Palaeoecology of the Miocene fauna from Paşalar, Turkey

Peter Andrews

Palaeoecology of the Miocene fauna from Pagalar, Turkey The environment ofdeposition of the Pa&x Miocene sediments indicates rapid deposition from sources close to the fossil site. There is a widt six range ofbones, from insectivorr treth andjaw to proboscidean limb hones. and there is some evidence of taphonomic bias during the accumulation of the bones. Mammals constitute the main part of the fauna. and palaeoecological interpretations are therefore hased on them. The high diversity of mammalian species (58 specks known) is sufficient to suggest some form ofwoodrd hahitat in a tropical or suh-tropical seasonal climate. The size distribution of the Pagalar fauna and its communit) structure confirm this, whik excluding the possibility of any of the tropical habitats with evergreen or semi-cverqreen forest vegetation. The closest similarities in all of the analyws arc with suh-tropical forest faunas from India, with seasonal, summer-rainfall climates and semi-deciduous forrst corer. dense ground vegetation and frequent open glades. ;Uso similar are the deciduous woodland (or forest) faunas from tropical Africa, and these also occur in highly seasonal climatic regions and have similarly simple tree canopies and abundant ground vegetation. It is concluded. therrforr. that the climate at Pagalar during the Miocene M~S seasonal, sub-tropical, and perhaps with summer-rainfall rains; and the \,cgrtation consisted ofsemideciduous forest with abundant ground cover and many open areas.

Recei\ ed 12April I989 Rrvision received 23 .July 1989 and accepted I August 1989

Palaeocolog> ,

Kqwords: hliocrnc. Struct~lrC

mammals,

communit)

qfffmnn f+olurm ( 1990) 19,569-.%X

Journnl

Introduction The

middle

varied

Miocene

deposits

mammalian

fauna.

and only a few lower based

on

the

palaeoecology,

basis

of correlations

of the fauna with

& Bernor,

Before

making

its composition.

the

geological

time

living

taken

in more

of

The

geological

The

ecological

bones

than

one

setting

habitat

or their

an extremely

been

found,

analysis

described (Flynn

and

cases

papers

in this

to be about

& Jacobs,

to be

in some

in the earlier

is estimated

rich and

no invertebrates has perforce

taxonomy,

that

of a fossil fauna, or taphonomic criteria make

in the source

to accumulate

accumulation of bones potential bias.

been fauna

assessment ecological

complexity

Their

concensus

Miocene

any

the

yielded

have

15Ma

on the

1990; Gentry,

1990:

1990).

any ecological

accumulation

and/or

other

and

have

by genera1

1990; Sen,

of the fauna

altered the

date

have

remains

et al., 1989).

of their

The

in Turkey

or plant

and so the palaeoecological

(Alpagut

issue.

source

vertebrates,

mammals

interpretations

Fortelius

at PaTalar

No pollen

the

bones;

type. transport

of the Pa$alar

likely up

it is necessary

to have

the

of the bones;

and

derivation

sediments

site constitute

shows

that

are:

ecological

of the

they

may

important

fauna

modifications

to the burial

which

been

Pasalar

area

Taphonomic

to consider

modifications

bones

the have

during

topographic

change from

affecting

during animals

either

a second

the

source

accumulated

of

rapidly

from the hill slopes to the west and south of the fossil site (Andrew & Alpagut, 1990). This is the location today of the main outcrops of Pagalar marbles and Kiziltepe metamorphic rocks which

are the main

source

0047.-2484/90/040569 + 14 $03.00/O

rocks for the fossiliferous

sediments.

They

outcrop

@ 1990 Academic

Press

along t.imited

a

belt up to 2 to 3 km from the site. ‘l’aphononnic the

Pagalar

fauna1

(Andrews

CG E:rso).,

and topograph!, dipping

(30-40

degree)

of differences

between

diKerences

occurred

types

possible

Topographic showed

minor

zones several

basin

from

part

not have There

during

extended

the Miocene,

composition

but

short.

assemblage emplacement (Andrews

to accumulate are

which

provided short,

& Alpagut,

a period

because

of weathering,

observed

would ecological

the

not be achieved.

assemblage

on the lower

been

of each

of the

granitic

1990). of the than

The

rocks

sediments

mountains,

and

they do today,

any conceivable

differences

insuffIcient

‘I’his

is reason

the edge of the Gonen

slopes

under

other.

but there

of intrusive

in the Miocene

habitat

in their

geographicall)

the accumulation

forms

Sr Alpagut,

of the mountains

source

the)

model

due to topographic

to have

fauna1 assemblage

to this:

the

of the

bones,

at their

present

time

greatly

ofweathering longer

burial

to have

and

In ecological been

the

of

relief

affected

tht

both

to

interpretation before

transport

the lack ofweathering alteration

deteriorate and

and

is considered

beyond

the spectrum these

periods

of

of the bones

all

recognition of weathering

are too short

for

likely.

from this discussion

that

the likelihood

of the Pagalar

is not high. The short time ofaccumulation source area of both sediment and bones the preservation bias, including

consists

of the death

of transport

of the bones

the bones terms,

likely to have

on the geological

pedogenic

shorter

time

site. The latter

of bone surfaces,

and other

by burial;

is considered

of accumulation

and

than a few days based

of root marks

of years:

derived from mixed habitats geographical extent of the

so that

during

is composed

extent

have

unprotected

ecological variation during some degree of taphonomic

distance

that

zones

are

of’Pagalar,

but the time of accumulation

The degree

change

factor massif

(Andrews

the Pagalar the

1990),

of tens

It may be concluded

a significant

components

no more

and the low frequency

suggest

major

two

of the sediment/bones

must have been longer. teeth,

would

zones

(1964) because

fauna.

