Two Crocidura and three Sorex species are reported from the Early Pleistocene Somssich Hill 2 locality (Villány Hills, Southern Hungary): C. kornfeldi KORMOS, 1934; C. obtusa KRETZOI, 1938; S. minutus LINNAEUS, 1766; S. runtonensis HINTON, 1911 and S. margaritodon KRETZOI, 1941. We compared the number of the Crocidura and Sorex occurrences in the 50 layers of the site and drew a sketch of the surrounding ecotypes. Layers 50–47 represent warmer climate with deciduous forests. From layer 46 cold climate and open grassland (steppe) appeared in the surroundings of the locality. In layers 33–4 mixed vegetation with forests and open grassland areas occurred varying in composition. Within this period layers 20–4 are characterized by the stronger dominance of the open vegetation. In layers 3–1 there are not enough remains to make a certain conclusion concerning the vegetation

Mészáros, Lukács

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IntroductionThe Somssich Hill 2 site (Villány Hills, Southern Hun-gary) is one of the most important Pleistocene vertebratelocalities of Hungary. It was discovered by Dénes Jánossyand György Topál in 1974. The excavation led by thembetween 1974 and 1984 yielded rich Pleistocene fauna(approximately 1.0–0.9 Ma old). The locality and the history of the excavation is describedby BOTKA & MÉSZÁROS (2015, in Hungarian) and a briefEnglish overview is given in BOTKA & MÉSZÁROS (2014a).The preliminary lists of the fauna were given in JÁNOSSY(1983, 1986, 1990). After the years of the excavations onlysome mammal groups were described by HÍR (cricetids,1998) and JÁNOSSY (1983, 1990, lemmings and arvicolids).Moreover the molluscan fauna of the locality was com-pletely elaborated by KROLOPP (2000).The rich material collected by the excavation team ofJánossy is kept in the Department of Palaeontology andGeology of the Hungarian Natural History Museum.Most of the Somssich Hill 2 material is under elabo-ration by the cooperative research group of the HungarianAcademy of Sciences, the Hungarian Natural HistoryMuseum and the Eötvös Loránd University (OTKAK104506 research, project leader: Piroska Pazonyi). On thebasis of their studies some preliminary results (BOTKA &MÉSZÁROS 2014b, BOTKA & STRICZKY 2014, MÉSZÁROS et al.2013, PAZONYI et al. 2013a, b, PAZONYI & VIRÁG 2013a, b,PAZONYI 2015, SZENTESI 2013) and articles (BOTKA &MÉSZÁROS 2014a, 2015a, b; STRICZKY & PAZONYI 2014;SZENTESI 2014) have been published.In the present article a palaeoecological study is demon-strated based on the occurrence of the species of two Soricidaegenera (Crocidura and Sorex) in the layers of the site.Material and methodsSome of the shrew specimens from Somssich Hill 2locality are more or less complete mandibles, but most ofthem are small mandibular and maxillary fragments orseparated teeth. Due to this fact it would have been a hardtask to settle the minimum number of the individuals. Thatis why an effective method of using soricid remains in thePalaeoecological reconstruction was suggested by MÉSZÁROS(2015). The number of Crocidura and Sorex teeth wascompared layer by layer without species attribution. On thebasis of the Crocidura-Sorex teeth rate a preliminaryecotype reconstruction of the site was given in the article. Later in 2015 a detailed elaboration of the Crociduramaterial was accomplished by BOTKA & MÉSZÁROS (2015b).This study provided a revision of previous taxonomicalPalaeoecology of the Early Pleistocene Somssich Hill 2 locality (Hungary) based on Crocidura and Sorex (Mammalia, Soricidae) occurrencesLukács MÉSZÁROSEötvös University, Department of PalaeontologyH–1117 Budapest, Pázmány Péter sétány 1/C, Hungary, e-mail: lgy.meszaros@gmail.comHantkeniana 10, 147–152., Budapest, 2015Barnabás GÉCZY Jubilee VolumeAbstractTwo Crocidura and three Sorex species are reported from the Early Pleistocene Somssich Hill 2 locality (VillányHills, Southern Hungary): C. kornfeldi KORMOS, 1934; C. obtusa KRETZOI, 1938; S. minutus LINNAEUS, 1766; S.runtonensis HINTON, 1911 and S. margaritodon KRETZOI, 1941. We compared the number of the Crocidura and Sorexoccurrences in the 50 layers of the site and drew a sketch of the surrounding ecotypes. Layers 50–47 represent warmerclimate with deciduous forests. From layer 46 cold climate and open grassland (steppe) appeared in the surroundings ofthe locality. In layers 33–4 mixed vegetation with forests and open grassland areas occurred varying in composition.Within this period layers 20–4 are characterized by the stronger dominance of the open vegetation. In layers 3–1 there arenot enough remains to make a certain conclusion concerning the vegetation.Keywords: Early Pleistocene, Crocidura, Palaeoecology, Somssich