ANNALES HISTORICO-NATURALES MUSEI NATIONALIS HUNOARICT Tom us 05. Budapest 1973.

New Species of Episoriculus from the Middle Pliocene of Osztramos (North Hungary)

B y D. JÁNOSSY, Budapest

Abstract — Two new species, Episoriculus borsodensis sp. n. and E. tornensis sp. n . are described from Osztramos, N . Hungary. Both originate from the younger and older Middle Pliocene. A detailed description as well as a systematical-evolutionary sketch of the members of the tribe Soriculini are given.

An ancient stock of Soricids lived during the Pliocene and survived through the Lower Pleistocene (the groups Blar inini and Soriculini) in Europe; one of the character­istic systematic units of this group is the evolutionary line fo the tribe Soriculini K R E T Z O I , 1965, now confined only to Eastern and Southern Asia.

As known, H E X S E L ( 1S55) was the first to describe a member of this group as „Sorex" similis from Sardinia. B A T E (1945) established later, for this and related forms, the new genus Nesiotites, with the two further species corsicanus and hidalgo f rom Corsica and the Balearics.

From the European mainland only one form was hitherto known namely P E T É X Y I ' S "Crocidura" gibberodon (1864), originating from Beremend and later newly described, on the basis of the stratigraphically younger material from Villány, by KORMOS (1934) as "Soriculus kubinyii" (the identi ty of the two was shown by K R E T Z O I 1936). This single species was noticed to appear from the Middle Pliocene of Poland (Podlesice, KOWALSKI 1956) up to the Lower Pleistocene of Hungary (the stratigraphically last fossils f rom Locali ty 3 Villány, K R E T Z O I 1956). Finally K R E T Z O I ( 1959) proposed to establish for Sori-cuius gibberodon a new genus, Asoriculus, an opinion that was not followed by R E P E X X I X G (1967), who ranged this species wi th the genus Episoriculus.

KOWALSKI (1956) mentioned the size of the Podlesice-from to be somewhat smaller, and I also found an equally small form in the material of Osztramos J (JÁNOSSY 1972). I t was impossible to ascertain on the basis of these fragmentary remains, whether the differences constitute taxonomic or biostratigraphic values or not.

A well preserved finding of this group from Locali ty 13 Osztramos (a skull w i t h fu l l dentit ion together wi th the mandible of one side and a humerus of the same individual) threw new light upon the matter.

Locali ty 13 of Osztramos represents, in contrast to the other ones of the same h i l l , not a fissure but a level of carstic cavities, observable in the whole quarry system. I t con­sists of more or less eroded and horizontally situated holes (average thickness one hal f to one meter), partially filled wi th yellowish-brown sandy clay, parts of which containing the remains of bats in large numbers. The Episoriculus fossils collected by TOTAL , are the only ones which do not belong to Chiroptera (the description of these latter w i l l be pub­lished by TOPÁL).

The dark r e d clay of Locali ty 1, Osztramos (JÁXOSSY 1972), apparently penetrates the mat r ix of Loc. 13, a clear geological argument for the stratigraphically lower age fo the latter than the former one.

A detailed analysis of the dentition of the remains originating from Loc. 13, Osztra­mos, as well as a comparison w i t h the similar bones of the type of Soriculus kubinyii KORMOS , the material collected newly from Beremend and from Loc. 7, Osztramos, Csar-nó t i an age (for a faunistico-stratigraphic picture of the latter see JÁXOSSY 1973) decided the systematico-biostratigraphic: position of these forms. I consider them to represent a new species, described as follows.

Ann. Hist.-nat. Mus. Nat. Hung., 1973, 65

4 Természettudományi Múzeum Évkönyve 1973.

Episoriculus tornensis sp. n.

D e r i v a t i o n o m i n i s : tornensis, named after the Carstic Terr i tory w i t h the Osztramos H i l l in Torna.

T y p e l o c a l i t y : Loc. 13, Osztramos H i l l , N E Hungary (48° 30' Nor th . L a t . : 38° 25' East Long.)

T y p e l e v e l : Lower part of Middle Pliocene (or somewhat older). H o l o t y p e : Viscerocranium w i t h ful l dentition, broken behind the pterygoids.