There

have been extremely

a short

to latitudinal

in the vicinity

have been minor

they

of the PaSalar

The time taken

within

rocks

to the tops

may therefore

species

occur

a greater

similar

communit\

Simpson

altitudinal

accumulated,

metamorphic

have covered

that

in two plant

are analogous

of the mountain

Kiziltepc

The full scalr

variation.

the

may

the

of mammalian

Because

to the sediments

the

they must

diversities

on the mountains

the Pa%alar sediments

derived

erosion.

been

major

higher

while

fauna.

to ecological

of a mountain

types

today

did not contribute

would

have

\.egt*tation

the site as stccpl)

but it is likely

resulting

sedinienls

influence

near

ofmoisture.

assessed,

arc at present, rise

;ts the

vegetation, retention

in mammalian

vegetation.

habitat

The

in which

although

has not been

this may not have been

fauna.

which were

that

regions

and

effect may be observed Pagalar

with better

give

source

are exposed

a lime-tolerant

also

that the bone comprising

same

two rock types

with

differences can

these

plateaux

habitats

on the slopes

climate

condensed,

these

the

the marbles

in the past as there

mountainous

the altitudinal on

slopes

suggests

from

day,

extent:

sloping

complesity

that

to believe

dip

cvidcnce

derived

1990). At the present

form gently

with

was

to a considerable

metamorphics

effects

assemblage

largely

fauna

ofthe Pagalar fauna. There has, however, loss of almost all weak or easily destroyed of isolated

teeth,

some

being

and the restricted limit the possible

mandibles

and

been bone, robust

postcranial elements. This degree of destruction does not appear to have had a differential effect on either the size distribution or the taxonomic composition of the fauna except for the loss of some of the smallest size category of mammals. Some predator activity is indicated by modifications on the bones, and this may have produced the bias against

PA$ALAR

small

mammals,

brought

about

hut it is not possible the high

degree

to say whether

PaSalar

compiled Forstrn, $en,

fauna

from

includes

made

although

this aspect

individuals

at this stage

(MNI)

or lower,

.Indrews

& Ersoy

specimen

and

The

fragmentary

‘I‘hc)~ appear

have the thick-enamelled diet.

Four

most

common

woodlands.

are glirids

with

dominated

by forms with low-crowned

the small

mammal

canoids.

omnivorous

forms

deinotherc

hrrbiage,

while

teeth with smooth

high

have

number

been described of species,

(hyaenids),

omnivorous

The

(Flynn

rounded diets.

is a small the larger

are present species

with

gomphothere

diet.

1976). There

is also a single

it includes

1990a).

the

moist sprcics,

Sciurids,

1990; i’lnay, I D9Ob). the rodent farina is

cusps.

Criretodontids

in

up about

(Schmidt-Kittler, primary

hall’

1976).

predators

(viverrids)

is surprisingly

(ltilids)

and

high, especially

Both

at Pagalar,

a gomphothere

bilophodont

teeth

more

of the mort

has massively

can be said species

to hale

(MN1 = 21, Forsten, 1990), and it has browsing rather than grazing adaptations. abundant grazing

of the

rhinocerotids,

adaptations

adapted

thick-enamelled browsing

of hyracoid

hydracoid-like Chalicotheres Begertherium,

(Fortelius,

and a deinotherc.

apparently

1990a).

teeth,

adaptations

with low-crowned

groups of perissodactyl most of which were also apparently quid .Inchitherium is the next most common large herbivore

with

1990).

like the canoids.

of proboscidean

for a tougher

most

the)

inhabitin,q

and they make

from Payalar

and

of carnivores

(GaAry,

probably

1977).

frugivorous

&
clnay.

small-mammal-predators

The diversity

adapted

Tht,

& ‘l’obien.

forms

&.Jacobs, rare, and

are comparativeI>,

likely to have had frqi\,orous

of carnivore

carnivore/scavengers

species

1978;

bird

and dentall>-

1990), and of the 13 rodent

(Tobien.

1 oi’

a single

fauna.

species

is a relatively

Two

to living

or right.

in ‘I‘ablc

to a hard object

1980; Flynn

related

of

based

will be mentioned

skeleton

are also present teeth

l‘ound which (Andrews

are prcxsent ($rn,

abundant

hypsodont

are considered

This

most

although

Pagalar

(Engesser,

ofleft

to mammals,

related

groups

account

postcranial

described

of Schi~o,&+s

(taking

elemc’nts, numbers

taxonomic are listed

from

in their

Minimum

the MNI’s been

is probable

and spalacids

strongly

particular Sistecn

been

have

described

of lagomorph

the

is restricted

which

is a species

Two species

ctcnodactylids, Spe[+cs

have

of which

crictstodontids

been

adaptation

insectivores

available,

adaptations

1990:

reconstruction

of the fauna1

in the text.

type for any taxon

remains

1990;

1990; Girbiiz.

only, and no correction

abundance

for some but not all ofthe

analysis

have

to have arboreal

records

has been

& Jacobs,

for the palaeoccological

in relative

\Vhere

reptilian

where appropriate. Two species of primate

or transport

Pagalar

(Flynn

1990; Gentry,

as appropriate

of any one tooth

(1990).

& Bernor.

1 give presence-absence

or juvenile).

list from

of the ,journal

the basis

have been calculated adult

‘The f:tunal

issue

for differences

will be mentioned

on the
in this

19900, b; Fortelius

here. The data in Table

has been

weathering

of the Paqalar fauna

of mammal.

papers

1990a, b), and it forms

1990; LInay.

attempted

58 species

the preceding

1990; Fortelius,

predation,

I

of breakage.