Hill, Sorex, Soricidae, Villány Hillsattribution of some teeth formerly defined as “Crocidura”.The preceding palaeoecological model settled by MÉSZÁROS(2015) needed modification in some points because of thenew data which is shown in the present article.Some alterations in the calculation methods were madeconsidering the previous paper. The rate of the Crocidurateeth is figured out according to the formula given here:The ocurrence of Crocidura and Sorex teeth in the layersof Somssich Hill 2 locality is shown in Table 1.TaxonomyPhylum Vertebrata LINNAEUS, 1758Classis Mammalia LINNAEUS, 1758Order Eulipotyphla WADDELL et al., 1999Family Soricidae FISCHER VON WALDHEIM, 1817Seven shrew species from Somssich Hill 2 site werementioned by BOTKA & MÉSZÁROS (2014b): Beremendiafissidens (PETÉNYI 1864), B. minor RZEBIK-KOWALSKA, 1976and the here discussed two Crocidura and three Sorexspecies. However, the Beremendia species are not includedin this palaeoecological research. They are described indetail by BOTKA & MÉSZÁROS (2014a). Subfamily Crocidurinae MILNE-EDWARDS, 1874Genus Crocidura WAGLER, 1832Dental formula is 143/123. The teeth are not pigmented.The first upper antemolar is larger than the equal-sizedsecond and third antemolars, all of them are unicuspid (afterBOTKA & MÉSZÁROS 2015b, REUMER 1984, REPENNING1967). It is present in the European mainland from the EarlyVillányian (MN 16) to Recent.Crocidurinae prefer relatively milder and more aridclimate compared to Soricinae (RZEBIK-KOWALSKA 1995).According to REUMER (1984) Crocidura prefer dry terrains.That is why these forms are good indicators of more-or-lessopen grasslands in the warmer phases of the Plio-Pleisto-cene (RZEBIK-KOWALSKA 1995). However, in this study wesupposed that low temperature limited their occurrence,even if steppe areas appeared in the colder periods of thePleistocene.The Somssich Hill 2 Crocidura material is described indetail by BOTKA & MÉSZÁROS (2015b).Crocidura kornfeldi Kormos, 1934(Figure 1)Morphology:  the coronoid process of the mandible isblunt and low, the coronoid spicule is small, situated highand not distinct. The anterior part of the coronoid process148 MÉSZÁROS, L..: Palaeoecology of the Early Pleistocene Somssich Hill 2 locality based on Crocidura and Sorex (Mammalia, Soricidae)Table 1. Ocurrence of Crocidura and Sorex teeth in the layers ofSomssich Hill 2 localityleans slightly backwards and the ramus sometimes has awidening in the middle part. The condyle reaches farbackwards and is comparatively large in buccal view. Thecondyle is not high in posterior view, the interarticular areais short. The internal temporal fossa is large and open,reaching halfway up the coronoid process. The regionunderneath the internal temporal fossa (the subfossa) isexcavated, separated from the fossa by a protuding ridge.The dental morphology of this species is very similar to thatof C. obtusa, the only differential character is the higherposition of the buccal re-entrant valley on M1–M2 (BOTKA &MÉSZÁROS 2015b).Crocidura obtusa KRETZOI, 1938(Figure 2)Morphology: the coronoid process of the mandible ishigh, its tip leans strongly backwards, the coronoid spiculeis situated high and distinct. The upper part of the condylereaches far backwards in buccal view. The condyle is high inposterior view, the interarticular area is long. The internaltemporal fossa is large and open, reaching halfway up thecoronoid process. The subfossa is sometimes present. There-entrant valley of the first and the second lower molarsopens closer to the buccal cingulum here than at C. korn-feldi.Genus Sorex LINNAEUS, 1758Teeth are pigmented with dark red to light orange colour.Fossa temporalis interna of the mandible is large andtriangular. There is an entoconid crest on the lower first andsecond molars (REUMER 1984).Dental formula is 1–6–3/1–2–3. M3 talonid is basined.Lingual cingulum is well-developed on M1–M3, but it isstraighter than on Crocidura forms. Articular facets of themandibular condyle are slightly to moderately separated(REPENNING 1967).The genus is present in Europe from the Late Miocene toRecent (RZEBIK-KOWALSKA 2003).Sorex endure much colder temperature than Crocidura(RZEBIK-KOWALSKA 2015). S. minutus, the only Sorexspecies in the Somssich Hill 2 fauna that is still living isfound even in the northernmost and coldest regions ofEurope. According to REUMER (1984) the fossil Sorex is indi-cative for a moist environment with well-developed (forest)vegetation and Sorex minutus suggests the presence ofwoody covering.We accept REUMER’s opinion, however, we cannot besure in the ecological role of the genus in the site before theexact specific determination of Sorex specimens, because