Pliocene insectivores from Węże

A CT A P A L A EO N T 0 LO G I CA P 0 L O N I C A

Vol. IV I 9 5 9

ANDRZEJ SULIM SKI

PLIOCE NE INSECTIVORES FROM WE;ZE

Stud y on the Tertiary bone brec cia fa u na from

W~ze

near Dzialo szyn in Poland

PART XI*

l';o. 2

Abst r act . - This paper con tain s a descri ption of 18 insect ivore speci es recovered fro m the bon e bre cci a at We ze nea r Dzialoszyn (prov ince of L6di). The desc ribe d form s include four ne w spe cies an d one new gen us: Erinaceus samsonowiczi n. sp., Blarinoides mariae n. gen., n. sp., Neom ys soriculoid es n. sp., Suncus zelce u s n. sp. The write r exp la ins his me thod s of work an d discusses the age an d origin of the brecci a, as well as the probl e m of the pal a eogeographic distr ibu tion of some spe cies

fou n d in the We ze bon e breccia.

INTRODU CTION

The insectivor e bone materi al e n trus t ed t o the w riter to be de scribed co mes f ro m t he bone breccia discovered at t he village of

W~ze

near Dzia- los zy n . Work on the pr eparati on of this material has been carried out during t he la st three years in the Palaeozoological Laboratory of the Polish Acade my of Sciences in Warsaw , It has been su pple me n t ed by mater ial fro m the same locality , handed over to the wri t er in 1955 by the l ate P rofessor E. Wilkus of Lublin' Universi t y , al so by that prepar ed in the Depar tmen t of P alaeoz oology of t he Wroclaw Institut e of Zoolo gy .

The writer 's most sincere t han ks are her e conveyed to Professor R oman Kozlowski f or hi s valuable advice and suggestions throughout t he preparation of this paper , also for his crit ica l comments after its com- pl etion ; t o P rofessor J an Sams on owicz for reading through th e ch apt er

*

Pa r ts I-V - see Acta Ceol. Pol., vol. II-V/1952-55; parts VI-X - Acta Palaeont. Pol., vol. I-IV/1956-59.

120

AN D R ZE J SULIMSKI

concerned with the st ra tig r a phy and or igin of t he W eze bre cci a ; to P r o- fess or Zbigniew Ryzi ewicz a n d hi s co-worker s in W r oclaw fo r help in pr oviding s u pplem en tary materials and information a bo ut new find s in Weze ; to ,P r ofessor Augu st Dehnel fo r t he permission to e x am ine t he rich Bialow ieza coll ection of rece nt shrews, and for di scussin g man y var iation pr obl ems in t his ani mal grou p ; to Professor Claud e W. Hibbard of t he Mich igan Univ ersit y for t he generous gift of co m parative m a terial cons ist ing of rece nt an d foss il America n shrews, a lso for hi s valua ble com men ts on the new spe cies and ge n us Blarinoid es mariae. T he au thor expresses hi s sincere gratitu de to P r of. A. H alicka , Di r ect or of Museu m of the Ear th in Warsaw , for t he loan of mat eri al for in vestigati on s .

Wa rmest t ha n ks are also due to Dr. K. K ow al ski for suggestions co ncerning fossi l insec tivores and for the ac cess to his ric h libra r y in Kr a- k6w; to Dr . W. S er afinski fo r ki ndly le ndi ng co mparative m at erials of recent hedgehogs ; to D r . J . Kulc zyck i for h elp in a na lysing pr obl ems of dental term ino logy and structure of cranial an d m andibular ele me n ts , as w ell as for pr ov iding t he missing literature items ; to Mrs . J. Humnicka fo r doin g t he En gli sh trans lation of the pap er , and f in ally to Miss M . Czar- nock a fo r t he photog raphy .

C

ON DITI ONS

OF

THE DEPOSIT

The bon e brecci a fro m which so man y in sectivor e remains have been re covered comes fr om a karst doline in t he side of a hill , called Zelce , at the village Wftze nea r Dzialoszyn . This is the n or thernmost point of the Kr ak6w-Wielun Jurassic Hi ghlands. P ap ers by J . S amsonowicz (1934 ) . an d K. K owalski (1951) give a detailed ge olog ical desc r iption , section of t he karst dolin e wit h the breccia , an d t opogr a phy of the nei ghbourhood of th e Zel ce hill.

The karst doline wi t h Ter tiary fa u na, abo u t 4 m in dep th a n d 5-6 m in di ameter , was filled in by la yers of two t ypes : re d bed s wi th a larg e ad m ix tu re of w eathered clay and b ean ore (terr a ro ssa) , and light beds , t he so-c alled "grey" breccia , strong ly calcifi ed . Thes e bed s, about 0.5 to 1.6 m in thicknes s , were re pe ated al ternately. The upper part of the doline was fill ed by cine reous soil an d abou t 1 m of calcareou s ro ck debris con t ain ing a mixed Plio-Pleistocen e fauna . The lower p ortion consis ted of thin lenticular, r ed and "g rey" breccia beds , an d of barren beds of lime st on e crust and calcite . Th e bot tom breccia la yer, probably of r ed co lou r at ion, rest ed directl y on Jurassi c limes ton e (sec ti on fig . 1).

. The bulk of the breccia (ca . 11 tons ), ex cavate d in 1933 by Professor

Samsonowicz , is now deposi t ed in t he Muzeum Ziemi (Museum of t h e

Ea r th) in Warsaw. Th e m at eri al s, at fi rst systematically so r ted an d stored

PL IOCENE INSECTIVORES 121

under t he gu ida n ce of P r ofessor S amson owicz, are now badly mixed up , ow ing to cond itions prevailing dur ing the war and im med iately aft er it . T he re main der of the Wf:ze brecc ia was ex cavated in 1955 , and deposi ted in t he Inst it u t e of P alaeozoology in Wroclaw.

A part of carnivorous remains yielded b y t he W eze breccia have been worked out b y J . Stach (195 1, 1952, 1954 , 1957), those of reptiles - by

Fig. 1. - Longitudinal section of karst doli ne wit h bone breccia (after J. Sarnso- nowicz, 1934)

1 soil. 2 limestone debris, 3 red br eccia with bean ore (terra rossa), 4 ,~ 5 "gr ey " br ecc ia.

between "grey" brecci a bed s th in lenticula r red breccia la yer s , 6 pinki sh -yell owi sh ca l ca reous crust, 7 red cla y with bone breccia and bea n ore, 8 ca lc ite, 9 limestone incr ustations and

cal cit e, 10 Ju r a ssic limes t on e.

M . Mlyn ar ski (1953, 1955 , 1956), while T . Czyzewsk a has published note

on some ungulate (1958). T he remaining mammalian fa u na, including

insect ivores an d ro de nts, is bei ng worke d out in Warsaw, t he ot h er

ung ulat es - in W ro claw . Stu dies on ca rnivo res and bats are carried on

in the Zoological In s ti t u te of the P olish Acade my of Sciences in K r akow .

122 ^ANDRZEJ SULI MS K I

AGE OF THE WF;: ZE BR ECCIA

The first ge olog ica l investigation s of t he Wf:ze breccia (Samsonowicz , 1934) h av e sho w n t he occurre nce there of a nearly complete Pli ocene section. The lower bed s of the breccia h av e been dated as Lower Plioc en e , perhaps the Upper Miocene ; th e upper bed s weathered and calciferous rock de br is - as Upper Pliocen e , possibly as early Pleistocene .

The su p pos it ion of Samson owi cz (l. c.) are , on the whole confirmed by later, palaeon t ologi cal in vesti ga ti on s . The carnivorous Arctomeles pliocaenicus Stach is never reco rded fro m t he Mioc en e . Stach (1951) believes tha t form to have bec om e ex ti nct at the clos e of the Pliocene ow ing to the cooling down of climate . Th e sm all bea r Ursus wen zensis S tach , 1952 has been referred to t he Middle or Lower P lioce ne , while the pos ition of Ny cter eutes s p. (St ach , 1954) has li kew ise be en placed in the Pliocene . Finall y , A griotherium interme dium Stach ; 1957 must have lived from the Upper Miocen e to the Up per Pliocene. S tudies on t u r tl es and lizards (Ml ynarski , 1953, 1955 , 1956) have pr ovided evidence for t he P li ocene ag e of our br eccia . Teeth of a young r hi noceros Dicer- or hinus megarhinus (Christol) , described by Czyzewska (1958) , end up the present list of publications on the Wf:ze fauna and p rovide ad ditional sug ges t ions to confirm the early Plei st ocen e age of t he up per breccia layers .