The species composition The

.)I

PAt.EOECOI.OG\

fiJr_ soft

presumabl!, of their

teeth

teeth

and three

adapted for browsing. ‘l’he after the bovids and girafids teeth,

low

are extremely has

relative]>,

‘I-he other

crowned rare

and

hypsodont

two rhinos

with

at PaSalar. teeth. wer(’ less

Table

1

Pa*alar

,Sioapithecus darcoini ,Sioapithetus sp. Soricid species Desmanodon cl: minor .Schi;ogalerix pasalarensis Erinaceinae Prolagus oeningensis dlloptos anatoliensis Sciuridae indet. Spermophilinus bredai l‘amiini cf. Eutamias cS. Paleosciurus Alicrodrvom_vs sp cl: hf. miocaenicu Peridrvomvs Sc+rms sp. cf. S. iintermedius Turkomv’s pasalarensis ,Ile~gacricetodon Democricetodon Pliospalax marmarensis Chalicomvs jaegeri .4mphicvon CC mcjor Hemic:von sansaniensis Plithocyon I~rsauus cf. primaevus ITrsavus aK, intermedius Plesiogulo Proputorius Trochictis Lutrinae Protictitherium intermedium protictitherium aff. gaillardi Protictitherium new species Percrocuta miocenica Percrocuta new species Pseudailurus cr. quadridentatus Pseudailurus larteti Orvcteropus Deinotherium giganleum Gomphotherium angustidens Pliohvras Anchitherium aurelianense Begertherium tekkavai Aceratherium aff tetradaclvlum Brachvpotherium brachypus Chalicotherium grande Listriodon cf. splendens Listriodon sp Conohyus simorrensv Taucanamo inonuensis Dorcatherium Palaeomeryyx hlicromervx Stephanocemas Hispanomeryx Ciraffokeryx aff punjabiensis Caprotragoides stehlini fivpsodontus pronaticornis

fauna1 list

PA$ALAR

common,

one

having

having

typical

There

are four

omnivorous

an apparent

browsing species

mammals

because

of uncertainties

bilophodont species

mixed

of suoid One

at Payalar,

of the

in the Paaalar

teeth

adaptation,

all of which Listrioidon

suoids,

fauna,

anterior

adapted

with

for a folivorous

has more bunodont

were

and

small

splendens,

the othn

molars

apparently

better

another

adapted

most

its MN1

Listriodon

teeth.

but

the

to calculate

posterior diet,

to medium-sized

is amongst

but it has not been possible

in .associating

apparently

at Pagalar

grazing/browsing

morphology.

herbivores.

common

.5 73

PALAEOECOLOC\

closely

has

related

for an omnivorous

diet. The common as abundant also

bovid

relatively

(Gentry,

1970,

compared

with

taken

to indicate

this

and

1987). and

The

(see Table

this

all

less

of the three

the presence contemporary

European

Methods The

Pagalar

faunas.

mammalian

These

together

fauna

include

with an additional

including

several

from

comparisons

with faunas

and

Asia,

central

coniferous

but

forests,

monsoon

of India

and

I. Semi-deciduous, main

canopy,

meadows

2. Deciduous et ul.. 1979). dominated

forests, East

forests

some

Asia.

(1990) than

of recent

and southern from

habitats,

original

et al. (1979)

from all over Eurasia. has been placed on

faunas data

India,

with summer

winter

for some

by one or a few tree species,

vegetation.

The

has wtlrc

comparative

by Andrews

of central

tundara

The

over the entire trees

and the subtropical for this

comparative

rainfall

with

the four

(monsoon)

7-8 months.

There

a discontinuous

are interspersed

Europe

impoverished

is a

lower tree

extensive

herb-

and

1967).

in tropical

It is highly

range

described

also includes

forest in sub-tropical

(Schaller,

forest,

deciduous

boreal

dominated

and rich ground

grass-rich

to a wide

faunas

sample

South

extending

tree canopy

Gentry

elsewhere, but for reasons of space the faunas from to the Pagalar fauna are illustrated here, as follows:

seasonal is long,

cervids,

individuals

conditions

from forest and steppe habitats 1984). Emphasis (Artemiou,

the comparative

will be presented most comparable

The dry season single

Turkey

the

22

of analysis

African

from the richer

the northern

forests

sample habitats

28 faunas

open

is

for grazing

but

1990),

species.

more

twice

sites.

will be compared

the 23 tropical

abundant,

bovoid

of drier,

than

adapted

(Gentry,

common

at Pagalar

apparently

is also

common

is more

1990). f@sodontus

& Ersoy.

form

GiruJ&eyx

are

181 individuals

1 in Andrews

is a hypsodont

giraffid

tragulids

in, for instance,

is Cuprotrugoides stehleni, which

at PaHalar

species

abundant,

palaeomerycids

presrnt

species

as any other

Africa

seasonal

usually

with

by one or a few species,

referred

to as tropical

a long dry season.

no lower

canopy

There

but grass-

woodland

is a single and

(Andrews tree canopy,

shrub-rich

ground

vegetation. 3. Deciduous with

temperate

a wet winter

trees at lower altitudes and herb-rich ground 4. Semi-deciduous season.

