An early Pleistocene fauna of small mammals from the Kadzielnia Hill in Kielce (Poland)

P A L A E O N T 0 L O G I C A I 9 58

KAZIIMIERZ KOWALSKI

P O L O N I C A

No. I

AN EARLY PLEISTOCENE FAU NA OF SMALL MAMMALS FROM THE KADZIELNIA HILL I N KIELCE ' (POLAND)

Abst ra ct. - A des cript ion is given of the Lns ect ivo r a , Chiropter a , Lagom or p ha and Rod en ti a fou n d in clay dep osits fill ing up karst poth ol e s in the Kadzielnia Hill of Kielce.Th e collected fauna is of early Pleistoce n e .age and is proba bly referable to th e Gunz-Mind el Inte rgl ac ial. The following species have been identified : Talpa minor Fr eud enbe rg , T.jossili s Peten y i, Sorex d . runton ensis Hinton, Sorex sp., Ber em endi a fissi dens (Pet e n y l), Petenyia hungarica .Kormos , Rhinolo phus cf. jerrumequinu m (Schre ber ), Myot i s cf. exilis Hell e r, Myo t i s sp., Hypolagus brachygnathus Kormos , Pli ola gus cf. tothi Kretzo i, Sci urus sp., Dolomys epis cop alis (Mehely) , D. kretzoi n.sp ., Mimo mys pli ocaenicus (F. Maj o r ), M. reidi Hinton, M.ne wton i F. Ma jor , Apodem u s sp., Muscardinus sp, The desc r ip ti on of a new sp ecies - PromimO'mys insu lijerus n.sp., previou sl y indeti f ied by the writer as

"Mimomy s d . pusillus (Mehely )" , supplem ents his earl ier paper on th e fauna from Podlesic e .

INTRODUCTION

Th e Kadzielnia Hill (20^{0 3}8'E

, 50

0 5 2 ' N )

is an elevation of 295 m a.s.1.,

lying within the precincts of the town of

K

ielce, in t

he Hol

y

Cro

ss

Moun tains

region .

Th

e central part of that h ill , which is also its

cu lmi n ation point, con sists of rocky Upper

De vonian (Middle Frasnian )

li

mestone.

Limestone qu arries in the Ka

dzielnia H

ill

h

ave

been worked

fo

r m

an y years

p

ast,

bu t its central portion is a p

rotected area. I n the

cour se of the' last war the nature protection regul ations w

ere v

iolated ,

in

consequen ce of which par t of th

e reserve was

de vastated. K

ar s t

phenomena occur in the rocky Upper

Devon ian limestone bu ild ing up

the Kadzielnia peak, such as numerous tunnels , dolines and pot

holes a

s

well as two large caves. Th e cave de

posits

have not thus far

b

ee n

investigated and have mostly been preserved untouched. The deposits

filling in the potholes and do lines, however, are

.

partly damaged

t h rough w ork at th e limestone quarries. Sections of the material filling

up some of these p oth oles are still observab le in places where th e

working op erations have ceased,

i.e .

on the sides of a rock-wall within

the protected area. In th e south-western portion a large dol ine (funnel-

-like pit) is readily noti ceable, filled in b y red clay. Its upper layer,

2 KAZIMIERZ KOWALSKI

severa

l

tens of centimeters in thickness,

is

repl

ete

with bones of mam m als, among which fragmentary remains of the Leporidae predom in a te. Bones from this layer are to be

seen throughout t

he outwashed surface of the pit, as well as at th

e

foot of the quarry below it where the bones h av e been transported by rain waters.

Bones of mammals have also been coll

ected

fr om potholes in th

e

north-western part of the protected area. They have be

en

yielded by the sandy clay depo

sits

filling in the vertical outwashed fissure. In the systematic par t, the sites of these two finds are referred to as the dol ine and the fis

sure.

Furth

ermore,

near the mentioned f issu r e, some fragments of a bone br eccia have been d

iscovered

un der remnants of the karst pothole destr oy ed during the working of the quarry. T hese bone br eccia r

ema ins ,

however, proved so few and unsatisfactorily preserved as to b e unidentifiable and have, henc

e,

b

een

left ou

t

from the General Remarks and from the Syst

emat ic

D

escriptions

. They consist of teeth of Myotis sp.,

Apodemus

sp

.

and some more closely indeterminate Microtinae and probably Ochoton

idae.

The caves within the Kadzielnia Hill area h av e been known for a l ong time. As early as in 1926, E. L. Niezabitowski descri bed the carpus bone of a rh inocer os Coelodonta antiquitatis (Blum.), discovered in the K

adzielnia Hill of Kielce,

which was presen ted to him by J

.

Rostafiilski

.

In 1932, J . Czarnocki wrote abo ut the caves and poth

oles

of Kadzielnia which yielded a fauna containing the r

emains

of a rhinoceros, rein deer and a lemmi ng

, Dicrostonyx torquatus

(Pall.). A mention of this P leis toce ne fau na is also made in

a

note by the same author, p rin ted in 1949. And again, during the sam

e

year

,

J

.

Czarnock i wrote about the karst

I

p otholes of K adzieln ia "with an interglacial fauna of step pe origi n". The correctness of the identifications given in th es e n otes canno t be verifie d owing to l ack of descri ptions or fig ures of the fo un d foss il r ema in s.

If

they h av e not be en misplaced bu t actually do b elon g to yo u ng P leis tocene forms t heir p resence wo u ld suggest the occurrence in the Kadzielnia H ill of a yo unger Pleistocen e fa un a too

,

tho ug h we do not kn ow w hether it was ob tain ed from t he area of the pr esent nature reser ve or f r om other pa r ts of th e hill , now devasta ted.

E arl y Pleistocene fossil re mains wer e found by t he p r ese n t aut h or

in 1950. Duri ng fiel d-wo rk don e in 1956 an d 1957 he collected cop ious

m ater ial fro m d ep osits filling in the large doli n e in the sou the rn wall of

the qu ar ry

,

as well as so me bon e remains fro m t he potho les an d fi ssures

in the n or th -western part of t he pro tect ed area . In his p aper on the

f au n a fro m P od lesice (1956) the autho r mad e m en ti on of the occurrence

in Kad zielnia of

Ber em endia fissidens

(Pe teny i), an ea r ly Pleistoc ene

spec ies. Verteb ra e an d scales of a re ptile, found b y h im in assoc ia tion

PLEISTOCENE FAUNA OF SMALL MA MM ALS 3

with bones of small mammals, have by D r . M. Mlynarski been identified as

belonging to Ophisaurus

cf.

pannonicus

Kormos. The collected m ater.al also comprises detached teeth of th e indete rm in at e Mustel idae and some few remains of snails which have no t so far been investigated.

The w r it er h ere con veys his thanks to

Dr. M. Kretzoi,

Director of the Geological

Ins

tit u te of Budapest, for the h elp shown during the work of identify ing a part of the micro tine m at erial , as well as for his friendly comments and r eadiness to cooperate. The writer w ishes also to th ank Mrs. J

.

Humnicka fo r t he Engl

ish

transl a tion of the Polish text, and Mr. J. Swiecimski for t he

pain s

h e has ta ke n

in

pre pa ri ng t he draw ings.

ORIGIN AND AGE OF THE BE DS

The v ertebra te remains in t he dep os it s of the k ars t doline

in the

Ka- dzielnia Hill w ere probably ac cum ula te d b y th e action of rain water which carried th ere bones of animals who p erished in the vicinity of t h e pit or perhaps par tly in th e hol es of th e L epo ridae. Th e various bones were found lying s ide by si de, without anatomical or de r and usually str on gly cracked. The satisfactory s ta te of preservation e ve n of very small bones is du e to th e doline b ein g g rad ua lly fill ed up by lim estone residuum wi t h a h igh content of calcium carbon ate. Th e thinness of th e b on e-bearing bed undoub tedly suggests th e con tempo ra neousness of the r emains preserved th erein.