The insectivores , here reported upon , cover th ree fa milies, with 10 genera and 18 speci es . Though age determination of the breccia on in sectivore ev idence only , is inadeq u a te , yet i t permits t h e confirmation of earlier inferences an d the assignment of Pliocen e ag e to the greatest part of the Wf:ze breccia. At the sa me time it seems reasona bl e to conclude tha t its low er layers r ea ched int o the Upper Miocene.

On the one hand this is indicated by species which exhibit many features in common with Mi ocen e s pe cies and bel on g to th at group of xerophilous forms e xisti ng in ar id ste ppe e n viron men t: Erinaceus samso- nowiczi n . sp., partly Desmana nehringi Kormos , remains of Galemys(?) sp., Blarinoid es mariae n. ge n ., n. sp., ex tens ively N eomy s soriculoid es n. sp. ,' pa rtly Beremetulia fisside ns (Petenyi) , r emains of Crocidura s p ., and both species Sun cus pannonicus (Kormos) and Suncu s ze lceus n. sp. On the other hand, the remaining s pecies , char acte r ist ic of moist palaearctic belts , with an ad m ix t u re of some of t he abo ve m en ti on ed spe cies , distinctly date t he breccia as U pper Pliocene or early Plei stoc ene.

As regards the new genus Blarinoides, it is mos t likely a Miocene

r elict. It probabl y became e x t inct owing to the cooling of climate towards

the clos e of t he Middle Pli ocen e, and did n ot leave descendants . A sim ilar

situation must have occ urred for Suncus zelceus n . sp .

PLIOCENE INSECTIVORES 123

Out of t he 18 s pe cies here re ported upon only t hree have

survive~

t h us fa r , the others are ex tin ct. Of the genera only three are ex t inct,. the o t h e r sev en ar e st ill living. The latest spe cies of the extinct genera mo st

likely su rvive d as late as the first Interglaciation period.

PALAE O G E OGRA P HI C DISTRIBUTIO N OF SOME OF THE WF;ZE INSE CTIVORES

Genus Er ina ceus L. is to - day enco u n te re d within Eurasia on ly , and the

Wf:~:e

fo rm E. samson owiczi n . sp. has an A siatic affinity. A sim ilar sit u a ti on is noted in the case of fossil s pecies b el onging t o ge n us Talpa L.

The su bfa m ily Desmaninae Thomas is now en cou n tered within a r est r icte d are a: Gal emys Kaup in the French P yr enees and the Ib erian P eninsula;

Desmana Giild. in t he sou th-easte rn parts of European Ru ssia . During the Pliocene t hey must have ranged ov er considerably wider areas , including central Eur ope .

Genus N eomys Kaup , though recorded already fr om the Pli ocene, does not seem to have been ver y a b u n dant during that pe riod , as is s uggeste d by t he few finds and publications. But copious populations of N. soriculoides n . sp . from Wf:ze distinctl y contradict them. It is not out of the question that th is species , re la te d with the south Asiatic genus Soriculus Blyth, may have its representatives also in other European sites.

Genus B er emendia Kormos occurred in grea t abundance during the Plio-Pleistoc en e over vast ex pa nses of sou the r n Europe , reaching as fa r north as abou t 51st or 52nd de gree of latitude. Its western range was roughly limited by the river Rhine and the w estern Alps.

The su bfa m ily Crocidurinae Milne-Edwards comprises a diverse group, at present ' ch iefly inhabiting African regions. Remains of Crocidura Wagler are no t numerous in t he Wr;:ze breccia , similarly as in Plio- -Pleistocen e bed s of sou th e rn Eur op e . Among others they confirm the s u p po sit ion wi th rega r d to t he occu r ren ce - during the formation of the breccia - of re peated period s with hot climat e favou ring the weathering proces ses of limest on e a n d t he form atio n of red clay (t erra ro ssa) . Species of gen us Suncu s Ehr enb. likewise su pport this su pposition.

The ge ographi c distribut ion of both species : Suncus pannonicus (Kor- mo s) an d S. ze lceus n. sp. in t he Pliocene was wide e no ugh t o include all so uther n an d ce ntral Eur ope.

The extinct ge n us Pet en y ia Kormos, oc cu r r ing in equ al abu n da n ce as N eomys sori cul oid es n . sp . an d B eremendia jissid ens (Peten yi) must ha ve li ved in the Pli ocen e over con siderable area s of so uth-easter n Eur op e , but n ot pa ssing be y ond t he upper Rhine and w estern Alps .

The new genus Blarinoid es here re por te d is probably Low er Pliocene ;

t he Weze find being it s first recorded locality. Genus Blarina G r ay, related

124 ANDRZEJ SULIMSKI

t o it , now lives in Central and North America only , with out , as a rule..

passing to th e north of the Great Lakes line.

The mos t interesting of all the in sectivo re W ez e forms are Blarinoide s mariae n. s p. and Ber emetuiui jisside ns (P et enyi). In t he Plio-Pleistocen e of Nor th America these spe cies h ave t he ir eq uiva le nts in Blarina bre- v icau da (S ay) and Para cry pt ot is r e: r Hibbard. The stri king m orphologi cal rese m bla nce between these species suggests two altern a tives. One is t hat of con vergen ce , the ot he r of fau nal exc hange due to t he re pea te d con nec- tions of Asia w ith North Ameri ca , near ly t hroug hout t he Terti a r y.

G. E. A. Dobson (1882- 1890) w as the fir st to advance t he se s ugges t ions.

He did n ot atta ch great sig nifica nce to the last early Pleist ocene con nection of t hese contine nts and to t he fau nal migration of that period ; bu t rather pos tula ted t he possibili ty of an earlier fa u nal ex cha nge duri ng t he T er t iar y . J. S . Ogn ev 's conce ptio n (1930-3 4) ra n al on g a si m ilar lin e . However , he ra the r stro ng ly favou red the s u p posit ion that convergence w as ch ie fly res po nsi ble fo r the m or ph ological sim ilari ties in man y spe cies of both contine nts .

Resemblances of cran ial and mandibular st r u cture in Blarinoides mariae n. s p. and Blarina brev icauda (Sa y) together w it h differences of t he t ime of oc curren ce - t he former being r ec orded fr om the Upper Mi ocen e to Middle Pliocen e , the latter fr om the Upper Pli ocene up to - rece nt times - are di stinctly sugges t ive of t he ir common origin. Eurasia

must hav e been thei r birth-place while the mi gration of t h is primitive stock towa rd s Nor th Ameri ca and Eu rope ma y have occ urred as earl y as the Miocen e. This s u p pos it ion will , howev er , r em ain merel y h ypo- t heti ca l until palaeontolo gic al evide nce m ay be for thcoming from Asia ,.

whence t hese forms have not as yet been repor ted. Both these genera may possibly be ge netically rela ted with ge n us Het erosorex Gaillard '

fr om th e Mi ocen e of France (Pive t eau , 1958) .

Early Pl eist ocen e mi gration of t hese forms was not possible during the la st con ne ct ion of t he co ntine nts ow ing to con sider able cooli ng of climat e .

A sim ilar phen om en on is obser vable in t he ca se of the European Plio-Plei st ocene Beremendia jissiden s (P eten yi) and Para cryptotis rex ' Hibbard , on account of ver y close sim il ar it ies in structu re of the skull and mandible and on contem po ra ne ity of occur rence as w ell. The former is known fr om the Lower Pliocen e , pr ob ably up to t he Gunz-Mindel Int erglacial , the latter - fro m the Upper Pli ocen e to Middle Pl ei stocen e.

The dating of the only intermediate link connecting these two s pecies ,.

i . e. B . sine nsis (Zdansk y) fr om Choukoutien, is not su ff icie n tly reliable

t o determine an an al ogous mi gration r ou te fo r these species as that

f oll owe d by ge nera Blarina G ra y and Blarinoid es n. ge n.

PLIOCENE INSECTIVOR ES 125

Upon comparin g t he ecology a nd geogra phy of li ving and f ossil insectivor es with those described fr om t he W eze fauna , two distinct gr ou ps m ay be distinguished which correspond fairly well with t he twofold characte r of the Weze breccia. The fi rst gr ou p comprises spe cie s exist ing in arid ste p pe en viron m ent and under continental climatic conditions ; t he ot her gro u p tho se th riving in wa rm , moist pal aearctic zon es with diver- sif ie d land re lief, probably res embling cond it ions now prevailing in Eurasia.

This divisi on of the in sectivores fits quite w ell into the concepts of Stach (1952 , p . 155-156 ; 1953, p. 133-1 34) and Ml ynarski (1955 , p. 202-203) concerning sim ilar du ality in the palaeoec ological and pal aeogeographi cal cha racter of th e rept ile and carni vor e fa u na.

