thick enamel, rotated canines, megadont

molars, and robust maxillae to be the result of parallel

evolution rather than being the synapomorphies defined by

Kay and Simons <1983). Pilbeam <1984a) uses facial features

and the palate of Sivapithecus to support his statements.

His analysis shows these features closely resemble those of

Pengo. But the same features of Sivapithecus are not at all

like those of~ afarensis, while the aforementioned dental

characteristics are very much alike between them. If the

traits common to Sivapithecus and Pengo are shared derived

characteristics that post-date the Asian and African

radiations, then parallelism can be documented for the

dental features <Ward and Pilbeam 1983:236).

 94

Whether these scholars cite specific fossil evidence to

refute hominid similarities or disagree on principle, the

consensus is that Sivapithecus was not a hominid. The

majority of sivapithecine characteristics are seen either in

Pengo, the extant African apes and/or AustraloPithecus.

Characteristics that were once considered to be shared

derived traits of Sivapithecus and Australopithecus are

currently being redefined as either the result of reversals,

parallel evolution, or retention from a generalized hominoid

stock. If the sivapithecines and australopithecines are not

linked by shared derived characteristics, then Sivapithecus

potentially has a phylogenetic link to the living extant

apes and Homo.

 95

B. Late Divergence Hypothesis.

Except for supporters of the early divergence

hypothesis, most criticisms of the late divergence

hypothesis as presented by Greenfield <1980,1983) concern

the identity of the last common ancestor. Again, part of

the discussion centers around what type of characteristics

are studied. If the traits that link Sivapithecus to Pengo

are primitive or general in nature. then it follows that

Pengo may have diverged from Sivapithecus. But if these

traits are specialized and unique, then Sivapithecus is

exclusively ancestral to Pengo and could not have been the

last common ancestor to Homo and the African apes <Lewin

1983). Lipson and Pilbeam <1982:547) also proposed that it

is not likely that Sivapithecus is ancestral to all of the

great apes, and that some of the ramamorph characteristics

are uniquely shared with Pengo.

If Sivapithecus is the last common ancestor of hominids

and pongids, then the following requirements would have to

be met <Kay and Simons 1983): (1) Sivapithecus would

exhibit basic elements of synapomorphies common to hominlds

and extant great apes. (2) Sivapithecus would not share any

derived features in common with any individual member of the

hominid-extant ape clade. Kay and Simons <1983) deny that

Greenfield~s <1980) hypothesis has satisified either of

these two requirements in his hypothesis. Here again,

non-agreement as to the type of characteristics important

for analysis is the central point of the discussion between

 96

the early and late divergence hypotheses.

 ,.
97

C. Sivapithecus (s an Ancestor to Pengo.

Kay and Simons <1983) use the same arguments to

criticize a Sivapithecus- Pengo ancestral relationship as

they do the late divergence hypothesis. These workers

present arguments against every trait that Andrews and

Tekkaya <1980) and Andrews <1982) claim is a shared derived

characteristic between~ meteai and Pengo. These traits

are labeled as primitive and generalized, not viable as a

basis tor an alternative hypothesis.

Shea (1985) analyzes the craniofacial morphology

between Pengo and the African apes. His study shows that

the greatest differences lie in the internal cranial

anatomy. But externally, Pan, Gorilla, and Pengo are

virtually the same. If the angle of the endocranial base is

shifted, then apart from size, Pengo and Gorilla are shown

to be alike.

In sum, a number of internal and

external features of the orangutan skull
appear to be related to the position of the
face on the braincase. These features should
thus be treated as an interrelated complex
rather than as a series of independant
characters <Shea 1985:338).

Shea <1985) does not view the skull morphology of Pengo

as derived from the general hominoid pattern, but rather as

an example of the primitive complex itself. Oxnard~~-

<1985:149) also postulate that the dental characteristics of

the African apes may be recent specializations. They also

hypothesize that some characteristics shared by orangutans

and modern humans are not the result of paral lellsm, as Kay
 98

and Simons <1983) assume, but are expressions of a basic

hominoid complex. This hypothesis does not support an

exclusive relationship between Sivapithecus and Pengo based

upon shared derived dental and cranial characteristics. It

also refutes Sivapithecus as a hominid. The same traits

that disallow Sivapithecus as an exclusive ancestor of Pengo

cannot be proven to be shared derived characteristics with

Australopithecus if they are proven to be part of the common

hominoid stock.

Von Koenigswald/s <1983:523) only criticism of Andrews

and Tekkaya <1980) is based on dietary considerations.

Modern orangutans have a diet of soft foods that result in

fine wrinkles in their dentition. He has observed that

Sivapithecus does not exhibit these wrinkles and their

absence indicates a diet of coarse food. This in turn leads

von Koenigswald <1983) to suggest that Sivapithecus lived in

a different habitat than that occupied by the modern

orangutan. This observation indicates an adaptative change

between Sivapithecus and Pengo.