Th e faunal composi ti on of the. m aterial filling in the dolin e, is as follows (figures refer to number of specimens ):

Insectivora

Talpa minor Freud enberg - 3 T.fossHis Peten y i - 3

Sorex cf. runton ensis Hinton - 2

Sorex sp. - 3

Ber em endip fisside ns (Pet en y i) - 3 Pete n yia hu n garica Kormos - 3

Rhinolophus cf. ferrumequinum (Schreber ) - 2

Chiroptera

My otis d. exil i s Heller - 3 Myoti s sp. - 6

Lagomorpha

Hypolagus brachygnathus Kormos- 38 Pliolagus cf. tot hi Kretzoi - 7

Rodentia

M.rei di Hint on - 2.1 M.newtoni F. Major - 4 Apodemus sp . - 2 Muscardinus sp . - 1 Sciurus sp. - 1

Dolomys epi scop ali s (Mehel y) - 4 D. kretzoii n.sp. - 8

Mimomys pli oca enicus (F. Major) - 29

Furt h ermor e,

scales of t

he

glass lizard,

Oph isaurus cf. pannonicus

Kor-

m

os,

ide ntified by M.

Mly n arsk i, h

ave also

been collected

t

he re.

4 KAZ I MIE R Z KO W AL S K I

T h e material filling in the fissure of the north-western p ar t of t he protected area has yielded bon es of

Sorex cf. runtonensis

H in ton ,

Pet en y ia hungarica

K or m os and

Mimomys pLiocaenicus

(F. M aj or).

H en ce we may suppose that this fauna is of the same age wit h that occurring in the doline.

WHh the exception of

Rhinolophus ferrumequinum

(S ch reber), recorded from a number

of early

Pleis tocene sites in Europe, bu t also s

till

living now, all th

e

other forms represent fos

sil

species.

Talpa min or

Freudenberg , usu all y refer re d

to in li

terature under the

synony mi c

name of

Talpa gracilis

Kormos,

is

known from early Pl

eis tocene

b

ed s

of Rumania (Puspokfurdo, B

r asso),

southern H u ng ar y (Beremend 4, Vill

an y 6,8,

Nagyharsanyhegy 4), Austria (Hundsh eim), Yug oslavia (Podumci), German y (Sackdilling er Hohle, Erpfingen, Gund

ersheim,

Mau

er,

Breit enberghohle), P

oland

(P

odlesice)

an d proba bly Italy (Verona) . The other spec ies of mo le,

Talpa fossilis

P eten yi , mor

e

oft

en

r

ecorded

under the sy no ny mi c name of

T.praeglacialis

Kormos, occurs either in as

sociation

with the last named for m or alone

in many early Pleistocen e

f

au n as.

Thus far it h

as

been r

ecorded

fro m Rum

ania

(Puspokfurdo, Brasso) , Hungary (Beremend 6, Csarnota 1, Villany 3,5,6,7,8,11, Nagyharsanyhe gy 4), Czechoslovakia (Gombas

ek

and Koneprusy) , Austria (Hundsheim a n d Laerberg in Vienna) and fro m G er m any (Sackdillinger Hohle, Gundersheim, Erpfingen, Mauer, H oh en- sulzen). To this spec

ies

may probabl y be referred fossil remains fr om D

od rech t in Holland,

from

the Fresh.

Wa

ter

B

eds

of England and from V

erona in Italy.

The rema

ins

of shrews from Kadzielnia have not been identifie d taxonomically with any cert a inty . They belong

t

o two species, one of w hich apparently s

eems

to be i dentical with

Sorex runtonen sis Hin ton,

known from early P leis tocene sites of West R u n t on and Backto n in E ng land

,

S ackdill ing er H oh le in Germ an y

,

H und sh eim in Aus tria , G ombas ek and K on ep r usy in Czech oslov ak

ia ,

B er em end 5, Csarno ta 2, N agyhar san yhegy 2,4 an d Vill any 3,5,6,7,8 in Hung a ry as we ll as from P odum ci in Yu goslavia. T h e ot her species of K ad zieln ia shrew approaches.

Sorex minutus

L., now living

in

E urope b u t also r ec ord ed as an early Pleistocen e fos

sil for m from

G

er m an y (Sa

ck dillinger Hohle, Erpfingen, Gaisloch

,

Westhofen, Br eiten b

ergh oh le,

Hohensulzen) an d from Hung ary

:

(V ill any 3,6,8

,

Csar n ot a 2), while during the

youn ge r Pleistocene

it seems

to hav

e

been a widel y d

is trib ut ed fo

r m .

Ber emendia fiss id ens (Peten yi) is a characteristic species

in early

P

leis.o cen e

f

aunas

of cen tr al Europ

e.

It has b

een

r

ecorded

from

Puspck Iiirdo

an d Bras so

in Rumania

,

from Cs

arnota

1,2,4, Beremend 5,

Villany

1,2

,3,5,6,7,8,11

al

so

Nag y h arsa nyhegy 2,3,4,5

in Hungary,

from

PLEISTOCENE FAUNA OF SMALL MAMMALS 5.

Sackdillinger Hohle and Gundersheim in German y

,

fr om Gombasek in Czechoslovakia and from Verona in Ital y .

Petenyia hungarica

Kormos

is asso-ciated

wi th the las t n amed for m in many e arly Pleistocene sites. It is known from Puspokfurdo in Ruma- nia, from Villany 3,5, Csarnota 1,2, Beremend 5 and Nagyharsanyh egy 2 in Hungary, from Gundersheim in Ge rmany and from Podlesice in Poland. A closely related or even p erhap s identical species has been describ ed by A. Pasa (1948) from Verona. Fin ally , M . Kretzoi (1943), on a figure published

in

a paper by A. Dubois and

H.

G. Stehlin (1933), has described a new species, the

Petenyia ste h lin i Kretzoi,

from the younger Pleistocen e deposits of Cotencher Cave in Switzerland.

If

Kretzoi's assignment is correct, it would indicate the persistance of genus

Petenyia

Kormos to the b eginning of the Wiirrn glaciat ion period .

None of the three species of bats discovered in Kadzielnia leads t o conclusions as to the age of the collected fauna.

Rhinolophus ferrumequi-

7tum (Schreber) is known from the early Pleistocene down to th e present time, while spe cim ens from genus

Myotis

Kaup are specifi call y indetermi- nate. One of them seems identical with

Myotis exilis Heller,

known from t he early Pleistocen e of Gundersheim in Germany, and probably also present in the fauna of Podlesice, Poland.

Hypolagus brachygnathus Kormos

is recorded from Csarnota 1,2, Villany 3,5,6,7,8, from Nagyharsanyhegy 2,4 and B eremend 1,4,5 in Hun- gary, from Puspokfurdo in Rumania, fr om P odumci in Yugoslavia, from Gundersheim in Germany and from t h e Tegelen Clay Beds in Holland.

Pliolagus tothi Kretzoi,

to which are prob ably r ef erable the fossil r emains.

of a sm all har e from K adzielnia, ha s thu s far been recorded onl y f rom Betfia in Rum an ia.