Inventary number V . 72. 114, Collection of the Paleontological Department, Hungarian Natural History Museum, Budapest. — P a r a t y p e : Left mandible w i t h fu l l denti t ion and processi, w i t h only the proc. angularis lacking.

D i a g n o s i s : Small Episoriculus species with relatively reduced anterior dentition (including upper and lower incisors and antemolars) and chiefly with an extremely diminished last upper unicuspid.

D e s c r i p t i o n : Typical features of Episoriculus E L L E R M A N et MORRISON-SCOTT , 1951 (Soricidae G R A Y 1821, Soricinae M U R R A Y 1866, Soriculini K R E T Z O I 1965) dental formula 1-5-3/1-2-3, upper incisor bifid. Fourth upper antemolar very small, as in recent species of the genus. P 4 with protocone placed close to para-style, lower incisor short with a small undulation beyond the terminal cusp. Pig­mentation reduced : merely top of unworn incisors, paracone of P 4 and protoconid of M\ orange-coloured. Condyles of mandible widely separated. (Other details see below).

I had the possibility to compare my exemplars, among others, with the re­cent species Episoriculus caudatus and E. leucops as well as Soriculus nigrescens, all collected by G. T O P Á L in India. Ah these species possess a diminished last upper unicuspid and the variation of this tooth in size is not very large (the largest being at most twice that of the smallest). The last upper unicuspid of the type of "Sori­culus kubinyii K O R M O S " 1934 ("Crocidura gibberodon P E T É N Y I " 1864) from Villány-Kalkberg is about three to five times larger than that of the specimen from Loc. 13, Osztramos. The same may be observed on well preserved skull and maxillary remains originating from Loc. 7, Osztramos, — also a typical S. gibberodon (see Plate I , Fig. 1-6; the sketch of the upper dentition of "8. kubinyii K O R M O S " re­drawn on the basis of the original type-specimen). Therefore the possibility of an individual variation is precluded and this feature must have an important evolu­tionary significance. The variation of this tooth in E. borsodensis sp. n. described below from Loc. 1. Osztramos, will be discussed in that place.

The range of variation of the outer length of the upper incisor seems to differ only statistically from that of E. gibberodon, but absolutely from that of E. borso­densis sp. n. (1.80 mm in E. borsodensis sp. n. 1.48-1.72 mm [n=10]; in E. gibbe­rodon 1.75-2.07 [n=10]).

The three anterior upper unicuspids are in the occlusal view — agreeing with those of E. borsodensis — more quadratic and not so compressed antero-posteriorly as in E. gibberodon (see Plate I , Figs. I -4). Thus the length of the row of unicuspids, together with the diminished one is about the same with that of E. gibberodon and E. borsodensis (1.72 to 1.8 mm).

Against E. gibberodon P 4 is reduced (outer length 1.15 mm in contrast with E. gibberodon showing 1.4-1.5 mm [n=3], and E. borsodensis with 1.27 mm). The ratio of the outer length of P^/M1-!!2 is quite different in the three forms : E. gib­berodon 1.40/2.82; E. borsodensis sp. n. 1.27/2.55; E. tornensis sp. n. 1.15/2.65.

M 3 of E. tornensis is more reduced than in E. gibberodon, and intermediate between these two species in E. borsodensis.

Fig . 1. Scatter diagram showing outer crowndength and maximal wid th of the unworn lower incisors of different species of Episoriculus: 1. „Soriculus kubinyii K O R M O S " , L O C .

3, "Villány; 2. Episoriculus tornensis sp. n., Loc. 13, Osztramos; 3. E. gibberodon P E T É N Y I ,

Loc. 7, Osztramos; 4. E. borsodensis sp. n. , Loc. 1, Osztramos; 5. E. gibberodon, P E T N É Y I

Loc. 5, Beremend; 6. The same, Loc. 2, Csarnóta

The skull of E. borsodensis is unknown, that of the only specimen of E. tor­nensis is narrower (4.46 mm at M2) than in E. gibberodon (5.0 mm in the two speci­mens from Villány 3 and Osztramos 7).