Greenfield/s (1980:364) answer to questions posed by

supporters of an exclusive Pongo relationship to

Sivapithecus is simple. Because he specifically proposed

Sivapithecus as the last common ancestor based upon general

characteristics, the late divergence hypothesis is not

falsified. In any case, even if Sivapithecus is

disqualified as the last common ancestor, the hominid-pongid

divergence based upon Pengo affinities to Sivapithecus fal Is

 99

within Greenfield/s <1983) own estimates of 5-10 Myr.

Pilbeam <1984b) addresses his comments to the study of

fossil populations in general.

I think we too have to pursue a pointilliste

reconstruction of paleoanthropologlcal
narratives, in the sense that we should
generally expect to be satisfied with broad
rather than specific insights. We dare not
peer too closely, because if we do the
picture wil 1 dissolve <Pilbeam 1984b:19).

Pilbeam <1984b) feels that two changes are needed in

the paleontological approach to fossil schematics. (1) The

fossil population should be analyzed with a biological

perspective as if it was a modern functioning species. <2)

The fossil species should be viewed on its own terms and

emphasize the differences between fossil populations.

Currently the trend is to try to categorize a fossil species

where it best fits, depending upon the current definition of

what Is "hominid", "pongid 11 •

11
dryopl thecine .. , or "Homo 11 •
 Chapter V

Conclusion

In the previous chapters, the following maJor

divergence hypotheses were presented: C1) early divergence

CMiddle Miocene) of hominids and pongids, C2> late

divergence CLate Miocene) of hominids and pongids, and C3)

two Pengo ancestry hypotheses for the phylogenetic status of

Sivapithecus / Ramapithecus. The early divergence and Pengo

ancestry proposals have maJor problems.

The early divergence hypothesis insists upon the

hominid or exclusively hominid relationship of Sivapithecus.

This narrow view does not allow for a thorough analysis of

anomalous ape-like characteristics also present in

Sivapithecus. Some of these traits are: C1) narrow

interorbital septum, <2) deep zygomatic region with a strong

lateral flare, <3) broad nose, <4) broad I1 significantly

larger than I~. <5) marked alveolar prognathism, and (6)

short upper face. The incorporation of Ramapithecus into

Sivapithecus further weakens this hypothesis. If

Ramapithecus alone resembles the hominids, and Sivapithecus

is reminsicent of the pongids, then it makes more sense to

keep the two genera separate. But Kay and Simons (1983)

deliberately join them, claiming that all of the

hominid-like traits are uniquely shared, derived

characteristics and the pongid-like traits are para! lelisms.

It would make more sense to me for Kay and Simons (1983) to

retain Ramapithecus as a separate genus CWolpoff 1983:663)

100
 101

because, by trying to emphasize a special relationship of

Ramapithecus with the hominids. they are only establishing

its common affinities ~.;ith modern hominoids. The conclusion

of this study proposes that Sivapithecus is not a hominid.

The problems with some of the Pengo ancestry hypotheses

are the same as those with the early divergence views.

Instead of the hominid-like traits being considered

specialized, it is the Pengo similarities that are viewed as

unique. But if the sivapithecine traits are considered

unique to Pengo, then Sivapithecus could not have been

ancestral to any of the other hominids. Any traits found in

common with the African apes or Homo are labeled as

parallelisms no matter how many times they may occur. For

example, a trait such as a dental / gnathic complex for

powerful mastication would have to have been developed

independently several times.

A variation of the late divergence hypothesis as a

valid response to these problems seems to be in order. If

the 5.5 Myr <Sankhyan 1985) date for Haritalyangar, India,

is accepted, then the Sivapithecus / Ramapithecus species

complex, excluding Kenyapithecus, spanned at least 5 million

years in hominoid evolution, beginning approximately 10-11

Myr. When Kenyapithcus is classified as a sivapithecine,

the Buluk, Kenya, date of 17.2 Myr <McDougal 1 and Watkins

1985) increases the time span to 10-12 Myr. Suggestions

have been made that the 11

higher" organisms evolve faster

than more "primitive" forms. In hominoid evolution, the

 102

duration for the transformation between one genetic

population and another is estimated at approximately 2

mil lion years <Cronin 1983; Ciochon 1983). Considering this

transition time. even a minimum of 5 million years is a long

span of time for a hominoid to remain unchanged. Even so.

many of the current hypotheses appear to adjust the

hominid-pongid divergence estimates to either just prior to

or after the Sivapithecus time span. In the case of the

Pengo ancestry hypothesis. the divergence occurs in a

nebulous time span between the Pengo divergence and the

first appearance of ~ afarensis.

Based on these premises, adjustments to the late

divergence hypothesis can be proposed. If Sivapithecus is

not a hominid, the divergence of the Hominidae could have

been as late as 5.5 Myr assuming the Lothagam, Kenya, fossil

is classified as a hominid and dated at 5.5 Myr <Patterson

et sl. 1970) When the 1-2 million-year underestimation

factor is taken into account, the hominid divergence based

on the Lothagam fossil is 6.5-7.5 Myr. At Haritalyangar,

India. Sankhyan (1985) has dated Sivapithecus at 5.5 Myr.