ON THE OR IGIN OF THE WF;2 E BRECC IA

The bon e m at er ial in the Wf;ze br eccia ha s accumulated in a hap- hazard fas h ion , with ou t traces of sorting . Bones belonging to one or mor e individuals are very ra re ly disc overed in the pl ac e wher e they w ere buri ed. As a rule th ey are fr agmen t ar y, for t u itous ly disp ersed in the form of hi ghl y characteristic b on e detritus (pl. I , fig . 1). The mo st common colo uration of b on es is dark or black. 'Com plet e, undamaged skulls of Soricida e are an ex ce p tio n. The rostral frag me n ts are those most abu n da nt.

The re d layer of the brecci a , with t he ex ce ption of bed s strong ly impregn at ed by calcite carbo na te , may be as cribe d to the r esidual cav e sli me. The li gh t coloured ("grey ") , strongly calcified breccia , was formed during a peri od when bon e r emain s w ere deposited on t he cave floo r by flo od waters . This may a ppear a sim plifie d solu tion of the problem, bu t just thi s cha rac ter of the b reccia is actually s ugges te d by the st ate of p re servation of bon es.

The cracks and pot holes due to t he erosion of Jur assic limest on e w ere

s ometi mes ove rfloode d by water. Besid es roc k d ebris t he w at er carried

also rem ai ns of animals t hat had di ed in the close neighbourhood , or t hat

may have fa llen in to t he cracks by accide n t, st ill ot hers inhabited the

crac ks and fissures . Som e bon es of ro dents and insectivor es - Erinaceu s

e xcepted , sin ce it is know n to have no foes - may have been brou ght in to

the cave by carnivores inhabiting it. T he hed gehog's pr esence in the cave

is eit he r accidental, or more probabl y due to its perman en t in habitation

t he re . The main part of ins ect ivore r emains, how ever ; must have been

trans ported by action of water. P ossibly too , birds of prey may h ave, in

126

ANDRZEJ SULIMSK I

part, acc umu lated the small m amm alia n rem a ins . T h is is sugges te d by the absence of bra in cases in soricides an d m a n y ro dents

1.

Taking ac coun t of these d a ta t he W f:ze fa u na m a y b e arranged in t o the following groups: a ) pe rmane n t or seasonal f auna of the ca ve, b) for ms b r ou ght into the carve b y pr eda tors , c) for ms acc identally falle n in to t he ca ve , a n d d) animal rem a ins, comple t e or f r ag men t a ry, ca r r ie d in t o the cave b y water duri ng tem pestuous rainfalls.

The p r esen ce in t he cave of teeth re mains belon ging to a you ng rhinoceros Dicerorh inus m egarhinus (Chri st ol) is ac ci de n t al. They may r epresent the rem nants of a feast held b y its carn ivore- in ha bitan ts .

ME

T HO DS

OF W

O RK

The chara cter of the b r eccia a n d of its ca lcareous ce ment m ade diffic ult t he m echanical prepar ation of t he m a te r ial. The on ly h ere ap plica ble tech n iq ue is th at of chemical pr epa ra ti on . 10-15 p er cent ace t ic acid solution was u sed b y t he write r. Often , however , w hen pr eparing minute a n d delica t e ele ments , it was on ly an 8 or even 5 pe r ce nt so lution.

For p r otection agains t injur y b y the aci d durin g mace ra ti on the bones w e r e coa ted wi th paraffin e , aft e r each dr y in g of t he bl ock. Bone s t h us se parated , washed a n d dri ed , were soaked in saturated so l utio n of shellac in alcohol.

Specim ens b elon ging to S oricidae and Talpida e were m e asur ed w it h a micr ome te r sca le unde r a binocula r m ic r oscop e , an d t he ob ta i ne d nu- m erical d ata r ou n ded to 0.1 mm. L a r ge r ele ments w er e m e asured , u sing fi ne cal ip ers , with a ccuracy up to 0.1 m m . Small spe cimens were m easured under a pprox im ately tenfold magnification ; in t he case of l a r ger spe ci mens t he magnific a tion w as fivefold.

In ad dit ion to the stan dar d leng th m easur e t he total a n d the cardin a l j aw leng th m e a sur ement w as constan tly us ed . The tota l jaw m easurement was taken fro m t he en d of t he incisor to the furt hermost post e ri or p oint of t he uppe r articu lar su rfa ce of the condyloid process , an d t he cardin al length on the lingual s ide of the j aw on ly, from t he anterior b order of t he m andibula r b on e to the same p oin t , as in the total measurement (fi g . 2A) . In order bett er to in dicate t he relatio n ef condyloid pr ocess to t he corono id p r ocess , m e asur ement ta b les are d one giv ing the h ei ght of the condyloid p r ocess fr om its base at t he contact wi th the angu lar process to t he upp ermost p oint of the articular surface in t he condylo id p r ocess (fig. 2B ).

I

Accordin g to a personal communication of Professor Ryziewicz, the breccia

has recently a lso yielded remains of birds probably belong ing to the Falcones or

Ga lliformes.

PLIOCENE INSECTIVORES

127

The mea su rement of the to tal jaw leng th is here less impor tant owing to the variable incisor len gth changing with individual age. Cranial measuremen ts were made accordin g to common standards. In view of s tr ong len g th , wi dth and heigh t variati ons in the skull of fossil soricides,

b

a

A

·1

B

Fig. 2. - Schematic illus t ration

of m

easurem e n ts

.A length of mandible, a total, b car dinal; B height of articula r processes, a coro n oid process , b condyloid process, c length of angula r process.

the num erical measurement data are only roughly approximate. The length measurements of t he maxillary to ot h-row, the length a nd width measu- reme n ts of th e palate, as w ell as pr oporti on s of teet h were done at the ve ntr al side of the skulls .

The illust rations were prepared wi th the help of the same opti cal instruments as those used for measurements, and with camera lucida for t he skulls and jaws of Erinaceus and Desmana. A constant 10 mm measu- r ement scale acc ompanies t he attached plates and photographs.

DE NTITIO N, TOOTH-COLOURATION, AG E CLASSES A N D MO R P H O L O G Y OF JAW ELEMENTS IN SORICIDAE

Som e subje ct iv ity in studies on fos sil soricides , particularly when carrying ou t mea surements or in th e determination of diagnostic charac ters , ser ious ly hinders re liable inferences. Differences in measurement data due to diffe r en t measur ement te chnique may be errone ous ly interpreted as an ex pression of geog raphic variation (H. Schaeffer , 1935).

In view of the mentioned difficulties which, as a r ule, ten d t o

individual in terpret ati on by the wri ters , the n eed seems obvi ous for the

128 ^{AND R ZEJ SULIMS KI}

introduction of detailed desc riptions of m easu remen t methods, of complete and universal numeri cal dat a on new spe cie s and, if possible , of thei r variation g r a phs .

The id en tification of t he penultimate un icuspid toot h as p 2 in Sorex L . a n d Blarinoid es n. ge n ., an d as pi in ge nera Ber emendia K ormos , Pe- tenyia Kormos and Neomus Kaup , is suggested by the s u p position that t he numerical re duction of unicu spids mus t have pr ogressed fr om ^{p :l} to Pl. This red uct ion is a lso asso cia ted with t he rostral shor tening of the sk ull , and wi th the sim ultaneous overgrow t h of th e 2nd and 3rd uppe r in cisors in gener a of the Cr ocidurinae gr ou p.

As compared to the individual age classification of S orici da e give n by H. T. Jack son (1928) , t hat acc epted by the pr esen t w ri ter has been sim pli fied . It resem bl es t he classi fic ation recog nized by A. Dehnel (1949 ) w ho distin guish es two ca r din al classes : t he yo u ng (adult) and t he o ld.

f. ptq .

" \'!t"+-- l im.

b

fc.c.sup.

o

l.i nto-t.

fc.c.inf. ~",--

fc. c. sup.

Fig. 3. -

a

Mor phology of cond yl oi d proces s (articu lar)

jc. c. sup. facies condyli sup er ior. tc c. in], facies con dyli Inferior, l. in tar t. lam i na Int er - artl culares .

b Morphol ogy of ar ti cu lar proces ses an d poste r io r pa r t of ma nd ibl e

inc. S. sup. Inci sura slgmoldea superio r, inc. S. info Incisura slgmo ldea in ferior , f. ptg. fossa pterygoldea, tor. mnd. for amen ma n dlbulae, lim. limula.

This sim pl ifica tion r esult s f ro m t he cha racter of so m e rem ains pr eser ved in t he br eccia w he re t hes e two classes are easily r ecogniza ble on the e x te n t of t he wear of t eeth.