The forest

forest

in southern

and hot dry summer. and coniferous vegetation.

tropical

rain

is multi-canopied,

Europe, It is often

trees at higher

forest

in Africa,

with

moderate

extending zoned

eastwards

altitudinally,

altitudes; seasonal,

there with

to the Caucasus, with

broad-leaved

is a single

tree canopy

a moderately

long

to high tree species

diversity,

dry

greater

than

any

of the

evergreen

rain

‘1’11~Pasalar simplest crude,

of habitat

variations

ecological

complexity

into

organisms. account

sources,

span

source

area

many

(equitability)

some

within

include

groups,

the major within

the range

with

size is estimated range

of recent

on postcranial

on proportions. habitats species

diversity

and

stability

within

disjunct

regions

animals

occupying

&mated, ephemeral ecological

trophic

strategy

guilds

(Harrison,

than

through

a given

habitat

geology,

soils,

Previous

comparisons

palaeoecotogical during

is more

of individual

of the communities

species provides

of the individual

1966)

and

of the habitat

is or

across

time

with

and

geographicall!, between

lasting place,

to the nature their

of

of the

itself (as controlled

at a given species

patterns

relationship

and longer

guide

from differing as a result

distinctive a better

adaptation morphology

Consistent

habitats

Rod)

area for a wide

on molar

types.

of body

than and

by the

so the

of the f:Ltuna

living

counterparts.

woodland

to

of Paqalar palaeoecology

interpretations

the Miocene

molar

et al., 1979). ‘I’h is adaptive

etc.)

the

basis.

in communities

for similar

(Olson,

in terms

1987). I ,ocomotor is based

strategies

and the structure

Analysis

conditions

time

siztxs

and between

on an empirical

the habitat

are found

1962; Andrews

presence/absence structure

of these

distinguish

in an!’

but the relative within

on second

adaptation

and

changes

from almost

communities

types

rt al., 1979; Legendre,

and

adaptive

topography,

and its ecology

in mammalian

and dietary

either

1987).

of body weight

proportions

to categorize

and ruminants,

from quite

diversit).,

all faunas

or

diversities

1987). Faunas

of size change

to habitat

ma)’ in the

by Order

of species

ma): reflect

(Legendre,

take mixed

change

of the fauna

For example,

habitat

from

Fossil faunas

tasonomic

g-roups

carnivores

and other

richness.

distribution

of relative

or both.

and

(Andrews

The relative

species

greater

must

of the fauna

grouping

of the

tasonomic

present

morphology,

Ibr mammals

accumulated.

1973;

and

give rise to ecological high

patterns

Fleming,

for fossil faunas

et al., 1979; Legendrc.

type can be related

mammals

are used

would

the taxonomic

on the basis of regressions

based

both

as derivation

and the degree

climate

1964;

if r.athc.r correlated

variability,

of richness

the fossil fauna

rodents,

of adaptations

size, diet and locomotor

richness,

such

order

groups,

(Simpson,

topographic

species

regime,

groups

these

are correlated

Finally,

the higher

or climatic

to be a powerftd, with differences

in terms

similar

structure,

found

ecology

;\t III{.

(in the sc’nsc

l975),

a misleadingly

may

in size within

habitat

hence

te\~ls.

has been

correlates

which

(Andrews

variations

of the species

of years,

have

01’ \~t~tt~~r

1964; May, greater

e&cts,

the groups

habitats

community

diversity

and

these

information

disparate

di\,ersity

at three

richness

level of fauna1 analysis,

provides

spccics

ofovcratl

of time over which and

tr‘ec

compared

latitudes,

taphonomic

of the fossils

have

(Simpson,

about

thousands

the

been

all lead to increased

possible

than

gives a measure

topography

Inferences

At the second Family

Species richness

Lower

and the length

easily

fhunas

ofspecies

in latitude,

et al., 1979).

les4

PI (I/.. 1979).

and recent

or heterogeneity).

Andrews living

(Andrews

fossil fauna

measure

hul

examples

types

level, the total number

of richness with

preceding

forest

have

of Pagalar

indicated

(Becker-Platen

dry et al.,

1975).

steppe

Schmidt-Kittter

(1976) considered that the environment may have been even more steppe-like of hyaenids in the fauna, but he had difficulty reconciling this opinion with

on the basis his opposing

view

carnivores

that

the

high

diversity

of canoid

(=

Arctoidea

of Schmidt-Kittler)

is

PASALAR

Table 2

PAI.AEOECOL0G-a

Species diversities

.)

of mammalian

faunas

Number of Number (:limatic

tape

Vegetation

Ckmiferous

temperate

\Varm

zone

specks

forest

Coniferous forest Deciduous forest hlixed forest Swampy deciduous woodland Stcppc-woodland steppe

temperate

Deciduous dry forest Semi-evergwen lbrest c\er,green forest

Subtropical

‘Tropical

Africa

Grassland Scmi-arid bush Deciduous woodland Deciduous forest Semi-deciduous forest Ecrrgreen forest hlontane forest

‘Tropical

Asia

Evergreen hlontane

deciduous forest

I)ata for tropical African faunas are I>tbers are from Andrews (in preparation).

indicative

of closed

forest

from the systematic and

of

samples

Iksert

Tundra Cold

/5

conditions

papers

forest

from

Andrcus

(Schmidt-Kittler,

and

1976). Conclusions

in this issue have generally

this will be commented

rl ol. (1979),

stressed

about

the openness

Pagalar

of the habitat,

on in its place.

Species diuersit_y The

analysis

species

presented

richness

here

of the taphonomy

(Andrews

and preservational

factors.

modern most

habitats

in Table

fossil faunas,

faunas.)

not exceed

savanna cast

Pazalar

fauna. are

2. (Because

fauna,

with

The

Pagalar

tropical

interpretations fauna and

species

most

fauna

richness

steppe

faunas,

have

comparable faunas

for a variety

from Pagalar,

of

as well as from

of modern

comparative

greater

than is present

from warm is also higher

temperate than

both

climates tropical

and this of itself is sufficient

indicated

to the Pagalar from

level of

by depositional

richness

is considerably

forest

which

subtropical

of species of bats

1. This

but from the discussion

from all the analyses

species

latitude

in Table

to be little altered

ranges

of the absence

excluded

and middle

The extant the

been

recorded

day habitats,

1990) it appears

for even the richest

41 species.

on previous

richness

& Ersoy,

present

figure of58 mammalian

and grassland

doubt

on the 58 species

with many

It is compared

they have

The Pagalar

in any temperature does

is based

is high compared

steppe fauna

deciduous

conditions in terms and

to

for the of species

semi-evergreen

forests.