Pliolagus beremen d ensis

Kormos, stron g ly r es embling it and differ ing in smaller size only, is kn own fro m Csa rnota I , Villany 3 and Beremend 4 in Hung ary . It

is interesting to note

that - as s tated by Kretzoi

(1941a) -

in the old er ea r ly P leistocene fau nas of Hunga ry w e may e n counter small numb ers of

Pliol agus

Kormos associate d wi th the

;predomin an t genus

Hypolagu s

D ice (Csa rnota, Villany, Berernend). In

Pusp okfurdo (Rumania), together with t he dom in a tin g ge n us

Hypolagu s

Dice, we m ay occasionally encou nter

Lep u s L.,

while

Pliolagus

K ormos

still p ersis ts there. In t h e still yo unger faun a fro m Be tfia,

Lepu s

L .

is

pred om in an t,

Hypolagu s

Dice cons titutes 22 per cen t of the total num ber

of L

eporida e, w

hile on ly 3 per ce nt are ma de up by

Pliol agus Kor

mos,

h er e rep r esented by P .

toth i

Kre tzoi. I n Nagyhars an yhegy ,

Pliolagus.

Korm os is a lready absent,

Hy polagu s

Dice is distinctly scarc e,

Lepu s

L .

.

b eing th e p r ed om in an t gen us. F in all y

, in

Brasso an d s t ill y ounger fa unas,

ge n us

Lepu s

L

.

is th e only r e p r

esentati ve of

the L ep or idae. H ence

, eve n

6 KAZIMIERZ KOWAL SKI

the composition of th e L ag om or ph fa una in K ad ziel n ia suggests its assignmen t to the older perio d of the ea rly P leis tocen e.

Genus

^{SciU TU S}

L . is kn ow n fro m the E arl y Ter tiary u p to Recen t times. In ea rly P leistoce ne fa u nas it is extre mely

'r are. It h

as been recorded fr om Ko n eprusy in Czechosl ov ak ia , Betfi a in Rumania, S ack- dilling er Hohle an d

Gu nd er

sh eim i n Ger m an y and fr om W es t Runton in England. Its oc cu rren ce in vari ab ly in dicates the pre sence of a fores t.

Dolom ys epis copalis (M

eh el y) is known fr om Rumania (Puspokfu rdo, Brasso), from Hung ary (Beremen d 6, Nagyharsa ny hegy 3,4, Villany 3,7,8, Ostramos, Bud ap est) , fro m Czechosl ov aki a (Gombasek

,

K on eprusy) , fr om

A

us t ri a (Hundsheim) , f r om Ital y (Veron a) and from

Ger m an

y (S ack- dillinge r

H

ohle, M auer , Esc

helbronn,

Erpfin gen , Gaisloch , Breitenberg- hohle, Hohenstilzen) . O

.

Fejfar

(1956a)

r egards it as a sylvan xero t he r mic

speci

es .

Dolom ys kret zoi

n. sp . is a spe cies new to scie nce

,

on w

hose

relationships on ly very li ttle can be said.

Mimomys pliocae ni cus

(F. Maj or), abun da nt in K adziel n ia, has so f ar been r ecor ded fr om E ast Run t on , Norfolk and Suffolk in Engl and, fr om numerous borings i n H oll and, fr om Va l d'Arn o in Italy , f ro m Seneze and Her ault in France, fro m Gund ersheim in G erm an y and f ro m P uspo kf'urdo in Rumania. Specimens from early Pl eist ocene sites in Hung ary (Berern end 4,5, Nagyha rsan yhegy 1,4 , Villan y 3,5,11) have b y Kretzoi (1956) been na me d

Mimomys mehe lyi Kretzoi.

In that aut hor's opi nio n M.

pliocaenicu s

(F . M a jor ) is r

estricted

to the Cal ab r ian hor izon (Villafr ank ian)

,

whil e M.

mehelyi K

r etzoi

oc cu

rs in the yo unger fau nas of H ung ary

.

In the lack of a des crip t ion

M. mehelyi

Kretzoi m ust b e regarde d as a nomen nudum an d it would seem that t he Hung arian specim ens const itute no mor e than a dis tinct s u bspecies.

Mimomys reidi H

in ton

is

k now n from Tri mingha m in E ngl and, fro m Bred a an d s' Gravendeel in H oll and , from Gu nde r sh eim in G erm an y, fro m Veron a in Italy, fro m K isl ang and B

erem en d

4 in Hun g ary.

Mimom ys new toni

F . M aj o r , has, so far

,

been r eco rded fr om W est Run ton and Norfolk in E n gland , f r om Tegelen in Holl and

,

Sen eze in Fran ce, Gunder sh eim in Germ an y an d fro m K islang, Nagyhar sanyhegy 1,4, V ill an y 3,5,11 an d Beremend 4 in Hu nga ry . The H ungar ia n spe cimens h ave by T. K ormos (1938) b een refe rred to a n ew s ubspecies,

Mimomys newtoni hungaricu s

Kormos, w hile Kretzoi (1956) eve n con sid ers them as a sep arate species,

Mimomys hun garicus Ko

rmos. M.

newtoni

F. Ma j or is as a r ule as s ocia ted with the species M.

pliocaeni cu s

(F. Ma jor) and M.

reid i

H in to n .

G enus

Apod emu s

K aup h as been repo rted from m an y localities

beginning w ith the ea r ly Pl ei st oc en e , t h ro ugh to Recen t times.

T

he

K

a-

PLEISTOCENE FAUNA OF SMALL MAM M ALS 7

dzielnia specimens seem to belong to

A. alsomyoides

Schaub, r ecorded from Vill any an d Beremend in Hungary and from Magyarko in Rum ania,

while its occurrence in Puspokfurdo is also possible.

Genus

Muscardinus Kaup

is of rare occurrence in early Pleistocene localities, probably owing to their mainly steppe character.

It

is an animal distinctly connected with forest environments.

It

remains have been reported from Koneprusy in Czechoslovakia, from Moggaster Hohle and Sackdillinger Hohle in Germany and from Puspokfurdc in Rumania.

Ophisaurus pannonicus Kormos has probably been recorded as early

as from the Miocene, and

"subsequently from the Pliocene and early

Pleistocene of Hungary and Poland.

The time distribution of the particular species constituting the Ka- dzielnia fauna clearly indicates its early Pleistocene age. On evidence of the stratigraphic column of the Pleistocene faunas of Hungary as given by Kretzoi (1956), the Kadzielnia fauna is referable to that author's Villanyium horizon, equivalent to the Gtinz-Mindel Interglacial. The Kadzielnia fauna is distinctly younger than that from Podlesice

(K.

Ko- walski, 1956) which contains a number of archaic forms such as

Baranomys loczyi

Kormos,

Parapodemus coronensis

Schaub,

Promi- momys insuliferus n.sp., but lacks representatives

of genus

Mimomys

F

.

Major. Hence the Podlesice fauna is in all probability referable to th e ea rl ies t period of the Giinz-Mindel Interglacial,being thus contempora- neous with th e Hungarian fauna from Csarnota. Naturally

,

as compared with the Hungarian faunas , our fauna presents d istinct differences re ad ily interpreted by th e considerable distance se p ara ti ng these two

'

are as

.

Thus, e.g.

Dolomys episcopalis

(Meh ely ) is not en coun tered in Hungary before th e Biharium

,

th a t is t o say i n faunas eq u iva len t to th e Minde l glaciation period .

In view of our po or knowledge regarding the ea rly Pleistocen e faun as it is yet to o soon to attempt an interpretation of their geographical componen ts.

If Hypolagus brachygnathus Kormos really b

elongs to thi s genus, we ar e then dealing here with a r epresentative of ani ma l forms common to th e Pl eistocene faunas of both Europe and North Americ a.

Other species from the fauna of Kadzielnia are mostly known from localities in central Europe, partly also from thos e in Western Europe.