One of the pr obl ems n ow under co nsi derat ion is the d iagnostic sign i- ficance assignable to teet h co lo urat ion in Soricidae. Dat a obta ine d b y research work of several authors (Dehnel , 1949, 1950 , 1952 ; Ku bik , 1951;

S ch aeff er , 1935, and ot hers) show tha t the occu rrence and t he in ten sit y

of coloura ti on vary in the li m its of one species . On the ot her ha nd , in

PLIOCEN E INSECTIVORES 129

so me fo ssi l form s of su bfam ily Crocidurin a e , t he col ouration occurs (Hibbard , 1950) but has a pparen tly no importan t t a x onom ic significance . This re m a ins an ope n question calling for more precise ob servation s . Dental col ourati on in species Suncus pannonicus (Kormos) and S . ze lceus n. sp. may b e cited in su p port of Hibbard's (l. c.) suggestion fo r the event u al unifi ca ti on of t he two su bfa m ilies of Soricinae an d Cr ocidurinae in to one taxonomi c group.

Ano the r open q ues tion is the influence of the h abitat on changes in te eth coloura ti on. S tudies b y M . G. All en (1938) ar e interesting but req u ire addi ti on al conf ir mation an d m ore d etailed investigation.

The fo llo w ing new terminolog y h as been introd u ce d by the w ri ter in systematic de scriptions o n speci fi c level :

a) in cisura sigmoi de a su perior and incisu ra sigm oid ea in fe rior - for the upper and l owe r interproce ssal sig moid notches of the jaw (fig.

3a );

b) lam ina interarticularis - for the interartic u la r list of th e condyloid p r ocess , u su all y called " b on e b ridge" (fig .

3b) -

a name that does no t adequa tel y inte r p ret t he stru ctu re of this jaw ele m en t.

SYSTEM AT IC DES CRIP TIO NS

Order Insec tivora Bowdich, 1821 F amily Erinace idae Bon apa r te , 1838

Subfamily Erinaceinae Gill, 1872 Genus Erina ceus Linnaeus , 175 8

Erinaceus samsonowiczi n. sp .

(pl

.

II, fi

g.

1 a-c

& 2)

Holoty pus:

specimen No.10511, left lowe r jaw

bea r in g P:

1-M:1 and frag

men tary

a

r ticu la

r processes.

Paratu p us: spe cimen No.1052,

left mandibl e w it h P:l

-M₃ a

n d

condyloid process.

Der i vatio no mi nis: samsonowiczi -

in hon ou r of Professor Jan Samsonowi cz,

di

scoverer of

Weze breccia.

Material. - 8 lower jaws represen ted b y fr agme nts of as ce n d ing r amus and horizo ntal ra m us with incomplete dentition ; f a irly numerous d eta ched , low er a n d upper t eeth and two ro stral fr ag ments of sk u lls: one with ri ght porti on of m a xilla, and incomplete dentition p r ob ably belon ging t o this speci es.

Descrip tion . - H ori zontal ram u s set at an a p prox ima tely r ig ht angle t o ascen ding ram us, t a pe ring an teri orly fr om Pt, . P osition of m ental f oramen betwe en P, and M

1

lower than in the living h edgehog . Height

1

All the here fig ured

and

d

esc ribed

specimens

a

re housed in the

Mu ze u m Ziem i

(Museum of the Ea rth)

in Warsaw, a

n d numbered provi s ion ally b

y

the pres ent auth

or.

Ac t a Pal a e o n tologl ca Pol on lc a - vol. IVj2 9

130 ^{ANDRZEJ SULIMSKI}

of jaw branch in P t, to M

₂

more or less uniform, but lower be yond M

₃•

Condyloid process strong, articular facet broad and nearly vertical to asce nding ramus. Shape of articular facet nearly regularly elliptical, lingually slig h tly expanded. Upper portion of coronoid process curved to the rear, blunt. Angular process broad and massiv e at t he base. An oblique crest placed labially to ascending ramus extends to the con dyloid process. Mandibular fossa s m all, oval. First lower incisor basally broad , elong ated , wi th a conspicuous, sle nder and pointed cutting edg e. Cingulum of incisor strong, but anteriorly almost ves tigia l. Alveoles of 2nd incisor notably sm aller t h an the preceding o ne. Alveole of canine large , nearly round , sligh tly labially pushed .

p;J

one- cus pe d, wi th con e ve r y blunt, cingulum con spicuous , r aised in posterior part to form a kind of cusp.

P, three-c us pe d, paraconid high , pointed , rising almost vertically ; pro- t oconid hi gher , fairl y pointed, directed posteriorly; metaconid more or less con spicuous . Anterior edge of paraconid sligh tly pointed. Labially cingulum le ss w ell developed. First molar with trigonid narrower than the talonid , bearing th ree w ell developed cusps - all notabl y blunt , the p ro toconid ex ce pte d - and a t alonid also bearing two rat her blun t cu sps. Structure of crown in t he next molar resem bles th at of M

I ,

all its cu sps being markedly blun t. In M

1

and M

2

cingula labially b etter marked . M:

₁

is the s mall est molar , consisting only of th e str ong ly re d uce d trigo n id with not more than one or two cusps and one r oot. In this molar th e paraconid is eit her very much r educed or absen t. Mandibular sy m physis oblique, b r oad. Tooth pigmentation da rk brown or nearly black . Skull (rostral fr ag m ent with right-side den tition) , com p rising al so ri ght -por ti on of pala te , nasal bon es, so me premaxillar y bones , fro n tal bones a n d part of t he zygomatic ar ch. Premaxilla not joint with fron tal bones . Lacrimal for am en sim ila rly pl ac ed as in t he li ving hed gehog . Nasal bones narrow, sle n de r, fron tal bon es relativel y broad , nasal forame n wid e , wi th probably elongat ed contou r. M axillar for amen pus hed so mewhat anteriorly abo ve the an ter ior canine root .

Dim ensions - see Table 1.

Remarks. - The

W~ze

spe ci m ens are rat her smaller than a ny spe cie s of Erinaceus L. thus fa r describe d from t he pre-Glacial of E urop e , with the exce ption of Erinaceus lech ei Kormos. On t he ir morphology and di- men si on s it has been ascertaine d t hat t he here cons idered re mains r esem ble t he Chinese Plio-Pleist ocene for m fro m Ch ou kou ti en E. algae Young a nd the li ving E. europaeus amuriensis Sc hren k (C. C. Young, 1934).

Specimens of Erinaceus sp. described by C. W. P ei (1940) an d Teilhard

de Cha rd in (1940) belong to ano th er grou p of hedgeh ogs (dea lbatus ) an d

markedly differ from the

W~ze

spe ci me ns . Our s pecime ns are smaller

than th e Mioc en e Palaeoerina ceus d . rec tus Ma tthew & Gr an ger (B . Boh-

PLIOCENE INSECTIVORES

T

a

ble I

Erinaceus sa IllSOIIOII ';C ;:; n.

sp.

- dimension s of mandibles

(in mm )

Man dible s --->

Specimen Nos. - )

131

T

ot a l

length of mandible

- -

^{- ca.38.4 - - -}

Ca

rd ina l length of mandi ble

33.8

36.1 - ca.35.5 -

- ^-

Length of:

II-M3 -

-

^{- 24.1a* - -}

^-

C-P4 18.0a

-

^{- 18.8a - 17.7a}

^-

12-M3 18.8a - -

20 .0

a - 19.8a -

P3-M3 ^16.0II 17.3 16.3 16.4 - 16.1a -

P4-M3 _14.1a 14.7 14.0 14.2 14.

3

14.4a 14.6

MI-M.1 _11.8a 12.1 11.5 11.5 11.8 11.8

a

12.2

M1-M2 10.5 9.8 10.0

9.8

10. 1 10.1 JO.2

M2-M3 _6.5

a 6.6

6.6 6.7 6.7 6.6

a

^6.7

12-P4 _7.4a - - 8.6a - 7.5

a

-

Lengt

h

of P

3 2.2 2.2 2.4 2.2 -

- ^-

Breadt h of P3

2.0

2.0 2.0

2.0

- -

-

Len

gth

of P

4

2.9

2.8

2.9

3.0

2.8 2.8 3.1

Breadth of P4 2.2

2.4 2.