The assessment of ecological diversity or community structure by taxonomic order, locomotor and dietary categories of the PaSalar fauna are shown in Figure 1.

size.

Taxonomic order

Sue category

Locomotor adaptation

%

Feedlng adaptation

5Or 40 Pasalar

30 20

1 ti

SubtropIcal deciduous

semi-

30

forest

20

N=2

Tropical

30

forest

20

N=3

Ir

IO

deciduous

1L

LL

IO

5or 40 Temperate

mlxed

30

forest

20

N=7

JbL

IO ic

U

t

I& RIPACO

IL

i_.b bl

40 Troplcol

semi-

30

evergreen

forest

20

N=3

IO

bL bkb ABCDEF

I-S A%0

I FBGCO

Figure 1. Community structure of mammalian faunas, showing the means from four recent hahitat types compared with that of the Pagalar mammalian fauna. The recent faunas represent, from bottom to top, the mean of 3 semi-evergreen tropical rain forest environments in tropical Africa (Andrew el al. 1979). the mean of 7 deciduous and mixed forests from southern temperate regions of Turkey. southern the mean of 3 deciduous forest/woodland environments in Germany, southern France and Spain, tropical Africa, and the mean of2 semi-deciduous forest areas in the sub-tropical zone of India. These habitats are described more fully in the text. The vertical scale shows the percentage numbers ofspecirs. The four categories are as follows: taxonomic-R, Rodentia; I, Insectivora; P, Primates; A. Artiodactyla; C. Carnivora, and 0. others; size-A, O-1OOg; B, IOOg-I kg; C, I-10 kg; I), IO-15 kg; E, 45-180 kg, and I;, > 180 kg: locomotor-L, larger mammals wholly confined to the ground; S, smaller mammals that arc mainly terrestrial but also live on the lower parts of trees and bushes and on fallen trees; A, arboreal; SC, scansorial, and 0, others (fossorial, aerial, aquatic); feeding-l, insectivorous: F, frugivorous; R. browsing herbivory; G, grazing herhivory; C, carnivorous and 0. omnivorous.

PABALAR

Tcuonomy.

The

artiodactyls

taxonomic

and rodents,

different habitat

analysis

a pattern

shows the dominance

that is widespread

types today. This indicates

f‘orms, but it provides

31

PAI..4EOEC:OI.OC:\

little information

of the fauna by carnivores,

in mammalian

faunas from man)

that the Pagalar fauna is consistent about

/

its palaeoecology.

The

with living

last category

of

“others”

is relatively high in the PaSalar Iiuna, and this is due to the unusually hi,$ of perissodactyls (three rhinos, one equid, one chalicothere) as well as two proboscideans, one hyracoid and one tubulidentate. ‘There is no palaeoecolo,+cal number

significance

immediately

.Si;e. The analysis

analysis

because

adaptation.

apparent

in this pattern.

of body size is potentially

body size can be estimated

the most useful category

more accurately

of’ ecological

than locomotor

The Pagalar fauna shows a high level of size equitability

(Figure

or dietar\

1). The bodi.

size classes are more evenly filled than are those of any of the recent faunas, partlv due to for which there are less than half tlrc the low number of class A mammals (0-lOOg), cspected any

number unless the size structure

living mammalian

reconstructing

past ecologies),

the smallest

mammals

under-represented, structure

of the Paaalar fauna is completely

but a more likely possibility

at Pagalar.

If this is correct,

and by correcting

that is similar

which have similarly

at a different

herbivores

relate

carnivores

do so at second

dietary

directly

could b(

for the PaSalar

to subtropical

to the vegetation hand when

in the

seasonal

size

tijrests

predator

for example,

the two groups might therefore

be expected

the analysis,

there was a trend in modern habitats

open habitats

abundant

in tropical

during

(Bernarz,

(Andrews,

1983).

ofincreasing

S maller

mammals

following the pattern

were more abundant

&

1983). The

representative

(Andrews, feeding,

1983): whereas

(and sometimes

and prey is complicated

other

by non-

of the predators 1988; Mikkola,

It was found that when carnivores

forest habitats,

large mammals

habitat

to provide overlapping

if analysed

in more

together.

can be considered

to the strength

hunting by the predator

by Andrews

alone (Andrews,

they eat the herbivores

between

relating,

or to cooperative

man)mals

level from that of the herbivores

information

habitats

A mammals

is obtained

fauna from Payalar published

was that the carnivores

trophic

size relationship

considerations,

equipment

01

high levels of equitability.

reason for making this distinction

The

the category

to that of the faunas from tropical

‘I‘obien ( 1977) was based on the non-carnivorous

carnivores).

from

means

is the presence of a bias against

this bias a pattern

A previous size analysis of a less complete

of the habitat

difftrent

fauna. This could be true (in which case we are left with no

killing

1983). and

and maybe conflicting were eliminated

diversity oflarge

fi-om

mammals

were still found to be most

in Figure

1, while in more open

than small ones. The seasonal

forest faunas

were intermediate, and the Pagalar fauna analvsed in this way was also found to 1~ intermediate, with approximately equal numbers oflarge and small mammals (Figure I in Andrews, 1983). This lends support for the similarity fauna with seasonal tropical to sub-tropical forests. Another Legendre

way of analysing following

the size distribution

the methods

of Valverde

in size distribution

of fossil faunas

of the PaSalar

is that developed

b)

(Legendre,

1986). The body sizes of all mammalian species in a fauna are plotted in rank order (resulting in a “cenogram”) and the steepness of the line and the presence or absence of breaks in the distribution both provide an indication

of ecological

affinities

of the fauna.