The mandible of E. tornensis is stronger than that of E. borsodensis and i t agrees in this feature with E. gibberodon. Outer height of mandible below Mi : E. gibberodon: 1.18-1.46 mm (n=10); E. tornensis 1.33 mm ( n = l ) ; E. borsodensis 0.94-1.18 mm (n=10). Height of coronoid process: E. gibberodon 4.05; 4.40 mm; E. tornensis 4.05 mm; E. borsodensis 3.85; 3.90 mm. As mentioned in the diagnosis of the generic features, the articular facets of the condyle are widely separated. The lower articular facet is enlarged, in E. tornensis as. in E. gibberodon, and not so narrow as in E. borsodensis (see Plate I , Fig. 12-14). Related to the articular com­plex, the ascending ramus is weaker in E. tornensis than in E. gibberodon.

The lower incisor is in the outer (labial) view absolutely shorter and narrower in E. tornensis and E. borsodensis than in E. gibberodon (unworn incisor in E. tor-nensis: outer crown length 2.96 mm, width in outer view at middle undulation 0.83 mm; in E. gibberodon the measurements are same: 3.38/0.90 to 3.85/1.01; in E. borsodensis 2.65/0.82 to 3.07/1.00 mm ; see scatter diagram Fig. 1). The inner pos-

terior (unguium of the incisor subtends an acute angle in all specimens of the ex­amined material of all three species, as well as in recent Episoriculus, and it is not rectangular as described and illustrated by K R E T Z O I (1962 Plate I I , Fig. 1) as char­acteristic to the genus Asoriculus from Csarnóta — perhaps a southern endemic form (Plate I , Fig. 7 - 1 1 ) .

The lower antemolars of E. tornensis are reduced (length 1.45 mm), as in E. borsodensis (length 1 . 32 -1 .48 mm) and in contrast with E. gibberodon ( 1 . 7 0 - 2 . 0 0 mm). The first antemolar is more diminished as compared to the second one in E. tornensis than in E. borsodensis. The lower molars of E. tornensis stand, in their proportions as well as in the development of the endoconid, between those of E. gibberodon and E. borsodensis : thinner and weaker than in the former but thicker and stronger than in the latter one. The talonid of Jf3 is more reduced in E. tor­nensis than in E. gibberodon and E. borsodensis.

R e m a r k s : These rather subtile differences substantiate in my opinion not only the taxonomic independence of the new species originating from Loc. 1 and 13, Osztramos, but they also have an important evolutionary significance, to be discussed later.

As mentioned in the introduction, géomorphologie arguments speak for a low­er stratigraphie age of E. tornensis than for the other two forms. The taxonomic and stratigraphie arguments concur in speaking with a large probability for an older Middle Pliocene age, and perhaps also for a somewhat earlier stratigraphie level.

I do not know of other closely related fossil species than the two forms compared above in detail (E. gibberdon and E. borsodensis). As mentioned earlier, I had the possibility to compare my fossil findings with the hitherto known two recent spe­cies of the genus Episoriculus) E. caudatus H O R S F I E L D 1851, and E. leucops- H O R S -F I E L D , 1855. as well as with one species of the genus Chodsigoa: Ch. cf. salenskii K A S T S C H E N K O , 1907 (mandible of latter material originating from owl pellets, coll. T O P Á L . N-Vietnam) and finally, with the only species of Soriculus s. str. : S. nig-rescens G R A Y , 1-842.

A comparison of our fossils with Chodsigoa may be excluded by reason of the more reduced upper dentition (with only three upper unicuspids), by the very dif­ferent shape of the lower incisor, the weeker cingulum on the outer side of the lower molars, the relatively more reduced J / 3 and the differences in the ascending ramus of the mandible, etc. (For further details on this latter, see R E P E N N I N G 1967). The differences in dentition are considerable also in respect of Soriculus nigrescens. In Soriculus — among others — the last lower antemolar and M1-M2 and M\ is, in relation of the other molars, much more reduced than in Chodsigoa, Episoriculus or the fossil forms. A comparison with members of the recent Episoriculus reveals that our fossil species stands in several details, and chiefly in the reduction of the anterior dentition, nearer to the living species of the genus (chiefly E. leucops of the same size) than to the fossil E. gibberodon or E. borsodensis. Although the well expressed differences in the morphology of dentition etc. of the recent forms ex­clude unambiguously a specific identity, no a generic distinctness obtains.