1 2.3 2.2 2.3 2.3

Length of MI 5.4

5.5

5.0 5.4

a

5.1 5.4

a

5.5

Breadth of M

l

at tr

igon id

3.3

3.1

3.2 -

3.2

- 3.2

Breadth of MI at

talon id 3.5 3.5

3.5

-

3.4 - 3.3

Length of M2 4.9 5.0 4.7 4.9 5.2 5.0 5.0

Breadth of M2

a

t

trigo

nid 3.3

3.0

3.2

3.0

3.3 3.0 3.1

Breadth

of M2 at talonid 3.3 3.2 3.3

3.0 3.3 3.0 3.3

Len gth of M3 _2.3

a

2.0 1.9 2.0 2.2 1.8

a

2.0

Breadth of

M3

_1.7a 1.8 1.6 1.8

1.8

1.5a 1.5

He

ight

of

mand ible beh ind :

M

l

7.2

7.2 6.9 7.1 6.9 7.2 7.0

Mz 6.6

7.1

6.5 6.8

6.7

6.9 -

M) 6.2 6.7 6.0 6.3 6.2 6.2

6.3

Thickness of man dible below :

M1

3.2 3.2 3.2

3.0 3.0 3.2 3.2

M3 3.1

3.4 3. 1 3.2 3.3 3.4

3.3

Bread th of

cond

ylo id process 5.6

5.0 -

- - -

-

Height of ascending ramus 17.6 ?* - -

-

- -

-

Length ofangular

proce ss

9.6 -

- - ^{- -} -

D

istan ce

between condy lo id and a

ngular

processes 11.4 - -

- -

-

-

"*^{iI -} measuredalong thealveoles, P - measurement uncerta in.

132

^ANDRZEJ SU L I MS'K I

lin , 1942; J . Vi r et , 1938) a n d their denta l p ropor ti on s ex h ib it marked dissimilarities. Neithe r is a cl ose r cor re latio n possible b e twee n the

W~ze

species an d E . pmegla cialis Brunne r , 1934 , ow ing to i nade q uate k n owled ge of the latte r.

S om e man di b u lar fr a gme n ts of this newly recorded h ed gehog from Wc:;ze resemble E. sansani ensis Depe re t, 1887 d esc ribed from th e Mi ocene of France . Th e position of t h e ment al f ora me n is sim il a r, the lower mandibu la r edge is arcuately b ent dow n in both for ms, as ce n d ing ram u s likewise placed at a ri ght angle to h or izon t al ramus. Similar condition s are noted in conspecific specime ns described b y C . Gaillard (1899) f rom Grive-Saint-Alban . The W c:;ze specimens, howe ver , di ff e r n ot in size only , b u t als o in arr angemen t of teet h, larger dimensi on s of 1

2 ,

position of cusps on P 'o , l ow er depression of menta l foramen an d m ore r obust articular process es of asce n din g r amus .

The separation of th e Weze spe cimens in to a n ew s pe cies of hed gehogs is sugg es ted on the fo ll owing features : characteristic low p osition of mental foramen , s m a lle r leng t h of th e t ooth-row 1

₁

- M ::, s tro ng st r u c tu re of th e w hole lower jaw, small er dimensions of cardinal length , differences of length in tooth-rows P! l-M:

l

and P ,,-M:l, sh a pe of P: l an d PIt which, as a rule , are with a slig h tly pos t e r iorly curved protoconid , fin ally f requen t stron g reducti on of trigonid in M:

_l.

Erinaceus sp .

(pl. IV, fig. 11)

Material. - Two left mandibular fragmen ts l acking d entition and with strongl y in jured a r tic u la r processes . On e fragment wi th the 1,-M :J alveoles , t he ot her with M

2-M

:: al veoles .

Dimensions of two mandibles (in mm):

Mandibles - >

Specimen Nos.- >

Length of tooth-row:

C-M,

MI-M .I M2-M,

Height of mand ib le below:

MI

M

₂ M.I

Thickness of mandible below: Ml

M2

M ,

1060

16.4 11.8 6.6

5.5 5.7 5.3

2.9 3.0 3.3

2 1061

6.2

5.2 5.0

2.9 3.0

PLIOCE NE INSE CTIVORES

133

Rema rks. - In dim en si ons bot h frag me n ts approach Erinaceus lechei K ormos, 1934 , described fr om Hungar y . But the unsatisf ac t or y state of preservation an d mea gre materi als do not allow its s pe cifi c id entification.

F amily 'I'alpidae Gr ay , 1825 S ub family Talpinae Murray , 1866

Genus Talpa Linnaeu s , 1758 Tal pa m in or F reu d enber g , 1914

(pl.

I

V, fig.

10)

1914. Tal pa eUTop aea val' . minD!' Fre udenberg;

W .

Freuden be rg,

Die Sa ugetiere ...,

p. 209, pl.

19,

fig.

7,

32,

35-37.

Fur ther synonymy

in:

1956. Talpa minor Freu

d

enberg; K.

K

owals k i,

Insectivores...,

p. 341-3 42, pI.

1, fig. 1,

and

1955. Talpa minor Freudenb erg; M.

Kretzoi, Die Alt p le is toza ne

...,

p. 162,

,

1-92, 197-200,

232.

1953. Talpa minoTFreudenberg; K.

Kowal s k i,

An early Ple istocene..., p. 8, 9.

Mat erial. - About 15 mandibular fragments, in various state of preservation and with incomplete dentition. S everal specime ns with M cM

³

or C-P,., a few d etached molars , canines an d incisors , also lon g bones of limbs . A doz en or so of humeral bon es. Skulls unknown .

Dimensions of 5 mandibles (in mm) :

Mand ibles

- )

2

3 4

5

Specim en Nos

.~

51 8 565 567 569

300

Length of:

1[-M 3

10.8a*

11.2

a - -

-

M

[-M3

5.4 5.2 5.3 5.6 5.0

M[ 2.0 1.9 1.9 2.0 1.9

Breadth of M

I

1.2 1.1 1.2 1.3 1.2

Len gth of M3 J.5 1.4 J.5 1.6 1.4

Breadth of M3 0.8 0.7 0.7 0.8 0.6

Height of mandible below M2

1.7 1.7 1.7

1.7 1.6

T

hick ness

of mandibl e

below M2 1.0 1.1 1.0 1.1 1.0

Height of cor on oid process 5.2 1* 5.1

? -

- -

• <1 - measured along thealveoles, 1"- measurement uncertain.

Hum er us (in

m

m)

10.6 to 11.5 3.1 to 3.5 Lengt

h

.

Small

est brea

dth

134 ^{ANDRZEJ SU LIMSKI}

Remar k s. - Th e appe arance of horizon tal ramus on t he whole agr ees with those described by W. Freud enberg (1914) and K. Kowalski (1956 , 1958). Ascending ra mus set at rig ht or slig htly obtuse ang le to horiz on tal ramus. Po sterior m en tal foramen a ppro x imately between t he r oots of M

1,

but more often below t he posterior root of that tooth. Teet h an d humer al bones struct u rally approachi ng T . europaea L ., but smaller a nd m or e delicate.

Talpa fossilis Pet en yi , 1864

(pl.

I

V, fig

.

9a-b)

1864. Talpa vu lgar is tassili s

Petenyi;

J.

S. Petenyi, Hatrag yott Munkai..., p.53-58,

pI.

1.

F

urther sy

no ny my

in:

195B.

Talpa tassilis

Peteny i; K.

K

owa ls ki, An early Plei st oce ne..., p. 9-10, fig. 1.

Material. - About 10 incomplete lower jaws, also fragment ary maxillae in various state of preserva tion ; numerous detached molars , several canines and incisors; fragmentar y humeral bon es.

Dimensions of 3 mandibles (in mm) :

Mandibles

. ~

Specim

en

Nos.

.- .

1

95 1

2 952

3 953

Length of

:

I I

C-M

.I 12.6a* ^I - -

M.-M

} 6.8

I

_{6.3 6.8}

M.

2.5

I

_{2.3 2.4}

Breadth of M

.

1.5 I

I 1.4 1.5

Length of M,

2.0 I _{1.9 2.1}

I

Breadth of

M_J 1.0

I

0.9 1.0

Height of mandible below

M

z

2.2

I

^{2.1 2.0}

Thickness of mandible below Mz

1.3 _I 1.2 J.J

• a- measured alo ngthe alveo les.

H

ume r us

Maximum len gth

Minim u m

Prox imal b

read th

Distal

(in

m

m ):

13.6 to 14.0 3.9 to 4.1 11.4 to

12.2

B.Bto

9.2

Rem ark s. - Mo rp holo gically thi s mole closel y resembles liv ing T alpa

eu ropae a L. It is on ly on s tronge r str u ctur e of teeth a nd of th e mandibular

hori zon tal ra m us , and some di ff erenc es i n in di vi dual mola rs that these

f oss il rema ins can be di stingu ish ed fro m recent m oles . In t he Weze

PLIOCENE INSECTIVORES 135

s pe cimens t he pr otoconids a n d hypoconids are som ew hat ext en ded an - te ri orl y. Cingulum of molars is more conspicuous and b etter d eveloped anteriorl y. In r e cent moles t he molar t a lo n ids are narrower a n d t he row with molars di stinctly shor ter. Post erior mental fo ramen is , as a r ule, b elow the a n terior root of MI' Along M i-M

3

horizontal ra m us o f nearl y uniform he ight . The same a pplies he r e to thickness of mandible . The canine di stinctly hig her t han the premolars, one-topped , n a r row , wi th sp icule directed so mew h at p osteriorly. Premolars (Pi-P I,) are w it h di stinct ci ngul u m , poste ri o rl y stretched out so as to f or m sm a ll cusps. The coronoid proces s relati vel y low , sim ilarly as in T. pra eglac ialis Kormos (Ko rmos , 1937 a, b ; Heller, 1936 a, 1954). S truc tu r e and dimensions of humeral b on es u sually typical of t h is ge n us.