The steepness

of the line is, of

course, the product of the fauna1 diversity: the bigger the fauna the more species have to be fitted into the cenogram and so the shallower the slope. This analysis also separates the carnivores

from the non-carnivores,

for the same reason as just described.

Species Figure 2. Cenogram of Pagalar fossil fauna, showing distribution of body weights of individual sprcies. The natural logarithms of body weights are shown on the vertical axis, and the rank of species in decreasing order is given on the horizontal axis. I(e)-: R, Rodentia; I, Insectivora; P, Primates; A. artiodactyla; S. Suidae; 0, Carnivora and n , Others (including all Perissodactyla. Prohoscidra. Tubulidentata and Hyracoidea).

A cenogram from

tooth

analyses

has been

published but the break

non-forest

faunas.

seasonal

form Large

the ground), fauna that

aquatic, contrasts

of small

conspicuous monsoon

than

live on the surface

similarity forests

of India

and

encompassing

the smaller

in shallow

low bushes, but other

perhaps

a

deciduous

Another mammals

(excluding

namely

1), and with

terrestrial those

woodlands

of the (<5 kg)

completely

and scansorial

of species,

(Figure

and

time on

30%

and so on. Together

Arboreal

classes

by small fauna

holes

trees,

by terrestrial

all of their

mammals.

from

both

these species

those

represented. This distinctive forest faunas, which have

adaptations

are dominated the Pagalar

fallen

from Pagalar.

of locomotor

from

with the

type,

is dominated

that spend species.

of the ground,

the tropical

fauna

species

of the mammalian

numbers,

which

habitat

shared

fauna.

adaptations, are poorly with tropical evergreen

between

estimated

forest and woodland is a feature

an intermediate

as larger

make up 80% of the fauna

on the other,

the body weights

tropical

of the Pagalar

50%

rocky areas,

in the distribution

Diet. In contrast

indicates

mammals,

in low but significant

faunas

using

specifically

for the PaSalar

(defined

ground

in banks,

fauna,

of the size distribution

spectrum

fossorial and aerial on the one hand

equitability forest

locomotor more

species

are present

of forest

constitute

species),

habitats,

also, therefore,

mammals

habitually

terrestrial

This

ground

consists

fossorial

of the recent

near the mid-point

of woodland

Locomotion. The species.

for the Pasalar

of ecological diversity (Figure 2). Comparison with the ( 1986) s h ows the steepness of the line to be similar to those

by Legendre

from the more wooded Africa,

prepared

size for the analysis

with

pattern greater

temperate There

is a

the subtropical

of Africa.

to the locomotor structure just described, the distribution of dietary classes in the Pagalar fauna shows a high level of equitability (Figure 1). Carnivorous species are most abundant, but insectivores are relatively scarcethis may be the result of the probable size bias against small (category A) size class mammals, in which most

PASALAR

insectivores

fall. Browsing

types,

grazing

with

This

proportion,

significant

and

aspects

comparative browsers

the

faunas,

there faunas,

with

single-canopy

forest position.

result

proportions

it corresponds

to the less seasonal

Significance Pazalar

fauna

taxonomic

tests

on these

and

any

analyses.

the dietary

There

modern

The

tropical

semi-evergreen

freedom

for 3 by 5 contingency forest

There

are

Pagalar approach

possibilities here

to which

fauna

also depend

in the middle on the unified

being

a selected

several

different

ones,

two

considerations must

communities,

and

portion

at an

faunas

as a

faunas,

and

at 0.001

(chi-squared differences

fact

that

past

changed

Secondly,

might

satisfactorily

way,

interpreted

for tht

8 degrees

the

of

nature

of the

The comparative

communities

with living

if the PaSalar

fauna

or through

to interpret.

the

and

can

the time of accumulation

community

answered,

in some

the

be

and the first consideration

since

be hard

fauna

38.92,

mammalian

the comparisons

of one original

the

and

from the subtropical

affinities.

day communities,

size

any of the faunas

in interpreting

here of its ecological

between

the

the Pagalar

no significant

of the fossil fauna:

similarities

between

between

be considered

may have

be measured these

difference

of the Pagalar fauna

the

Miocene.

and

of frugivores

differences for both

affinities

as present

nature

most recent

falling

forest

forest.

the comparisons are

communities

on

criteria

faunas

the seasonal

deciduous

must

communities

through

faunas

difference

provided

depends

by the same

the extent

that

and the evidence

used

interpreted Payalar

three

fauna

two the

in deciduous

of high proportions

no significant

with

Palaeoecological

the

of frugivores

to tropical

showed

was significant table),

or the tropical

relative

dietary browsers.

In

carnivores forest

comparative

locomotor

forests

and

matches

in terms

was also no significant

distributions.

semi-deciduous

forests

are

fauna.

dominance

semi-deciduous fauna

than

1).

comparisons

of the

from

and carnivores

(Figure

PaSalar

of grazers

of the Pasalar tropical

of grazers

the

of change

the seasonal

of browsers

represented

of frugivores,

of

to dominance

The structure

and low proportions

proportions

adaptations

is a gradient faunas

are the next most abundant

less commonly

high

dietary

forest

intermediate ofhigh

considerably

relatively

of the

in tropical

and frugivores

herbivores

herbivores

,579

PALAEOECOLOGl

degree

in comparison in general

Thirdly, of similarity

with habitat

communities

is biased,

being

is of the either

a mixture

of

and if the first with

differences

recent

from

and climatic

other

terms.