The definition of the climate prevailing in the Kadzielnia Hill at the

time o f the formation of d eposits filling up the karst depressions

is by no means easy. Practically all the faunal components th ere are

fossil forms about whose climatic requirements

1110

direct statement may

be made. The presence of

Rhinolophus ferru m equinu m (Schreber)

, a bat ,

a nd

of Optueaurus pannonicus Kormos,

a doubtlessly xerothermic reptil e,

8 KAZIMIERZ KOWALSKI

suggests a warmer cl

ima te

th

an t

hat now prev ailing

in

Poland

,

poss

ibly

one approaching the Mediterranean climate. The material which has

y

ielde d

t

he consid ered fau n a, namel y a red product of the chemical weathering process of limestone, belonging to the typ

e

of terra rossa

,

leads to th

e sa

me

suggestions.

The pres

ence

of typically sylvan mammali an gen er a

, such as Sciuru s

L.

an d Muscardinus

K

aup

are

ev

id ence of the ex is ten ce ther

e

of

fo res ts.

The s ca r ci ty of these forms,

however, sugges ts that the wooded ar eas

constitu t ed only

a

secondary

ele men t of the land scape.

SYSTEM AT IC DESCRIPTIONS

Inse ct ivo ra

Bowdich, 1821 Family

Talpidae

Gray

,

1825 Subfamily

Talpinae

Murray

,

1866

Genus

Talpa Linnaeus,

1758

Talpa minor Freudenberg,

1914

The sy n o ny m ies hav e been given in K. Kowalski's paper of 1956, Insectivor e s..., p. 341; also

1956. Talpa minor Fr eud en b e rg; M. Kretzoi, Die altp lelstoza n en Wirbeltierfaunen..., p. 162, 192, 197-200, 232.

1957. Talpa gracilis Kormos; G. Bru nn er . Die Breit enberghoh le..., p. 360, 363- 365, fig. 3.

Material. -

2 humeri

, one com ple te, t

he

ot her da

m age d, also 7 mand

ibul ar

fr

a gment s

w

ith P,-M3 and 2

detached t

eeth ; all

from the dolin

e.

Descripti on. -

In st r uc tur

e

th

e s

tudied humerus r

esembles

that of

Talpa europae a

L

..

differing in sma ll er dim

en sions only.

S

tructu r e

of mand

ibl e and teeth also ap p r oachi ng t

hat

observed in recent species.

F

oramen

m

en tale unde rlying t

he

tr igonid of

p

^-t-

Dimension s. -

Len g th of the

co

m pl

et e

humer u

s

11.6

mm, wi

d th

3,0

mm,

t

h at

of the da

maged

specim en 3.1 mm.

Dim ens ion s of ma

ndibul ar frag

ments (in mm) -

see table on p. 9.

An

other

m

andibular

fr agmen t cont

ained P"

P

2,

P

, with a len

g th of

1.2 mm, 0.7 mm and

1.0 mm

respectively.

Sy stem ati c position . -

The d im

e nsions of

hu me r

i an d of the

m

an-

dib

ula r frag ments are

disti n ctly

smaller tha n those

of T.

europaea L., suggesting the ir assignment to

T.

minor F

reu den berg

w

h ich

is a

smaller

ear ly Pleistocen e

mol

e.

PLEISTOCENE FAUNA OF SMALL MAMMALS 9

Man di bles - -- - ... 2 3 4 5 6

height of mand ib le on the inner side

below Me 1.8 1.6 1.6 1.9

th ickness of same 0.9 1.0 1.0 0.9

Pi le n gt h 1.75

M_I length 1.9

M₂ length 1.9 1.5 1.6 1.6

~ width on trigonid 1.0 1.2 1.0 1.0

M_a length 1.6 1.5

M_a width on trigo nid 1.0 0.9

.

Talpa fossil.is Petenyi, 1864 (fig. 1)

1864. Talpa vu l garis tassi lis Pete n y i; S. J. Petenyi.Hatrahagyott munkal, p. 53-58, pI. 1.

1914. Talpa eur op aea var.major ; W. Fr eudenbe r g , Die Saugetiere..., p. 660-661, pl.

47, fig. 28-31, 34.

1930a. Talpa pr aegl aci a li s n.sp.;T. Kormos, Diagnosen..., p. 238-239.

1933. Talpa cf. praegLiacialis Kormos; F. Heller, Ein Nachtrag..., p. 61.

1934. Talpa praeglacialis Kormos; G. Brunner Eine praglaziale Fauna..., p. 307-308.

1936a.Talpa praeglacialis Korm.; F. Hell er, Eine ob erp lioc a ne Wdrbeltierfauna..., p. 106.

1936b. Talpa praeglacialis Korm.; F. Heller, Eine Forest-Bed-Fauna..., p. 5-6. 1937b. Talpa praegla ciali s Kormos ; T. Kormos, Revision de r Kleinsauger..., p. 25- 26,

fig. 1.

1938. Talpa tassilis Petenyi; M. Kretzo i, Die Raubtiere..., p. 91-92.

1939. Talpa pra egliacialis Kormos; F. Heller Kleiristiu gerreste , p. 11,f,ig. 5.

1943. Talpa prae gla cialis Kormos ; A. Schreuder , Fossil voles , p. 405-406.

1952. Talp a pr aeglacialis Korm.; W. Weiler , Plioz an ..., p. 158-159.

1954. Talpa pr aeglacialis Kormos ; F. Heller, Neue Fundstellen..., p. 470. 1956b.Tal p a tassilis Pet e nyi; O. Fe jfa r , Seznam druhu.., p, 27q.

195'6. Tal p a tassilis Pet en yi ; M. Kretzo i, Die altp leis tozanen Wirbeltie rfaun en..., p. 165, 169, 171, 187, 193, 195, 197-201, 203, 232.

Material. - 6

specime ns of humerus,

of which two complete, also 4

m

an dib ular

fr

ag men ts

an d som e detach ed

te

eth

represen ting P

I-Mao

All

th ese re m ain s were

collected from the ma

terial

fill

ing

in the k

arst

doline.

Description. -

In structur e an d d imensions

the collected specimens

of humerus do not

d

iffer fro

m

t

hose displ

ayed

by recent sp ecim ens of Talpa europaea L. Th

e st

r uct ure of the m

andible,

howewer, differs dis tinctly as comp are d

to

that in the r

ecent

form. The molars, with

a

h

eight similar to the height in recent sp ecimens

,

are

con

side rably

les

s

10 KAZIMIERZ KOWALSKI

broad, the talonid in M2 being particularly narrower. The tooth-row is sho r ter than those in T. europae a L. The tips of teeth do not show distinct forward curving.

Dim ension s. - The lenght of the two complete specimens of humerus is 14.0 and 14.1 mm, the width 4.1 and 4.1 mm respectively.

Dim en sions of mandibular fragments (in mm):

Mandibles 2 3 4

heightof'm a nd ib le on the inner side be- low M₂

thickn es s of same p) length

P2length P₃ length PI length M) length M₂ length

M2 width on tri go nid M₃ leng th

M;j wid t h on trigo ni d

2.1 2.1

1.2 1.4

1.4 1.0 1.0 1.5

2.4 2.4

2.3 2.J

1.2 1.3

2.0 1.0

Systematic position. - Four speci es of genus Talpa L. are known from ea rl y Pleistocene beds of central Europe, namely: T. minor Freu- denberg, T. fossilis Peten yi , T. stromeri Brunner, T. episcopalis Kormos.

T. min oT Freud enberg is distingu ish ed by particularly small dimensions, while T. episcopalis Kormos is distinctly larger sized. The other two species corres- pond in size to the recent sp ecies T. euro- paea L. and also agree with the here described fossil remains from Kadzielnia.