The Wr;ze specime ns are n otably larger than T . minor Freudenber g , but only slight ly so t han T. episcopalis Kormos (Kretzoi , 1956). In size they nearly agree with T. stro meri Brunner , 1950. In a str ong an ter ior curvature of the tooth tips the latter fo rm differs distinctly from T . fassilis P erenyi . The structure and size of humeral bones are sim ilar in these two species.

Talpa eurapaea fossil is L. described by A. Pasa (1950) is only sli gh tly smalle r than the her e st u die d species and is probably conspecific. Talpa sp. recorded b y C . W . Pei (1931) from t h e early Pleistocene of Choukoutien exhibits man y features in common with T. fassilis Petenyi, Its humerus , however, is sm a ller, while the vast geographical distance of their occur- r e n ce does not suggest a closer relationship.

Talpa sp.

Material. - Some incomplete lower jaws and ar ticular processes, on e incomplete humerus , a few detached molars .

Remarks. - The here represented specimens do n o t permit more

exact specific classification. Nevertheless in size of horizontal ramus they

ap pr oa ch nearer to the living Talpa europaea L. This is mo reove r su gg este d

b y the shape and size of th e humeral bone , morphology of molars an d

character of the prese r ved fragment s of articu lar processes . These m ol e

remains have been recov e r ed from calcareous rock debris i n the upper

portion of th e kars t d oline , probably referable to t he fi nal period of

for m a t ion of t he Wr;ze breccia.

136

^{ANDRZEJ SUL IMSK I}

Su~family

Desmaninae Thomas, 1912 Genus D esmana Giildenst aed t , 1777

D esm an a nehringi K or mos, 1913

(pI. II, fi

g .

3

a-d)

1913. Desmana (?) Nehri n g i

K

orm os ;

T. K

ormos,

Trois n

ouvelle s es

p

e

ces

...,

p

.

138, pl. 6,

f

ig. 1 a-f .

Further synonymy in

:

1956. Desmana neh r in gi

K

ormos;

K.

Kowals k i, Insecti vore s ...,

p

. 342- 344, pl. 1,

fig. 2

a-b,

3

a-b.

Material. - Several low er jaws, fairly well preserved ; one r epresents the right mandibl e with I:.! , P

2

-M

₂

and damaged articular pr ocesses ; also a nearly complete sk ull lack ing the zygomatic arches , with the skull roof partly damaged , with dentition more or less complete on both sides of the jaws .

D escr i pt i on . - Mandib le . The ma ndib u lar structure does not essen- ti ally diff er fro m the type described by Kormos (1913). T he differences co nsist in certain morphologic al features w hic h may be assigned to in dividual va riation. The more important differentiati ng features of the Wf;ze speci mens are: position of anterior m en tal for amen (between roots of P, and P :.! or be low P :.!), also that of post eri or men t al fora men (bel ow t he protoconid of M ) or bel ow t he anterior root of that tooth), st ruct u re of P

2

with we ll devel op ed cin gulum lack ing the par ac onid , st ruct ur e of P 3 wi th the lab ial cing ulu m m or e dis tinct and a kind of posterior cus p, str ucture of P, with d istinct paraconid an d a con spicuou s labial cingulu m strong ly cu rved over t he br anchin g r oots . In t he W eze specimens the molars are wit h high er prot oconids , w hile the ta lonids are broader t h an the t rigon id s w hic h h av e a strong labial cingulum. Last mo lar is t he lowest wi th unreduced talo nid . The two jaw b r an ch es of the Wf;ze specimens meet at right ang le , as in Hun ga ri a n specimens . The con dition is sim ilar in P :\ alveoles where t he posterior root of that toot h has been pushed somew hat lingually . The articular processes h ere are probably sha ped as in ty pical forms fro m Hungar y .

Sk ull satisfactorily pr eser ved , stro ng ly elongated, with occipital par t

broad and dom ed . F oramen m agnum ov al , hori zon t ally expanded. The

rem n ants of the zygomatic arch suggest t hat t he arches were delica t e

and sle nder. In t he upper or bital par t the sku ll strongly co nstricte d, nasal

bon es long an d narrow, shar ply terminating w it hi n t he suture wi th

fronta l bones. U p per incisors 1

^{1- 1}

lar ge , br oad at the b ase, in se ction

subtriangular . Their an terior and posterior ed ge s sharp. These teeth are

more or less de pressed posteriorly, the ir talonids low . The outer t oot h

surface covered by delicat e enamel mosaic. Na sal foramen horizontally

PLIOCENE INSECTIVORES

T a b l e 2

Desmona nehringi Kormo s - dimensions of sku lls (in mm)

137

Skulls ^~ 1 2 3

Specimen Nos. ----+ 1201 1202 1203

Total length ofsku ll 45.7 -

-

Maxima l breadth of skull 20.1 -

-

Minimal breadth of sk ull 8.5 8.0

-

Anterior breadth of palate 5.5 5.2 -

Posterior ,.

"

^{" 13.2 13.5}

-

Nasal height ofsk ull in pi 5.0 4.8

-

Rostral height of skull in M2 8.0 7.6

-

Maxima l breadt h of alveole II 2.9 2.5 2.8

Minimal

"

^{" "}

"

^{1.1 1.0 1.2}

I. Length of C 1.2 1.1a* 1.0

2. Breadt h of C 1.3 1.3a _1.I

1 :2 0.92 ca.0.84 0.99

I. Len gt h of pi 2.0 1.8a 1.9

2. Breadth of pi 1.3 1.3a 1.2

1:2 1.53 ca. 1.38 1.58

1. Len gth of pz 2.1 2.1a 2.2

2. Breadt h ofp2 1.5 1.6 a 1.6

1:2 1.40 ca.1.31 1.38

1. Length of pJ 1.3 1.3 1.4

2. Breadth of p3 1.6 1.5 1.6

1:2 0.81 0.90 0.87

1. Len gt h of p4 2.5 2.2 2.5

2. Breadth of p4 2.3 2.3 2.5

1:2 ^\ 1.08 0.95 1.00

Length of MI 2.8 2.7

-

Breadth of M! 3.3 3.3

-

Length of MJ 1.8 2.0

-

Breadth of MJ 2.0 2.1

-

Lengt h of:

Jl-M3 22.0 22.5a -

C·P4 9. 1 8.6a 9.0

MI_M3 7.2 7.0 -

Breadth ofbridge over infraorbital foramen 0.8 0.9 -

Bread th ofnasal part fo r 11• 1 6.2 6.4

-

• a - measuredalong thealveoles.

138

^ANDRZEJ SU LI MS K I

T a b l e 3

Desmana nehringi Kormo s - dimen sions of mandibles (in rnrn)

Mandibles --> I 2 J

Spec imen Nos. --> 1204 1205 ' 1206

,

Total length of mandible ca.26.3 - -

Length fromthe anterior border ofmandible bon es

to the posteri or border of M) ca. 16.3

- -

Length of:

I1- M) 20.0a" ^-

-

C-M) 14.3a - -

M,-M) 7.5a 7.5 7.3

Length of

M,

2.8 2.6 2.8

Breadth of

M

I at talon id 2.5 2.2 2.0

Breadth of M, at trigon id 1.8 1.8 1.8

Lengt h of

M)

- 2.2 2.0

Breadth of

M.1

at talonid ^- 1.6 1.5

Bread th of

M.1

at trigon id

-

1.6 1.6

He ight at protoconid of:

P 4

1.8 1.8

-

M,

_{2.1 2.1 1.6} ?*

M2 2.5 2.5 1.8?

M3 -

1.6

1.2

?

He igh t of ma nd ible below 1'.12 3.5 3.6 3.3

Thickness of man dibl e below M2 2.1 2.2 2.0

Bread th ofcondyloid process

-

2.6 -

He igh t of ascending ramus - ca. 11.3

-

Distanc e bet ween the ante rior and posteriorment al

foramens 6.5 6.2

-

• a- measured alo ng theaiveo les. ?- measurement uncer tain.

expa nd ed . Anterior edge of nasal bon es - over nasal foramen - slightly

conve x anteriorly. Bridge over infraorbital fo ramen narrow, fa intly

oblique ; its l ow er end is betw ee n M

₁

and M

₂.