The hominids may have evolved from only one sivapithecine

species. The pongids probably evolved from one or more

sivapithecine species, depending upon whether Pan and

Gorilla diverged from each other or concurrently from the

basic hominoid stock. Pengo probably diverged from an Asian

sivapithecine earlier than did Pan and Gorilla. Instead of

a hominid-pongid split at the end of the sivapithecine time

 103

span, the divergence may have occurred as species evolved

within the sivapithecine adaptive radiation. In other

words, the actual divergence could have occurred within the

Sivapithecus radiation. The diverging sivapithecines are

considered sister species until they evolve into distinct

genera. If Sivapithecus phylogeny is viewed as a continuum,

then it is easier to postulate the hominid-pongid divergence

as occurring within the sivapithecine adaptive radiation.

If the phylogenies are viewed as flowing from ancestor to

descendant lineages, then the intermediate forms will

overlap the boundaries of both. It is conceivable that the

variations between Ramapithecus and Sivapithecus are

evidence of this radiation process. Ramapithecus is usually

considered more hominid-like than Sivapithecus. But

Ramapithecus is so close to Sivapithecus morphologically

that they are often combined into the same genus. The

emergence of RamaPithecus may be part of the adaptatlve

radiation process of Slvapithecus towards the Hominidae.

The differences between the later, 8 Myr date for

Ramapithecus in the Siwaliks <Tauxe 1979), and the earlier

date of 8-17.2 Myr <Wu 1984; McDougall and Watkins 1985) for

Sivapithecus in China and Kenya, respectively, may be the

divergence of hominids and pongids.

Unfortunately, at this time, there is not enough

evidence to isolate any specific sivapithecine species as

ancestral to any individual subsequent hominoid species.

However, Wolpoff <1983) postulates a Pengo/ African ape

 104

split occurring in the Asian geographic radiation from the

Eurasian stock. and an African radiation resulting in the

Homo/ African ape divergence.

As early as 1951, Washburn had this same view. He felt

that the ancestors of hominids and extant pongids were

members of different radiations and adapted to their

environments accordingly. The direct ancestor would have to

be specialized, not generalized. Ciochon (1983:819) also

emphasizes the specialization of direct ancestors when he

states ~it may not have been development of new

characteristics that marked the adaptive shift of the

earliest hominids, but rather an emphasis on specialization

of an already existing suite of features".

Simons (1963:887) emphasized the transitional nature of

RamaPithecus. In 1963, he firmly stated the Ramapithecus

fossils could not be attributed to either the Pongidae or

Hominidae. His early views are more consistent with the

radiation/ divergence hypothesis than his later proposal,

which classifies Sivapithecus as a hominid <Kay and Simons

1983).

Some of the ramifications of this radiation /

divergence hypothesis are: (1) The pongid divergence is

actually a two-point split. Pengo left the major hominoid

stock earlier than did the hominoids which diverged into

Homo and the African apes. This premise is now commonly

accepted. C2) If Sivapithecus is related to Pengo, then

traits common to both genera are generalized from the basic

 105

hominoid stock. <3) Only one ramapithecine / sivapithecine

lineage could be ancestral to Pengo. It would also be an

Asian species. <4) If Sivapithecus and Ramapithecus are

distinct at the genus level, as proposed by Oxnard ~ gl.

(1985), then Ramapithecus may be the most recent genus to

diverge from the basic hominoid stock. But as Lieberman et

gl (1985) point out, it is unlikely that any fossil group

will ever be specified as an actual ancestor to the

hominids.

It is irrelevant to ask whether or not

this particular Ramapithecus is ancestral to
humans; indeed, it is irrelevant to ask
whether any specific fossil of this
approximate age is ancestral to an extant
form. The statistical chances involved in
the accidents of fossilization almost always
preclude any specific form from being such an
ancestor <Lieberman et gl. 1985:322).

<5) If Sivapithecus and Ramapithecus are separate

genera, then they are still representative of the

sivapithecine adaptative radiation. <6> The earliest

hominid was a special form of ramapithecine. But

Sivapithecus / Ramapithecus was not a hominid <Wolpoff

1983).

Allowing for the approximately 2 million years

estimated to be needed for a lineage to diverge and

distinguish itself from the preceding genus, the

hominid-pongid split should be 7.5-10 Myr. The

Haritalangyar, India, and Lothagam, Kenya, fossils are dated

at 5.5 Myr. But they are the only sivapithecines and

australopithecines dated 5.5-8 Myr. The fossil record is

 106

too sparse to either disregard or accept this date. The

underestimation variable should be added to both the 5.5 Myr

and 8 Myr dates resulting in a divergence estimate of 7.5-10

Myr.
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