~rnr.1 T.stromeri Brunner, described from Gais-

loch in Germany (G. Brunner, 1950), is distingu ish ed by the tips of molars being bent forward. In the Kadzielniaspecimens,

Fig. 1. Talpa jossHisPeten y i, however, this feature is lacking.As shown

fragment of mandibl e; slide an d by Kormos in his paper of 1930a, T. fossi-

top view. li_IS Petee enyi. IS. among at ers c aracterizeh h . d by the posterior portion of M3 being distinctly narrow. In the Kadzielnia specimens, the molars - M3 included - are narrower than the corresponding teeth in Talpa europaea L. This feature, therefore, distinguishes the collected fossil remains from the recent species and

PLEISTO CENE FAU N A OF SMALL MAMM ALS 11

re asonab ly refe rs th

em

to

Talpa fossi lis '

P e te ny i, a species widely dis trib uted in t he ea r ly P leist ocen e of Europe.

Fa m ily Soricidac Gray, 1821 Subf amily

Soricinae

Murray , 1866

Genus

Sorex Linnaeus,

1758

Sor ex cf. runtonensis Hinton, 1911

(fig. 2)

)mm

coronoid process meets the angle, i ts upper part being

Fig. 2. - Sorex d . runtonensis Hin- ton,fragmentofmand ib le ;sid e and top

view.

Material. -

Mandibular fragm en t with processes a nd M

1

and M

_a,

a nothe r with P

4-M 2 ,

and a de tached mandibular in cisive from the material filling the dolin e ; also a fragmentary processus coronoideus and a fragment of mandible with M

2

f rom the material filling the fissure in the NW part of the quarry.

Description.

- Anterior margin of the body of the mandible at a sli g h tly obtuse som ewhat in clin ed forward. Fossa

pterygo idea high , triangular. Facets of processu s articul ari s connected by a broad, slightly arcuate bone b ridge.

The poin ts of teeth pigmented to a redd is h-brown hu e. Talonid of M"

not reduce d

,

t his tooth be ing 5-cus- ped . Th e !'1andib ula

r

incisive h as

t h ree dis tinct t ubercles .

Dim ensions. -

The length M

1-Ma

is abo ut

3.7 mrn

,

h

eight

of m an di- bular bod y below

M2

1.3 mm , thick-

ness the re 0.8 mm

,

length of M

1

1.6

,-,-"_..:.-.c",,,

mm and 1.5 mm, length of

M2

1.2

a

nd

1.3

mm.

Sy st em at ic position. -

The pig- mentation of mol ars , lack of re d uct-

ion in talon id of M, an d t he shape of facets in the ar ti cular proces s

reason ably refer the collected remains to genus

Sorex L.

Its speci f ic id en -

tification , however, is st ron gly hamp ered by the fragmentary condition of

t he Kadzielnia specimen s a nd cannot as y et be definitely determined. The

sh ape of its articular process bars its as sign m en t to the group of S.

alpinus

Schinz, while the moderate d imensions shut off any comparability with

distinctly large forms s uch as e.g. S.

savini

Hinton or very small ones

such as S.

minutissimus

Heim de Balsac. This leaves us with the group

of modera te ly siz ed species, of th e dimensions of

Sorex araneus

L., but

from th is spec ies our fossil spec imens d iffer i n slightly smaller dimensions

12 ^{KAZIMIERZ KOWALSKI}

and

in

different shape of the coronoi d process. On the other hand, t h ey closely approach

S. l'untonensis

Hi nton

,

a species wid ely distributed over Europ

e

in early Pleistocen e.

Sorex sp.

(fig. 3)

Material. -

4 fragmentary mandibles with preserved processes and

McM3

from the deposits filling up the doline .

Description . -

An te r io r margi n of t he coro noid p r ocess meets the ramus of mandible at a ne arly right angle w it hout being c urved forw ard at its uppe r part. This p r ocess

is

b ro ad er and more rob ust than th a t in

S. minutus L.

Fossa pterygoidea h igh

,

tri angula r. The bo n e bridge betw ee n the facets wide, slightly incised, t he general sh ape o f this pr ocess resembling t hat

in S. minutus L.

Tooth po in ts p igmented to a reddish-ye llow co lour . M

3

five-c usped, he nce its talonid is n ot red uce d,

Dim ensions of mandi b u lar

fragments (in m m):

Mandibles -~ 1 2

I.

3

height of mandible on the inner side be - I

low M₂ 0.9 0.9 I _1.1

thickness of same 0.7 0.6 0.7

M₁ length 1.4

M₁ width on trigonid 0.6

M2 length 1.2 1.3

M₂ wiath on trigonid 0.5 0.6

M_alength 0.9 0.9

M, width on trigonid 0.5 0.6

Systematic position.

- T h e presence in M

3

of five cusps suggests the assignment of th

e

collected fragment to the subfa mily of Soriciriae, while on the

sh ape of facets in the arti cu la r proces s

we may reasonably refer it to genus

Sorex

L. The fragmen

tary

condition of the available material does not permit to

5mm

determine its specific position. The collected mandible resembles

in size

that of

S. minu- tu s L., from which it slightly

differs in the _ ~ propor tions of the coronoid process. The Ka-

_

dz

ielnia

sp

ecimens

are, in any case, referable

F'_19. 3_{. -} S_orex. _sp., f_ragment

to

'a

larger form than that described by the

of mandible ; side an d top

presen t writ

er

from Podlesice under the

view.

name of

Sorex sp .

(Kowalsk

i,

1956)

,

but

sm alle r than S. kennardi

H

in ton

from

th e

P leisto ce ne of England.

PLEISTOCENE FAUNA OF SMALL MAMMALS

Gen us Ber em en dia Kormos, 1934 Ber em endia fissidens (Petenyi, 1864)

(fig. 4)

13

]'864. Crossopus fissi de ns; S. J. Pet en yi, Hatrahag y ott munkei, p. 60, pi. I, fig. 5.

1911. Neom y s fi ssidens (Petenyi) ; T. Kormo s, Can is (Cerdocy on )..., p. 170, pi. 7, fig . 1-3.

1913. Neomy s fissidens (Pe L) Kormos ; J. Ehik, Die pr ag taziale..., p. 140.

1930b. Beremendia jissidens (PeL) n.g. ; T. Kormos, Beitra ge ... p. 57.

1930a.Neomys (?) fissi den s (Pet.) Ko rmos ; F. Heller, Eine Fo r est-Bed-Fauna ..., p. 254-258, pi. 15, fig. 1-3, text-fig . 2-4.

1-933. Ber em endia fissidens (Pet.) Kormos; F. HeNer,Ein Na cht ra g..., p. 61-62.

1934a.Beremendia fi ssidens (Peten y i ): T. Kormos, Neu e Insec1enfresser... p. 299-301, fig. 33.

1934. Berem en d ia fiss idens (Pet eny i): G. Brunner, Eine pragla ziale Fau na..., p. 311, pi. 6, fig. 6- 8, text-fig. 6,7.

1936a.Berem endia fissidens (Pete nyi); F. Hell er, Ein e ober plioc ane Wirbelt ier- fauna..., p. 107-108, pi. 7, fig. 1-2.

1941b.Bereme n d ia fissidens (Petenyi) ; M. Kretzoi, Weiter e Beitrage ..., p 110 1948. Beremendia fiss idens Petenyi; A. Pasa, I MammiferL..., p. 14-16, fig. II, 1-4.

1949. Beremendi a fissidens Pet enyi: lVL Friant, Les Mus arai gnes..., p. 256-257, fig. 17.

1956. Ber em en dia fissidens (Perenyi): K. Kowalski, Insectivores...,p. 349.

1956. Beremendia jissidens (Petenyi): 'M. Kretzoi , Di e altp leistoz anen..., p. 164, 169- 171,176,,1<80, 183, 184, ,187, 192, 193, 195, W7-201, 203.