Second and third in cisors

somewhat sma ller than t he talon of II , one-:to pped , one-rooted, w it h

di stinct labial cingu lum. C hi gher than in cisors, two-roo te d, one-topped ,

and a more or les s con spicuous postero-lingual cusp. p i lower than the

canine, wit h one root and t he cingulum labially more distinct. T he next

premolars of similar str u ct ure , with two roots , with la bially disti nct

cingulum and posterior small heels. Last premolar high, with a strong

protocone. In st r uct ure this toot h does not essen tially differ from p r. , as

has been observed in a type form from Hungary and in specimens fro m

P odl esice . First molar with su btriangular ou tl ine at top an d w it h elo ngated

talon. Metacone and m etastyle of M' joint by a sh ar p edge. P oste rior

PLIOCENE INSECTIVORES

139

mesos ty le a lso joint to the m etac on e along a short edge which cu rves out anteriorly. P ar as tyl e wi th a conspicuou s knob. Anteri or molar cingulum strong. Next m olars of sim ilar structure. Last m olar strong ly r educed, shorter and narrower.

Dim ensions of 3 skulls and 3 man di bles - see Tables 2 & 3.

Re mark s. - Diff er en ces in proportions b etw een C and p 3 of the Wf:ze s pecimens an d t hose descr ibed by A . Schr euder (1940) m ay , po ssibly , result fr om different measurement metho ds. These di ffer ences suggest t hat the skull be longed to another species. Neverthe less the study of other teeth per mits to ascertain t hat these ch ar acters are subject t o variation and may not be rega rd ed as diagn ostic featur es . Indices no ted in t hese f or ms have not been r eported in an y ot her s pecies of Gal emys Kaup and Desmana Gi.ild. The characteristi c arrangement of teeth and their structure , b read th of bridge over infraorbital foramen and the mandibular mor- phology - all indicate gen us Desmana and spe cies D. nehringi Kormos.

On the other hand , a relatively wide bridge over infraorbital foramen , shar p ante rior ed ge of protocone in p 4,

~nd

the weakly dev el oped cingula of low e r molars ex clu de gen us Galemys Kaup.

Genus Galemys Kaup , 1829 Galemys (?) sp.

Material. - Fragments of sk ulls (ros tral parts) wi th man y gaps in dentiti on , also f ra gments of lower jaws eit her with or without teeth. Few detached t eeth of the upper and lower jaw.

Dimensions of 3 sk u lls and 2 mandibles - see Table 4.

Remarks. - Th e fragmentary condition of the recovered remains doe s not perm it thei r su re specifi c identification , neithe r is their generic iden- t ifi ca t ion q uite doubtle ss. On measurements it may be ascertained that these forms are notably smaller than Desmana Gtild. , in size coming nearer Galemys Kaup , possibly one of the Hungarian for ms : G. semseyi Kormos or Mygalina hungarica (Kormos) (Schreuder, 1940) . Differences in sh ape of maxillar teeth , and the small er breadth of bridge ov er infra- orbi tal for ame n , do not permit their definite assignment t o one of these spe cies .

The presence in Wf:ze of gen us Gal emys Kaup m ay indicate a fa rther

no r thern range of t h e Pliocen e occu rrence of t his ge n us. It has been

actuall y re cor de d fr om Plio-Pleistocene formati ons nearly all over

Europe . In central and eastern Asi a it is no t known either as a living or

.ex tlnct f orm.

140 ^ANDRZEJ SU LThl S K I

Ta b le 4

Galemys (?) sp. - dimension s of skulls and mandibl es (in mm)

Skulls -->

I

^{I 2 3}

Specimen Nos. - -- -- >

I

^{130 1 1302 1303}

Anterior breadth of pal ate 4.4 5.0

-

Posterior breadth of pal at e

.

^{10. 6 11.6 -}

Length of too th-ro w with:

MI-MJ 6.3 6.5 6.3

MI-J'v12 4.5 5.0 4.8

M2_MJ 4.0 3.8 3.8

Thickn ess of bridgeoverinfraorbita lfor a men

I

0.6 0.7 0.6

Mandi bles --> I 2 3

Specim en Nos. --> 1304 1305 1306

Length of tooth -row with:

I

M,-M3

-

6.8 7.0:*

M2-MJ 4.5 ? 4.5 4.7

Length of M,

-

2.5 ^-

Breadth of M Iat talonid - l.l -

Breadth of M1 at trigonid ^- 1.0 -

Length of MJ 2.0 2.1 2.1

Brea dth of MJ at talonid 1.4 1.0 1.5

Brea dt h of M3at trigonid 1.4 0.8 1.4

Height of mandible below M2

I

3.0 3.1 3.0

Thickness of ma nd ible below M

z

^{1.6 1.6 1.7}

• ? - measureme nt uncertain.

Family Soricidae Gray, 1821 Subfamily Soricinae Murray, 1866

Gen us Sore x Lin n aeus , 1758 Sore x runtonensis Hinton , 1911

(pl. IV, fig. 5a- b)

1911. Sorex runtonensis Hinton; M. A. C. Hint on, The British FossiL, p. 532, pl. 25, fig. 8, 9;

textfig.

8-a.

1930. Sorex runtonensis Hinton; F. Hell er, Eine Forest-Bed-Fauna..., p. 259, pI. 15, fig. 5a-b; tex tfi g. 6.

1933. Sorex runtonensis Hinton; G. Brunne r, Eine pr a egl azi ale Fauna..., p. 309, pl. 6, fig. 1-3; textfi g, 1, 2.

1933. Sorex runto ne nsis Hinton ; F. Hell er, Ein Nachtrag ..., p. 62.

PLIO CEN E INSECTIVORES 141

1937. Sorex runtonensis Hin ton; T. Kor m os, Revision del' Kleins a uget lcre..., p. 31-33, fig. 3.

1949. Sorex

r untone nsis

Hin ton; 1\1.Fr ia nt, Les

Musara ign es...,

p. 239, fig. 3.

1958. So rex cf. Tll1ltonensis Hinton: K. Kowals k i, An early Ple is t oc ene..., p. 11-12, fig. 2.

Material. - About 25 mandibular fragmen ts, va rio us ly preser ved ; n u merous detached in cisors a nd molars , long limb bon es, also fragmentary upper ja w s, pr ob a bly conspecif'ic.

Description. - Morphologic ally th e preserved mandibular remains come ne arest to forms descr ibed fro m W est - Runt on (Hinton , 1911). T he lower incisor has th ree lobes , se pa rat ed by two dis ti nct notch es . The two posterior lob es are strong er t han t he anter ior one. The prem olar is two- to pped, with the h ypocon e not s o w ell dev el oped as in t he r emaining so ricides . M:: wi th five cus ps. The coronoid pr ocess sle nder, basally broad.

Ascending ram us meets horizontal ramus at right or sligh tly obtuse angl e.

Arti cu la r fa cet of the corono id process sm ooth , without narrowin gs.

Masseter crest as in li vin g Sorex arane us L . In the condyloid pr ocess the art icular facets resem ble those in S . aran eus L., with the lin gu al end of the low er f ac et extending farthe r down. Upper and lower sig mo id notches em argin ate d, conspic uo us. P ter ygoid fossa high , deep, triangular. Me ntal foramen in fron t of t he anterio r root of M , or bel ow P I,. Tips of t eeth discol oured or distinc t ly pi gm en t ed.

In on e cranial frag ment lacrim al foramen placed between the roo ts of MI. F ive unicuspid te eth . Other features as in the living Common Shrew.

Dimen sions of 5 m andibles (in m m) :

Mand ibles ~ 1 2 3 4 5

Specimen as. ~ 1 55 15 136 148b

Total lenght of ma nd ible 11.2 12.0 12.4

Cardinal

length of mand ible

9.0 8.6 9. 1 8.8 8.8

Length of

I-

M3 7.3 7.1 7.1 7.0 6.9

Length of MI

-M3

3.8 3.6 3.7 3.3 3.8

Height of mand ible below M2 1.4 1.3 1.3 1.3 1.3

T

hick ness of mandible

below M2 0.9 0.8 0.8 0.8 0.8

Height of coronoid process 4.0 4.0 3.9 3.8 4.1

Ind ivid

ual aile of specimen y 0 1I n g

Remarks. - Among rece nt Eu ropean sh re ws S orex caecuti en s Lax-

mann occupies an in te rmediat e positi on in wha t size is conc erned between

S . aran eus L. an d S. m in utus L. Hence, S . runtonensis Hin t on oug ht to be

nearest to that spe cies . T he he ight of the coronoid process is an al og ou s

with tha t in S . caecutien s Laxman n , but the ot her cha racters fi t in t o

the variation of S . aran eus L.