Communi[v change Mammalian ones,

communities

but there

is every

this is potentially community community.

in the past indication

many

of them

the case, some care is needed

type. The Pagalar The

can only be interpreted

that

most

notable

fauna

differs

difference

have

in making

in several

in comparison no recent

direct

respects

is the abundance

comparisons from any living

of suids,

with

present

counterpart.

Where

with any one mammalian

and particularly

the

dominance of Listriodon splendens and the closely similar but more bunodont Listriodon sp. These are omnivorous/herbivorous species with similarities in tooth structure to living tapirs. The dominance of the herbivore niche by an equid, a giraffid and two species of bovid

has similarities

with

tropical

African

woodland

habitats,

but the presence

of three

rhino and two proboscidean species and an abundance of small tragulid and cervoid species are not to be found in any recent habitat type. The way to proceed in this case is to

find common

terms

fossil fauna

and

ofreference

use these

between

as the basis

the habitats

considered

to be most similar

lor the palaeoecological

to th(,

reconstruction.

Taphonomic bias The nature

of the sediments

represents

and depositional

the accumulated

bones

distance

to the site during

a single

the short

time ofaccumulation

bones

for the Pagalar

some geological differences of predator

bias in the fauna

by transport existing

hand,

today.

but the distribution

This

could

place

those

of size classes

be due to predator

fauna and

in minor

biases

habitat

of single

relating

elements

followed

of the fauna. communities

to locomotion

and

the presence

of loss of very small

or loss of small

was

gives an indication

do not indicate

gives an indication

the

There

of the bones

is typical

classes

communities

selection

activity

1990). Weathering

of ecological

of living

have resulted

both add to the possible

of the PaSalar

the distribution

with

bias,

& Ersoy,

to its burial

area and

that provided

both in time and space. ofpredator

fauna

the short

catchment

communities

relief that would

area, and evidence

the distribution

Similarly,

diet are consistent

were restricted

(Andrews

of this material

On the other

The restricted

event.

the Pagalar

the site transported

that the mammalian

and topographic

the catchment

that

depositional

assemblage

variation

within

suggest

near

suggest

bone

environment

on the hill slopes

during

of

mammals.

transport.

Comparisons with recent communities The

community

both

temperate

semi-desert been

and other

shown

subtropical latter

structure

of the Pagalar

and tropical to

be

open with

forests country forest

than to the former.

These

Miocene. The faunas Kanha according During

I have

Park

to altitude is what

other

canopies abundant,

and

examined

from

the summer

also occur.

be sparse. there

There are

the moisture

The main

available

grass

is generally

vegetation

and

varying Much

in pure stands, closed

and lower

bamboo

may

1963).

African

meadows

(Eyre,

forest/woodlands

are also markedly

monotypic

and have the same

of a single

main

with

vegetation

interspersed

thick ground

are from

for tree growth.

deciduous

tree canopy

open

of India

most trees lose their leaves.

tree canopy

ground

the middle

and dry forest,

by Shorea robusta often

dominated

is thick

many

affect

from to the

and an account during

regions

of moist

of

have

and

are more similar

at Pagalar

rainfall

is a mixture

similarities

in common,

existing

and early spring,

Sal forest,

belt. They

typical

from steppe,

forest/woodland

amount

conditions

1967). There

in late winter

is called

rainfall

that

widely

The greatest

deciduous

have a certain

and soil, both of which

tree species

may

in the summer

from

It also differs

structure.

tropical

possible

(Schaller,

the dry season

the forest many

that

National

from

habitats

of these can be taken as indicating

may be distinguished

community

faunas

semi-deciduous

fauna

in all the analyses.

of but tree

be locally tropical

combination

with grassy

glades.

The climate is less seasonal in terms of temperature, but the impact of the dry season is probably more significant because of the lack of temperature variation: in the Indian summer-rainfall

forests,

much

of the dry season

is combined

correspondingly less effect on the environment. It may be inferred from this consistency of pattern

with cool temperatures,

that the climate

at PaSalar

during

with the

Miocene was subtropical at the very least, with a pronounced dry season. Less certainly it may be inferred that the rainfall pattern was one of summer rainfall, since the present climatic regime of winter rainfall in Mediterranean regions supports very different community structures of the mammalian faunas. The vegetation supported by this climate

was

probably

deciduous,

closed

forest,

interspersed

PA$AI.AR

PAI.AEOECOL.OC:\

dominated

b?- a single

with grass meadows

:iX

tree

and always

species,

deciduous

with abundant

I

to semi-

ground

vegetation.

Acknowledgements I am most grateful includes has

been

particularly with

taphonomy,

Mikael

I also

Financial

support

excavation Directorate

has been

who

Libby been

Department by these

and Alan

out on behalf and

who

has worked

Gentry,

who and

generously from

of the T.C.

Ministry

of Ankara.

ofPalaeoanthropolog)-.

and done useful

Boise

Financial

by

Fund.

of Culture

Faculty

many

commentrd

many provided

the

Martin,

on aspects

who have

provided

projects

the University

Lawrence

in the excavations

reviewers

for specific

in the field work at Pagalar.

the project,

Ersoy

has helped

consistently

assistance carried

Ayhan

Andrews

anonymous

of Antiquities, provided

who have helped who organized

the beginning,

has

with

and Geography, has bcrn

two

people

Alpagut,

Fortelius, and

thank

Foundation,

Rerna

us from

field identifications, trst.

to the many

who of the of the on the

suggestions. the The

Ixake) Pa~alar

and Tourism.

of Language,

support

This

Histor\,

for the csca\,ations

foundations.