Material. -

Fragment of mandible with M

cM2

and damaged processes, fragment of mandible with complete p

rocesses

and with M

2 ,

f

rag ment

of mandi ble with M

2-M3,

four detache d mandib ular incis ors, f

ragme nt

of maxilla with

P4_M2, fragment of m

ax ill a with

P

3_M3, two

·de tached

incisors II an

d

a detached maxil

lar

molar, all from material

filling

up

t

he

doli

ne.

Descript ion . - Anter ior mar gin of the

coronoid pr ocess

m

ee ts t

he

ra

m us

of 'the mandible at an o

bt use

angle. The man dibular body very

m

assive.

'T h e upper

fa

cet

of the articular process placed obliquely in relation to

the lower fa

cet,

the b one

br id ge uniting them

wide, concave.

Molars with points pigmented to a dark brown colour. Mandibular incisor sharp,

long,

with tip prominently curving forward and

s

t r on gly coloured, showing a distinct groove on the inner wall, without incisions

.on

the margin. M

3

with a r

educed

small talonid, without metaconid.

II large, with tip distinctly bifurcating and thus two-cusped. p

3

small,

protruding lingually from the tooth-row. P

4_M2

large, with pointed

,

dark-

.tin ted

cusps

.

14 KAZIMIERZ KOWALSKI

5mrn

Dimensions.

- Length of Mj-M

,

6.4 mm, height of mandibular ramus on the inner side, below M, 2.6 mm. Length of P

4_M2

is 6.7 and 6.6 mm respectively for the two cons idered specimen s.

Systematic position. -

The pigmentation of the molar points, to- gether with the reduction of the talonid

in

M, and the large dimensions of the r

emains

clearly indicate their assignment to genus

Beremendia

Kormos which embraces on

e

speci es on ly, namely

Beremendia fissidens

(Petenyi) record

ed fr om

a number of

earl y

Pleistocene localities in Europe. In 1955

, 1.

G. Pidoplitschko mentions the species

.Blarina ucrai- nica spe-

c. nov."

,

in the early Ple i- stocene faun

a fr om Czortk6w (we-

stern Ukraine).

It

rather seems th

at

P

idoplitschko

has committed

an error in identify ing

a specimen

of Beremendia fissidens

(P

etenyi)

with the American genus

Blarine

Gray, a form thus far never enco- untered in the Old World. Mention

sh oul d be made

h

ere

tha

t

C. C.

Young (1934) described fr

om Cho u-

koutien in China a new species ,

Neomys bohlini Young,

said to be

Fig. 4. - Bereme ndia fissid ens (Pet eny i),

d

istinguished

by a reduced talonid

fragm ent of mand ible; side and top view. . .

In

M

a•

This character bars the assignment of the describ ed form to genus

Neomys

Kaup and brings it nearer to genus

Beremendia Kormos.

Hence

it

is not out of the question that this genus had a wider range of distribution within the early Pleisto- cene than heretofore supposed.

G

enus Petenyia Kormos,

1934

Petenyia hungarica Kormos,

1934

The sy n on im ics have been give n in K. Kowalski's pape r of 1956, Insectivores...^r- p. 352; also:

1956. Petenyia hungarica Kormos ; M. Kretzoi, Die altp le is t ozane n Wirbeltierfau- nen..., p, 164, 169, 170, 175, 184, 187.

Material. -

Incomplete mandible w

ith

Pi-M , and with pres erved

processes, collected from th

e

material fill ing up the fissure in the NW

part of the quarry, also a mandible with Mt-M

_a,

fragment of mandible

with M

2-Ma,

detached mandibular inc

isor

and fragment of mandible

without dentition,collected from the material filling up the doline.

PLEISTOCENE FAUNA OF SM A L L MA M M A L S 15

Description.

- Tooth-tips strongly p igmented to a dark-brown colour.

M, with talonid strongly reduced, with one cusp only, lacking th e metacon id.

Dimension s. -

Length M1-M

g

3.7 mm and 3.7 mm resp ectively, height of mandibular ramus on th e inne r side below M

2

1.5 a nd 1.6 m m respectively.

Systematic position.

- The ab ove mentioned characters of the col- lected mandibles clearly indicate their assignment to g enus

Petenyia

Kormos. As has been stated b y th e present writer in his paper on the f auna from Podlesice (Kowalski, 1956, p. 353), the sp eci es d escribed by Kretzoi (1943) and Pasa (1948) are so poorly differentiated t h at th ey ought probably to b e r egarded as sy n ony mo us with P.

hungarica

Kormos.

It

is to this species, as the on ly r epresentative of gen us that the Kadzielnia specim ens are r ef erable.

Chiroptcra

Blumenbach, 1779 Family

Rhinolophidae

Bell , 1836 Genus

Rhinol oph u s

L aceped e, 1799

Rhinol ophus

d.

ferrume quinu m (Schr

eber, 1774)

Material. -

Mandibula r fragment with M

j-M ,

an d w ith damaged processes , some fragments of m axilla an d detached t eeth, all collected from the material filling up the d oline.

Description.

- The preserved dentition as well as fragmentary processes d

ispl ay

a structure ide n tical with that in sp ecimens of the Rec ent

Rhinolophus ferrumequinum (Schreber) from central Eu

rope and

in

early Pleistocene specimen s from Podlesice (Kowalski, 1956). The t eeth in th e collected mandibl e are s tr on gly used, indicating that they belonged to a very old in d iv id ual.

Dimensions.

- Length of M1-M

^g 6.1

mm, height of mandibular ramus on the inner sid e below M 1 1.9 mm.

Systematic position.

- The co rrectness of the assignment of the

Podlesice s pecimens to genus

Rhinolophus

Lacepede is beyond doubt,

while its d

im

ensions and agree me n t with r ecent s pecim en s furthermore

refer th em to sp ecies

Rhinolophus ferrumequinum

(Schreber)

.

As has

been sta ted by the presen t writer with more detail in his paper

published in 1956

,

evidenc e for t h e identity or differences between fossil

and recent sp ecimens cannot be obtained until more copious material has

been made av ail able for comparative studi-es.

Hi ^{KAZIMlERZ KOWALSKI}

Family

Vespertilionidae Gr ay , 1821

Subfamily

Vespertilioninae

Miller

,

1879

G

enus Myotis

Kaup, 1829

Myotis

cf.

exilis

Heller, 1936

(fig. 5)

Material.

- 6 fragm

ents of

mandib le w it h P

I-M_a, wit

ho ut processes, coll

ected

from the material filling up

the doline.

Description.

- The

alve oles indicate

the presence of

t

h ree one-rooted premolars. p

. is

slightly

elonga te d, the

tip protruding ab ove

th e tips of

other molars. M

3

does not show an

y stronger reduc tion

of talonid. For.

mentale

betw een PI

and P

2

of rather large size.

Dimensions

of mandibular fragm

en ts

(in mm

) - see ta

ble below.

Systematic position. -

The presence of three on

e-root ed

premolars in the mandi

ble

and the con

siderable

height of P

,

clearly indicate the

assignm en t of the collect ed

remains to genus

Myotis Kaup.

The small dimensions of these r

emains

r

estrict

the d

iscussion

conc

erning

their id

entification

to the

s

m allest forms of this ge- nus, to say

:

M.

exilis Heller ,

M.

ins ignis H

ell er

,

5",rn

M.

daubentoni (Kuhl),

M.

capaccinii (Bon ap ar -

t

e),

M.

mystacinus (Leisler in Kuhl). Of these

forms M.

mystacinus

(Leisler in Kuhl) shows

~.. . ' .'

altogether different proportions of

p.,

while

~~

analogies are observable be

tween the studied

species and the group of forms M.

daubentoni-

Fig. 5. - Myotis cf. e:l.'ilis

Heller, fragment of man- exilis -capaccinii.