S.runtonensis is reported from some localities as a member ofa steppe assemblage (OSIPOVA et al. 2006). Nevertheless it will not be necessary to fully convertthe model presented here, because S. runtonensis has itsgreatest abundance in layers 47–35, where steppe wassupposed on the basis of the rodent species (see below).Sorex margaritodon KRETZOI, 1941(Figure 3)It is a strongly built Sorex species with light orange pig-mentation on the teeth. Hantkeniana 10 (2015) 149Figure 1. Crocidura kornfeldi KORMOS, 1934, left mandible fragment withM1–M3, Somssich Hill 2, layer 30. Scale bar = 1 mmFigure 2. Crocidura obtusa KRETZOI, 1938, left mandible with I1 fragment, A2and M1–M3, Somssich Hill 2, layer 12. Scale bar = 1 mmFigure 3. Sorex margaritodon KRETZOI, 1941, right mandible fragment withM1–M2, Somssich Hill 2, layer 45. Scale bar = 1 mmMÉSZÁROS, L..: Palaeoecology of the Early Pleistocene Somssich Hill 2 locality based on Crocidura and Sorex (Mammalia, Soricidae)150Sorex minutus LINNAEUS, 1766(Figure 4)This Sorex species is a very fragile built one, it has darkred pigmentation on the teeth. Sorex runtonensis HINTON, 1911(Figure 5)It is a medium sized Sorex species, the cusps of the teethare light to dark red-coloured. Palaeoecological conclusionsCrocidura is used as an indicator of and open, grassyvegetation, while Sorex is regarded as the marker of forestsin the ecotype changing model. In addition we consideredenvironmental features shown by rodent occurrences in thedifferent layers (Piroska PAZONYI, personal communi-cation).While rodents indicate warm temperature, Crociduraspecies are totally missing from layers 50–47. That is whywe supposed warmer climate and deciduous forests withcomplete Sorex dominance. In the time of the deposition of the layers 46–34 steppevegetation is marked by the rodent fauna. However theabundance of Crocidura species is not high. Theexplanation of this fact is that their spread is usually limitedby low temperature, even if the ecosystem would beadequate to their preference. This is supported by the European occurrence of theliving Crocidura species, the diversity of which isdecreasing toward the northern areas – where they aremissing from the cold grasslands (RZEBIK-KOWALSKA 1995).So we may suppose that cold climate and open grassland(steppe) appeared in the surroundings of the locality in thisperiod. Sorex species tolerating low temperature well weresignificant here. In the interval represented by layers 33–4 there arefrequent Crocidura forms as well. On the basis of thealternating frequency of the two genera we conclude mixedvegetation with forests and open grassland areas occurringvarying in relation.Figure 4. Sorex minutus LINNAEUS, 1766, right mandible with I1 fragment, A2and M1–M2, Somssich Hill 2, layer 38. Scale bar = 1 mmFigure 6. The number of the Crocidura teeth relatively to the whole Crocidura and Sorex teeth material in the layers of Somssich Hill2 locality with the supposed ecotypes: 1 – warmer climate with deciduous forests, 2 – cold climate and open grassland (steppe), 3 –forest-steppe with less grassy areas, 4 – forest steppe with greater grassy areas, 5 – not enough remains for the ecotype-reconstructionFigure 5. Sorex runtonensis HINTON, 1911, complex left mandible with thewhole dentition, Somssich Hill 2, layer 45. Scale bar = 1 mmWithin this period grassy areas became larger in sizefrom layer 20. This is indicated by the greater significance ofCrocidura forms beside Sorex ones.  The remains found in layers 3–1 were not enough fordrawing a well-founded outline on the vegetation (Figure6).AcknowledgementsThe researches were supported by the HungarianGovernment, Ministry of Human Capacities (UT-2014/2015-0001 project) and the Hungarian Research Found(OTKA K104506 project). The author is indebted to theleader and the members of the OTKA Research Group fortheir valuable suggestions and kind help in the work:Piroska PAZONYI, Dániel BOTKA, Mihály GASPARIK, ZoltánSZENTESI, and Attila VIRÁG.Hantkeniana 10 (2015) 151ReferencesBOTKA, D. & MÉSZÁROS, L. 2014b: Beremendia (Mammalia, Soricidae) remains from the late Early Pleistocene Somssich Hill 2 locality(Southern Hungary) and their taxonomic, biostratigraphic, palaeoecological and palaeobiogeographical relations. — FragmentaPalaeontologica Hungarica 31, 79–111. BOTKA D. & MÉSZÁROS L. 2014a: A Somssich-hegy 2-es lelőhely alsó-pleisztocén Soricidae faunája. — In: BOSNAKOFF M. & DULAI A.(eds): Book of Abstracts of 17. 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Palaeoecology of the Early Pleistocene Somssich Hill 2 locality (Hungary) based on Crocidura and Sorex (Mammalia, Soricidae) occurrences

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