142 ^{ANDRZ EJ} SUL I MS K I

In for ms fro m Hundsh eim kn own as S. py gma eus P allas (Freud en berg, 1914) and definitel y id en tified by Kormos (1937b) as S . runtone nsis, t h e total len gth of the low er jaw is abo ut 15 mm. The length of t he

W~ze

spec ime ns su pposed to be 12.4 mm , bu t prob ably ranges fr om 11 to 13 m m.

S. praea raneus described b y K ormos (1934) ma y po ssibly be a sy- non ym of S. runtone nsis Hin t on .

So rex araneus Lin naeus, 1758

(pl. IV, fig. 8a,b)

Mate rial. - A score or so of fairly well pr eser ved mand ibles , a mo n g t he m one n early comple te wi th s lig h tly damaged artic ular processes ; detached upper and lower teeth. Cr anial fr agmen ts re presented by rostr al pa rts wi th unicuspid t eeth typical of t his species .

Dim ensions of 6 mandibles (in mm):

6 481 5

633 4

2 3

305 2

85 J

500

jvla ndi bles

~ " ----1-- - - : _

Specimen Nos. --~

Tot al length of mandible 11.8 11.8 11.1 11.4 11.4 11.7

Cardinal length of mand ibl e 9.3 9.3 9.2 9.2 9.4 9.5

Length of I-M J 8.1 8.2 8.5 8.3 8.5

-

Len gth of M1-MJ 4.0 4.1 4.0 4.0 3.9 4.1

I

Height of mand ibl ebelo w M₂ 1.5 1.6 1.5 1.4 1.5 1.5

Thickness of man dible bel ow

M2 0.8 0.9 0.8 0.9 0.9 0.8

Height ofcoronoi dprocess

-

4.6 ^- -

-

-

Individ ualage of specimen young old y 0 u n g

Rem ark s. - Cran iom etr ic dimensions and mandibular morpho logy almost identical as in t he Common Shr ew . Sli ght diff ere nc es, con sisti ng in somewhat g reater leng th of the tooth-row M, -M;\, have no sig n ifi cance here a nd fit into t he rather strong ind ividual variation dis played by th is species . T his also ap plies to t he posi tion of m en t al fora men , sh ape of articula r face ts in the condyl oid process , and t he mor p ho logy of t he incisor and ot her mandibular tee th .

The presence i n the brec cia of this s pe cies con firms th e su p pos ition t hat the materia l was mixed after excavation. This is a form recorde d from t he earliest P le istocene and , so far, never reported from the Plioce ne .

Sorex cf. minutus Lin na eus , 1766

(pI. IV, fig. 3a,b)

Materi al. - S ever al fragments of mand ib les with incomplete denti-

tion and partly damaged articular processes ; numerous long bones of

PLIOCENE INSECTIVORES 143

limbs, detache d u pper and lower teeth, also a dozen or so of frag mentary maxillae, probabl y of th e same species; com plete skull not known .

Dimensions of 6 mandibles (in mm):

6 327 5

331 4

335 3

179 2

325 I

207

Mandibles )

1 --- - - - - -- - ---,-,.--

Specimen Nos.- -_ - - - -

Total length of mandible 8.3 8.8 8.9 8.7 ^- -

Card ina l lengthof mandibl e 6.6 7.2 7.4 7.1 ^- -

Length of I-M3 5.7 6.1 6.2

-

-

-

Length of MI-J\h 3.0 3.2 2.8

.

-

-

^-

Height of mandiblebelow Mz 1.0 0.8 0.8 1.0 0.8 0.6

Thickness of man dible bel ow

!'viz 0.6 0.5 0.6 0.5 0.6 - 0.6

Heightof coron oid process 3.0 2.9 3.0 3. 1 3.3

-

Individu al age of specimen y 0 u n g old

Remarks. - Very small mandibles, with teet h and articular pr ocesses , so characteristic of this species , are n ot abundant in the breccia. Fr om Sor ex minutissima Heim de Balsac (1940) they differ in arrangeme nt of lower molars . The m ola rs of the Wf:ze specim e ns are w it h relatively low cr ow ns and with cusps no t so di st inctly anteriorly extended as in S. mi- nutissim a. From t he living shrew the y differ in somew hat sm all er size .

Sore x sp .

(p l. IV, fig . 6a-c. 7a-c)

Material. - More th an ten mandibles wit h inc omplet e dentition ; n umerous detached lower mola rs , also lower inc isors .

Dimensions of 5 mandibles (in mm):

Ma nd ibles- -- - - Specimen Nos.

I 1100

2 110 1'

3 1102

4 1103

5

1104

Length of I-M3 7.8 8.2 7.7 6.9 7.5

Leng th of MI-M3 4.2 4.0 4. 1 3.8 4.2

Heigh t of mandible below M

z

^{1.6 1.7 1.6 1.6 1.6}

Thick ness ofmand ible belo w M

z

^{1.1 1.0 1.2 1.1 1.2}

Heigh t of co ronoid process 5.3 5.7 - 5.2 ^-

Individual age of specime n yo u n g old you n g

Re marks. - The inco m plete state of preservation of these re mains

bars its comparison wi th thus fa r described s pecies. In size they approach

So rex savini Hinton (Hinton 1911 ; Kormos, 1937b ) and S . dehneli K ow al-

ski , 1956. Th e morpholog y of teeth an d articular processes , also of the

144 ^{AND RZ EJ SULIMSKI}

hori zontal r amus , agrees fa irly well w ith analogous ch a r ac t er s in the tw o ab ove named species. The de termina t ion of t heir system atic position calls for addition al material s . S ome fragmen ts m a y belong per ha ps to S. m ar-

garit odon Ko rmos , 1935.

Ge nus Blarin oides n . ge n .

Geno ho lo t ypus: Blarinoi des mal'iae n. sp.

Deriva tio nominis: Blari noides - after its rese mb la nce to the Amer ica n ge n us Blar i na Gra y .

Diagn osis . - A large shrew with dental formula 3 1 3 3 _ 3"

1 1 1 3 -

~.

Asce ndin g ramus and h ori zon t al ramu s of mandi ble massive. Lowe r mandibular incisor lon g , massi v e , wi th two d istinct lobes on the cuttin g edge a n d a small accessory l ob e just beyond the t i p of t hat t ooth . The canine sm a ll , flat ten ed . P I, large , with in d istinct two c us ps, with a w ell d eveloped poste r iorly extended cin gulum. M:\ t hree t imes sm a ll er than M

_1,

with five cusps (ent oconid sm all, v isible). Corono id proces s 'b r oad , gen tly anteriorly fl exed. Masse t e r crest w ith spine . Condyl oid p ro cess massive , interarticular li st broad , sligh tly linguall y n otched. U ppe r ar ticu la r fac et of condyloid proces s inclined at a n ang le of abo ut 45

^0•

Ascending ramus meet s h orizontal ramus at an ob t use ang le .

P rofile of sk ull gen tly slo p ing, typical of ge nera S orex L. an d Blarin a Gra y. Nasal f oramen hi gh , of n early u ni f orm wid th throughout its h e igh t . S tr ong external flexions obser va ble next to t he roots of p it . Lacrimal for a m e n abov e th e post er ior ro ot of MI. In fr a orbit a l fora men ab ove p it . An te ri or p al ate f or a me ns between

_P-~.

p oste r ior palate fo r amens just in front of ante rior ro ots of Ml -l. 12 and r

^l

la rge (the third somew h at l arge r than the se con d) with postero-lingual cusps . C an d P! a bo ut h alf the siz e of in cis ors , simil a r in stru ctu re . P '' m inute , p ushed below P it, late r all y n ot visib le. Molars slig h tly posteriorl y e x ca v a te d . M3 three-c us ped with reduced ta lo n.

Blarinouies maruie n . sp .

(pI. II

, fig. 4a- b ; pl.

III,

fig.6a- c; text-fig. 4, 2a-f)

Hol otypu s: spe cime n No. 803,

r

ostr al part of skull w

ith

MI- I_ M:l- :l, an

d

an incomplete row of incisors a

n d pr e m ola r s.

Par atypu s: specimen No.

178, left mandible

with ar t icular processes and

I

- M;\.

Ma

n di b le

a

n d skull probab ly b

elo nging

to

the

sam

e in

di

vi

dual.

Der i ua tio nominis: mari ae- from Maria, the name of

th

e aut ho r' s

m

ot he r.

Mate rial. - Five complete mandi bl es an d 10 rostr al p a rts of skull

with incomplete dentition. Al so a bo ut 40 m andibular and cran ial fragment s,