T

he K

ad zieln ia specimens are

dible; side and top view.

somewhat larger than M.

daubentoni (Kuhl)

Mandibtes - -- - -, 2 3 4 5 6

I

side I

height of mandible on the in ner

below M2 1.6 1.5

-

^- I_I ^{- -}

thickness of same 0.7 0.7 0.7 - - -

symphys is length --

-

^- 2.1 - 2.0

p. len gth - -

I ^-

^{0.7 0.7 -}

p. width ^{- -} -

I

0.6 0.6 ^--

M₁ leng th ^-- 1.3

I

^{1.3 1.3 1.3 -}

M₂ leng th 1.4 1.3

I

- - - -

M₃ len gth 1.2 -

-

I

- -

-

PLEISTOCENE FAUNA OF SMALL MAMMALS 17

an d M. ins ignis Helle r with dimensi ons mor e 'Closely appr oaching to thos e of M. exilis Heller and M. capaccinii (Bonaparte). A ver y certain de t er- mination of their sp ec ifi c posi tion is imp oss ib le ow ing to the fr ag men t a ry condition of the avail able spe cime ns, the agreement of dimensions, how- eve r , as well as the proportions of P,I sugges t their assi gnment to the ea rly Pleistocen e species M. exilis Helle r .

Myotis sp .

(fi.g 6)

Material. - 16 fragmen ts of mandible witho ut processes, cont ainin g P2-M3, 2 fragments of maxilla with MI_M3, collected from the mater ial filling up the dolin e .

Fig . 6. - Myotis sp., fragmen t of mand ible; side an d top vi ew .

Jmm

slightly elongated. P3 Description . - Alveol e of incisor oval-s h ap ed,

one-r ooted, moderately la r ge. p. with height alm ost that of molars, sub q ua d - rate in outline. Tal on id of M3 not re- duced.

On their dimensions the fragment of maxilla is reasonably referable to the same species. Molars without protoco-- nulus.

Dim ension s. - Length of P.-M2^4.0 mm, length of M1-M3 (in another spe- cimen) 4.1 mm, height of mandibular ramus on th e inner side below lVL 1.7 mm in two specimens. Two spec im en s of P, show the length to be 0.9 an d 1.0 mm, the width 0.8 and 0.75 respecti - vely.

Sy st em atic position. - Thestructure of molars, and more particularly so the presence of three one-rooted premolars, as well as the height of p. , indicate the assignment of all the collected specimens to genus Myotis Kaup. The app r oxi m a tely similar dimensions suggest that in spite of certain differences all the remains belong to one species. Their specific identification,however,meetswith difficulties in viewof the fr ag m en t a r y material. On their dimensions they are referable to one of the mod e r at ely sized forms, though they all displ ay a slig ht dissimil a r-ity 'n the proportions of P,.

.Acta Pat a eon t otogtca Polo n rca - vel.Ill!I 2

KAZIMIERZ KOWALSKI

Lagomorpha Bran dt, 1855 Family Leporidae Gr a y , 1821 Subfamily Palaeolaginae Dice, 1917

Ge n us Hy polagu s Dice" 1917 l-Iypolagus brachygnathus.Kormos , 193 4

(fig. 7, 8)

lS30b . Lep us brach y g nath us n.sp . (n ome n nu dum); T. Kormos, Beitr age...

1934b.Hy p o! agus brachygnath ll s n. sp.; T. Kormos, Zur Fra ge ...., p. 75"Dig. 2 a-a. 1934. Hypolagus brachygnathlls Kormo s; J. J. A. Bernsen & A. Schreuder, Eine

Rev is ion..., p.84.

1936a.Hypolagus brachygnathus Ko rmos ; F.Hell e r, Eine oberplioc ane Wirbeltier- fauna ..., p. 13'7-139, fig. 1-2.

1937. Hypolagu s brachygllathlls Ko rmos; A. Schre uder. Hyp olagu s.... p. 225-229, pI. 2, fig. 1-2.

1941a.Lagotherium brachygnathllm (Kormos); !VI. Kret zo i. Die unterpleist ozan e Saugetierf'auna.... p. 323-324, fig. 7.

19:54. Lagother ium berem endens e (Petenyi); M. Kretzoi. Berich t..., p. 248.

1956. Lagoth erium berem endense (Pet enyi); M.Kretzo i, Die al tp leis to zanen Wir- belt ie rfaune n.... p. 160, 162, 164, 169. 170, 176, 179, 184, 188, 193, 195. 198 , 201, 208.

Material. - 60 fragments of mandible with different parts of de n - tit ion, 3 ma x ill ae with complete row of molariform teeth and numerous fragments of max ill a e, nu merous detached incisors, upper and lower pr e- molars and molars, numerous, mostly fragmentary bon es of the skeleton, all collected from the materia l filling up the doline.

Descri ption. - Man dib ular incisor more st r ongly curved than that in Le pu s euro paeus Pall as , ex t re m e ly massive and broad. Dias tema very short. On the outer side of P"two enamel folds: the anterior fold shallo w, the pos t er io r penetrating to mid-widt h of the tooth. No enamel folds on the inner side of the tooth. P,-M" dis t inctly more massive than those in Lep u s europ aeus Pallas:

The first maxillar incisor flattened out; over one third of its surface there is a distinctly marked groove, shallow and without cement. The maxillar premolars more massive than the corresponding teeth in Lepus europaeus Pa ll a s. On the anteri or su r fa ce of p2 a deep central incision, another one on the outsi de. The re-entrant enamel folds on P3_M2 ex t end over two thirds of their width. M" str on gl y reduced'but always present.

Fo r dimensions of mandible see table on p. 20. For dimensions of premolars and molars see table on p. 19.

Sy st emat ic position. - In 1929 L. R. Dice was the first to attempt a reasonable division of the Leporidae - a family whose systematics presented considerable difficulties - by splitting it up into three subfamilies on the structure of the third lower premolar. Later on

PLEIST O C ENE FAU N A OF SMALL MAMMALS 19

Mandibles- -

, I

^{2 3 4 5 6}

I

lengthof toot h -row

I

_15.0

pt length 1.7 1.7

pt width 3.9 3.8

p' length 2.9 2.9 2.7

p:' width _{5.5 5.5 5.4}

pi leng t h 2.9 2.8 2.7 2.8 2.8

pi width 5.7 5.5 5.5 5.3 5.7

M' len gth 2.9 2.9 2.6 2.7 2.7 2.5

M' width 5.5 5.2 5.6 5.0 5.0 5.2

lvrt len gth 2.4 2.3 2.3 2.6 2.3

Mt width 5. 1 4.8 4.4 4.6 4.9

M' le n g th 1.1 1.2 1.1

M:' wid t h 1.9 2.0 1.8

20mm

Dice himself

d

ecided

that

,

in conside ration of the presence of numerous

inte r

med iate forms b

et w een

the s ubfam ilies Palaeolaginae and Arch

aeo- la

ginae,

that sub di v isio n ought to

b

e con

fined

to on ly two

cl

early

d

istinct subfa milies, na

mely th

e

P

al aeol a-

ginae an

d

t

he Lep ori n ae. In

the fo

r mer, th

e fol

d

ex ten d ing

from

th e outer edge of

P, re ach es ap pro-

ximately to the mi

d -width of

th at toot h,w

hile

in th e latter it extends as

far to

the in ner edge.

On evi dence

of

th

e

sep a r a t io n

made

by Dice, Kormos (1934

)

diffe renti a ted three genera

of fos- sil

L

ep orides fro

m th

e ea r ly Plei- s

tocene of

central Europe : Pliola- gus

K

ormos, Hypolagu s

Dic

e an d Lepus

L. The firs t named form

,

r

ep r esen ted

by spe cies

Pliolagus beremenden sis

Kormos, w

as

said

to

be dist ingu ished, in

ad dition to its s

ma ll d

im ension s, also

by the

Fig. 7. - Hy po! ag u s brachygllat h us Kor-

presence of a v

es tig ia l

re-en tra nt ma s; A inc om ple te ma n d ible, B maxilla.

fold on

t

he outer edge of P

a•

T

his fold is not, howev er

, always dis ce rnible.