References

.bdwxvs. P., I.lrd,~J. & Evans, E. hI. Iv. (1979). Pattwny ofrcoloqical divcrqit! in fossil ,md modern mammalian faunas. Hiol. ,/. /An. .Soc. 11. 177-205. :\ndrws. I’. CyrTohirn. H. (1977). New hlioccnc localit> in Turkey \\ith cvidcnw on thr wixins IrfRam;~I)itll~cu\ and Sivapithccus. ,V~z!ure 268, 599-701. :\rttwiou, C. ( 1984). Mammalian community palrorc~~logy: a wvirw ofrrcvnt methods \vith $prcinl wtiw~nct~ 11I Miorcnc mammalian faunas of Europe. ~~llvohiol. CIVIL. Rlontpellicr. 14, 91-l IO. Brckrr-Platcn. J. 11.. Sickcnherg, 0. & Tohirn. H. ( l’J75). Dir Glicdcrunq drr Kanozoischcn Srdimvnt~~ dvt Turkci nach \.c~rtrhratrn-FaunenRruppen. Gml. ,/cMt. 15, l9Ai. Brdnarz. J. C. (1988).Cooperative hunting in Harris’ Ha\\ ks f’orohukw unictwr[u~. .S&nce 239, 1525-I il’7 E:ngess~r-. B. ( lWC1). Inscctiwra und (1hiroptrra (hlamnlalix] aus drm Ncog-en drr Turkri. .S&r’ri,-. /‘CA. :16/r. 102. 47.-14% I+:\-rc. S. R. ( 1963). l+etafion and .Sooi/s. n Ctbrlri Pic&r<. b:d\\ard Arnold. London. Flrming. 7‘. H. ( 1973). Numhrrs of mammal spccirs in north and cc.ntl-nl .\mvrican forest communitic\ /+u/qr 54. ~5.55-.563. Flynn. I,. J. & Jacobs. 1.. I,. (1990). The mall mnmmn/.c ~~f~l’uplnr.,/. hum. f:‘w/. [III pms). Forstcn, A. ( 1990). :Inrhilherium from PaSalar, Turkc! J. hum. Gsd. 19 (in press). Fort&us, hl. (199On). Rhinoccrotidac from Pa$ar, middle Miownr of Anatolia. j. hum. 6~11. 19. 189-508. Fortelius. hl. (199(M). I.css common ungulate Eprcirr from Pqalar. middle hliorene of Anatolia. TurLr\ .,/. hrrn~ Ew/. 19, 4711-487. E‘trttlitrs, hl. & Bernor. R. ( 1990). A provisional s)-slvmatic ass~s~mcnt of the Rfiownc~ srmidra from P,~yd;lr. Turkr!,. ,/. hum. f+o/. 19, 509-528. (;azir) ( A. \V. (1976). ,]ungtertiHrr hlastodontcn aus Anatolien (‘l’urkci). (Al. ,/h. 22. I-143. (;entr>-, A. \Y. (1970). Thr Bovidac (hlammalia) of thr Fort Ternan f’auna. In (I.. S. B. I.eaky &,I. (;. %~\aq. Eds) b’o.cril I Prlrbrnfrr $i?f IU(I. ~1. 2, pp. 243-323. I.lmdon: Acadrmic Press. Grntr-y, ‘2. 12’. (1987). Rhinoccmscs from the hliocrnc of Saudi Arabia. R&l. nr. .\lus. nn/. I!;$!. iCAw/. J 41. 4OOL432 Gents). 11. \V. (1990). Ruminant artiodactyls of PaSalar, Tukq. ,/. hum.f+h/. 19. 529-5.X. <;iirhiiz. hl. (IWO). Preliminary list of Paaalar Carnivora.,/. hum. God. 19. 463-464.

Harrison, J. 1,. (1962). The distribution off&ding habirs ,m~ong animal> in a rt-opic.il t,~in 53-64. Legendre, S. (1986). Anal+ of mammalian communities from thr tats I’orcn~ and France. Palaeooer/ehrata 16, I9 t-2 12. Legendre, S. ( 1987). Concordance entrc pat&mtotqir ccmtincntalr ct tcs &+nimtnts C.K. Acad. .Sci. 304, 45-50. May, R. M. (197.5). Patterns ofspecies abundance and diversit). In (Xl. I,. (:ody &J. hl. Relknap Press. and Evolution of Communities, pp. 81-120. C amhridge:

tiltr.\t.,/.

.Irr/u/. I:;l)/. 31.

Ologor~~n~~ 01 ~outlltw~ pat~~)cr;ln~)g~~rplli(tu~.\. Diamrmd,

Ed\)

f:i~/nq~

Mikkota, H. (1983). Owls ofEurope. (T. & A. II. Pqscr, Eds). Cartton: Olson, E. C. (1966). Community evolution and the origin of mammals, Ecol. 47, 291-302. Schaller, G. R. (1967). The Deer nnd the Tiger, a .Slu(v OJ HiM/iji in India. Chicago: University of Chicago Prrss. Schmidt-Kittter, N. (1976). Rauhtiere aus dem .Jungtertiiirs Kteinasiens, Palaeon~ographicn 15, t-131, $en, $. (1990). hliddte Miocene tagomorphs from Payalar, Turkey. J. hum. Euo~. 19, 455-46 I. Simpson, G. G. (1964). Species diversity of North American rrcc‘nt mammals. .$w/. Zool. 13, 57-73. Tohien, H. (1978). New species of Cricetodontini (Rodent& Mammalia) from the hliocene ofTurkey, .Ilnin,-rr gemrim. .Ilil/. 17, 209-2 19. ~~nay, E. (1990n). Turkonyspasalnrmsi~ Tohien, its range of\.ariation in the type localit! at Pagatar. ‘Turkey ,,/. hum. EJol. 19, 437-443. ijnay. E. (1990h). A new species of Pliospulm (Rodentia, mammatia) from the Middle hlioccm~ of Pqalx, Turkey. J. hum. Ewl. 19, 445453.