Finally I received for comparison by courtesy of J . C H A L I N E (Dijon) remains of Nesiotites corsicanus B A T E , 1945. collected by C H A L I N E in the type-locality. The examination of these bones and dentition convinced me that Nesiotites represents a quite different evolutionary line than the Episoriculus group of the European continent (fresh, redcolored pigmentation of teeth, elongated lower incisor, as in Neomys, but strongly curved as in Chodsigoa. quite aberrant shape of the ascending ramus of the mandible etc.).

Episoriculus borsodensis sp. n .

D e r i v a t i o n o m i n i s: borsodensis, named after the administrative department Borsod wi th the locality Osztramos.

T v p e l o c a l i t y : Loc. 1, Osztramos Mil l , Ne — Hungary (48° 30 'Nor th . Lat . , 38" 25' East Long.)

T y p e l e v e l : Middle Pliocene, horizon of Estramontium (see Jánossy , 1972). H o I o t y p e: Left maxil lary fragment w i t h the penultimate and last unicuspids

and P 4 . Inventary number: V . 72. 115, Coll. Pal. Dep., Nat . Hist . Mus., Budapest. — P a r a t y p e : Anterior fragment of left mandible wi th the incisor, two antemolars and M1 (see JANOSSY 1972, Plate I I . , Fig. 3).

R e f e r r e d s p e c i m e n s : Nearly a hundred very fragmentary remains, all representing teeth and diverse mandibular parts.

D i a g n o s i s : A small Episoriculus species, with relatively reduced anterior dentition (chiefly upper and lower incisors), as in E. tornensis sp. n., but last upper unicuspid and first lower antemolar larger. Ratio of P4/AP~'~ and the articular fa­cets of mandible differing from that of all hitherto known fossil species.

D e s c r i p t i o n : I submitted a detailed sketch of the features of the genus Episoriculus as well as the comparison of E. gibberodon, E. tornensis sp. n. and E. borsodensis sp. n. in the course of the description of the former species. Suffice i t here therefore to emphasize only the most important features of the new species of Locality 1, Osztramos.

1 described these remains as ,,? Episoriculus sp." on the basis of a very scant material originating from the locality mentioned (JÁNOSSY 1972). Among the various maxillary fragments coUected earlier and in the latest material of exca­vations in 1972, I found one specimen with a well preserved last upper unicuspid and two former specimens in which this tooth is represented only by its alveolar holes (see Plate I , Fig. 6). I became convinced that the form from Locality 1, Osztra­mos, is in this feature clearly betwreen E. tornensis and E. gibberodon.

Beyond the diagnistic features, I have to emphasize the presence of quadratic (and not compressed) upper unicuspids, the less reduced first lower antemolar (against E. tornensis) and the narrower lower molars — all characteristics analyzed in detail in the course of the description of E. tornensis sp. n. given above. Pigmen­tation — if any — is the same as in E. tornensis: a light orange colour only on the top of the incisors and on the most prominent cusps of the last upper premolar and molars as well.

A comparison with recent species as well as with the insular fossil form Ne­siotites was also made above.

The remains are comparable stratigraphically and morphologically for Euro­pean fossil material, chiefly with the Podlesice-form (see K O W A L S K I 195(i. Plate I I Fig. 3) . This figure is for a comparison somewhat too small, but it still seems to show the brachygnathous form, which appears to agree with E. borsodensis sp. n.

Not only the forms from Beremend and Villány, but also the specimens ori­ginating from Csarnóta and Weze (by courtesy of M . K R E T Z O I , I received these latter from his collection) correspond with the elongated "dolichognathous" lower anterior dentition of E. gibberodon. Going by M E I N ' S exact figures of the recently published remains from Rhodos the characteristics are again the same, in spite of the smaller dimensions ( M E I N in: B R U I J N , DAWSON & M E I N 1970, Plate 11. Fig. lc).

As mentioned above one can hardly form a valid opinion of the evolutionary connection between E. gibberodon, E. tornensis and E. borsodensis. Against our hy-

pothetical supposition, the geologically oldest form — E. tornensis — is the con­siderably more evolved one (with the most reduced anterior dentition), and the stra­tigraphically youngest form E. gibberodon the most primitive.

Presumably, we have to do with collateral evolutionary lines which became extinct at different times.

References B A T E , D . M . (1945): Pleistocene shrews from the larger western Mediterranean Islands.