The seco nd

of the

her

e named early

Pleis to ce ne Lagomorphs is by Kormos (1934)

re

ferr ed

t

o

the fossil

Am

e r ican genus Hy polagu s

D

ice, w

hi ch h

e

called H.

brachy-

Hypolagus brachygn athu s Kormos Dim en s ion s of mand ibular fra gments (in mm)

~o

Mand i bles } 1 1

I

2 I 3

I

4

i

⁵

¹

⁶

^I

⁷

I

8

I

9

!

10

111

^{112 113 114 \15 116 117 118 [19 120}

~>

N~ [;i :0 N

o~ ::;::

~(JJ

~ 17.5

4.214.0 3.513.3 6.0I6.1

4.213.9 3.413.0 3.513.0 15.4114.5

4.014.0 1.0 2.3 15.0115.7 3.8

- I 30

i .

i 4.0

3.0

- 4.0

- 3.113.2 -

I

1-

-1 --

3.613.5I3.4I 3.413.4 3.313.5I3.0I 3.513.2 15.5

1.812.1

I I

15.0 -

1 -

^I

6.2 6.0, - -

-1 --

1.⁷

1 -

I - 13.3

I --1 3.0

3.7!3.0

I

3.0I 3.213.0I 3.3

3.513.3I3.4I 4.1I4,2I3.814.1 1.8

-I I

2.913.1I 3.0I 3.312.9I2.913.113.0I 3.3 3.914.0I3.613.413.6 14.214.2 13.8 14.1 3.0 3.3 2.9 3.0 3.3 3.1 3.1 4.014.0 13.613.5I3.814.2I4.113.914.2

3.8I3.9I 3.8 ._- 1.9I 1.812.1

1 -

3.213.113.0I 3.013.1I2.7 3.7I3.213,4I 3.4I3.3 3.5 3.6

3.1 3.4

3.81- 4.0 4.0I 4.0 3.1

I

^{3.3 3.3 3.112.8}

3.9I 4.1 4.0 4.013.9 3.0

I - ;

3.3 3.1 3.0 3.81-

I

3.7

1.9 - 11.6 3.3

3.914.0

I I ! ^{I I}

^!

19.5119.0118.0 - I - i - - - -- ^{i - - - -} I - - 1--

,20.5 - I :17.01 - , - - -

- 1 - - - - - -- ^--1

^{18.4 !17.8}

I I i i I I 1 I

114.6 14.8115.0115.0 14.5113.7114,0 13.7^{1 1 3 . 9} 14.2114.9114.2114.0 5.8 6.215.816.4 6.0 5.9I5.9 6.4 6.2 5.8 5.816.0

I

6.0

14.8 14.8 - 1ts.5 15.01.14.5 - -

-1-

^{I -}

^I

I3.7 -- - 3.71- - 1- -

I - - -

I3.1 3.4 3.313.7I3.3I3.3I3.113.1I 3.2 3.0I3.4

3.0I3.1 4.014.0 3.2 3.3 im p ression

an d M2

Wid th I P³ length P₃ width p{ len g th P₄ widt h M₁ len gt h M₁ width M₂ length M₂ width M₃ length

Thick n ess of same

He igh t of mand ible between PI Dis t a n ce f.ment ale to masseter

Length over P:l-M 2(on crowns) Lengt h of dia stema

PLEISTOCENE FAUNA OF SMALL MAMMALS 21

gnathus Kormos. It

is of the size of the European species L.

europaeus

Pallas, being distinguished, besides a completely different structure of P

g,

also

by considerably shortened mandible. The last named form, genus Lepus L., occurs in younger early Pleistocene faunas of central Europe,

while in the west of Europe it is known already from the Pliocene.

On the structure of the third lower premolar the above describ- -ed Kadzielnia fossils may undoubtedly be referred to the subfamily Palaeolaginae. Their massive, stout mandible points out to species

Hypolagus brachygnathus Kormos as is also confirmed by the great

conformity of dimensions within specimens from Hungary and Holland (Schreuder, 1937). A short diastema and strongly curved incisor are here particularly characteristic features.

The mandibular dentition of H

. brachygnathus Kormos has never,

thus far, been described.

Kretzoi (1941) uses for the here considered species the name of

Lagotherium brachygnathum (Kormos), stating that the generic name

8~SJ

^5mm

Fig. 8. - fIypolagus brachygnathus Kormos, P3; A-C from be- low, D from above.

of

Lagotherium Croizet &

Jobert has long been applied to the Leporidae with a short massive skull, recorded from Perrier. Kretzoi, however, does not say whether the structure of the third lower premolar in the Perrier specimens was identical with that in the Hungarian specimens, while Schreuder (1936, p. 227) referred the Lagomorphs from Perrier to genus

Lepus L. This problem must,

therefore, be still considered an open quest-:

ion. In his papers published in 1954 and 1956 Kretzoi introduces still another name, namely that of

Lagotherium beremendense (Petenyi)

which he regards as synonymous with

Hypolagus brachygnathus Kormos.

This opinion, however, calls for evidence, thus far not supplied by

Kre

tzoi .

The mandible, as figured by Petenyi (1864, pl. II, 1) seems to

differ from that in H.

brachygnathus Kormos, in

having its incisors

less strongly bent. There is, however, no description of it.

lmm

KAZIMIERZ KOWALSKI

Genu

s Pliolagus

Kormos

,

1934

Pliolagus d. tot hi Kretzoi,

1941

(fig. 9, 10)

Material. - 5 fragments of mandible, n

um

ero

us d

etached in cisors,

m

andibular an

d m

axillar

mola rs a nd premol ars

f

rom

t

he m

a terial recove re d f

rom

the kar st

doline.

Descrip ti on. -

Mandib le s mall, deli ca t

e.

In

ciso r

n

arrow, slightly c

urve d. On th

e outer s

ide

of P3

tw

o depressio ns

passing in to r

e- entrant fo

lds

filled by ceme nt.

Of the s e the

an terio

r

one sha llow, the

po

sterior

deep , r

eachingto m

id-wid th

of tooth . On t

he

inner side of P^3,

a dep r ession usu-

ally occurs opposite to the p

osterlor,

in

ne r r

e- en tran t

fold. Bo

th the outer and inne r edges of P³

di

splay

distinctl

y prot r udi ng

ridg es. P

r M 2

are

sm all, abo ut the same siz

e as thos e in

Orycto-

~ig. 9. - Pl i ola gu s d . tothi Kretzo i lagu s cuni cu lus L

. Mandibular in

ciso r

incomplete mandible.

strongly

fl

at ten ed.

A groove lacking cement

runs over one third of its wid th.

Dimensions

of mandibular fragm

en ts

(in mm)

:

Mandibl es 2 4 5

distanc e f. men tal e to ma sse te r

im p ress ion 16.0

length of diastema 14.0

height of mandible betw een P4

and M₂ 13.3 13.2

thickness of same 5.5 5.5

length over P_{3-M 3} (on crowns) 13.7

width I 2.2

P₃ length 2.9 3.0 2.7 2.6

P₃ width 2.6 2.9 2.4 2.6

P4 length 2.7 2.6 2.6 2.6

P4

^{wid t h 3.5 3.5 3.5 2.9}

M₁ leng th 2.6 2.9 2.7 2.6

M₁ width 3.3 3.5 3.2 2.9

M₂ len gth 28 3.0 2.7 2.7

M₂ width 3.3 3.4 3.1 2.6

M3 length 11