- Annales Mag. Nat. History, ser. 11 , v . 11; 738-769. B R U I J N , D E H . , DAWSON, M A R Y R . & M E I N , P. (1970): Upper Pliocene Rodentia,

Lagomorpha and Insectivora (Mammalia) from the isle of Rhodos (Greece) I I I . — Proc. Koninkl. Nederl. Akad. Wetensch., Ser. B, Vol . 73 . No. 5: 568-584.

H E N S E L , R. F . (1855) : Bei t räge zur Kenntnis fossiler Säuget iere ; Insectenfresser und Nage­tiere der Diluvialformation. — Z. Deutsch. Geol. Ges., 7 : 458-501.

JÁNOSSY , D . (1972): Middle Pliocene Microvertebrate Fauna from the Osztramos Loc. 1. (Northern Hungary). — Ann. Hist.-nat. Mus. Nat. Hung., 6 4 , 27-49.

JÁNOSSY , D . (1973): The Boundary of the Plio-Pleistocene based on the Microfauna in Nor th Hungary (Osztramos, Locali ty 7) — Verteb. Hung., 1 4 : i n press.

KORMOS , T . (1934): Neue Insectenfresser, F ledermäuse und Nager aus dem Oberpliocän der Villányéi- Gegend. - Földt. Közl., 6 4 , 296-321.

K O W A L S K I , K . (1956): Insectivores, bats, and rodents from the early Pleistocene bone brecea of Podlesice near Kroczyce (Poland). — Acta palaeont. Polonica, 1: 331-394.

KRETZOT , M . (1956): Die alfpleistozänen Wirbeltierfaunen des Villányer Gebirges. — Geol. Hung., Ser. Palaeont., 27 : 1-264.

K R E T Z O I , M . (1959) : Insectivoren, Nagetiere und Lagomorphen der Jüngs tp l iozänen Fauna von Csarnóta i m Villányer Gebirge (Südungarn) . — Verteb. Hung., 1 (2): 237-246.

K R E T Z O I , M . (1962) : Fauna und Faunenhorizont von Csarnóta . — Jatiresber. Ung. Geol. Anstalt f. 1 9 5 9 : 297-395.

K R E T Z O I , M . (1965) : Drepanosorex — neu definiert. — Verteb. Hung., 7 ( 1 - 2 ) : 117-129. P E T É N Y I , S.J. (1864): H á t r a h a g y o t t m u n k á i , I I : 37 -81 . R E P E N N I N G , CH. A . (1967) : Subfamilies and Genera of the Soricidae — Geol. Surv. Profess.

Paper, 5 6 5 : 1-74.

Explanation <>î Plate I . Rough sketch of the upper and lower dentit ion as well as the condylar process of different

Episoriculus species F ig . 1. E. tornensis sp. n . : r ight upper complete row, Loc. 13, Osztramos (Holotype). — Fig . 2. „Soriculus kubinyii K O R M O S " : the same, Loc. 3, Villány (redrawn on the original type specimen). — Fig . 3. E. gibberodon P E T É N Y I : r ight upper tooth row w i t h the de­tached incisor, the unicuspids, Pl and M1, Loc. 7, Osztramos. — Fig . 4. The same: r ight upper unicuspids and P 4 same locality. — Fig . 5. E. borsodensis sp. n . : the two last left unicuspids and P 4 , Loc. 1, Osztramos (Holotype). — Fig. 6. Right antepenultimate uni­cuspid, alveolar hole of last unicuspid and P 4 , same locality. — Fig. 7. E. gibberodon: outer view of anterior part of left mandible w i t h the incisor, the antemolars and the M1, Loc. 7, Osztramos. — Fig . 8. „Soriculus kubinyii", the same, Loc. 3, Villány (redrawn on the type specimen). — Fig . 9. E. borsodensis sp. n . , the same, Loc. 1, Osztramos. — Fig . 10. E. tomesis sp. n. , the same, Loc. 13. Osztramos. — Fig 11. Inner (labial) view of the same specimen. — Fig. 12. Posterior view of the condylar process of the same. — Fig . 13. E. borsodensis sp. n . the same, Loc. 1, Osztramos. — Fig. 14. E. gibberodon, the same, Loc. 7,

Osztramos.

Author's address: Dr. D. JÁNOSSY Paleontological Department of the Hungarian Natural History Museum 1088 Budapest, Múzeum körút 1 4 - H i . Hungary

Plate I.