taxonID	type	description	language	source
E12187BDFFCCD120C29B8FD9D38ABBC4.taxon	materials_examined	Material. MN 15: Včeláre 2 B: fragment of left md (ramus md), M 1. Ivanovce: right md with p 4 – m 3 (as Asoriculus gibberodon in Fejfar and Sabol 2005). MN 16: Javoříčko XI: complete left md except for the angular process. Javoříčko XVIII: right md without dentition. MN 17: Včeláre 6 / 1: fragment of right md without dentition. Koliňany 1: fragment of md without dentition (condylar process + part of corpus md preserved).	en	Pažitková, Barbora, Horáček, Ivan (2024): Shrews Of Tribe Nectogalini (Mammalia, Eulipotyphla, Soricidae) In The Fossil Record Of The Czech Republic And Slovakia. Fossil Imprint 80 (2): 190-228, DOI: 10.37520/fi.2024.017
E12187BDFFCCD120C29B8FD9D38ABBC4.taxon	description	Description and comparison. Overall, our material corresponds to the original diagnosis of Asoriculus gibberodon (absence or reduction of pigment, robust jaws, low coronoid process, pronounced spiculum, shallow upper sigmoid notch, compression of the unicuspid dentition, short i 1 with visible cingulum, compact m 3 with shortened talonid). We consider the significantly shorter distance between the condylar process and the ascending ramus (CdL) than in extant Neomys milleri to be an important diagnostic feature of Asoriculus gibberodon. Another notable difference is observed in the height of the condyle (Cd 1 Cd 2), which is lower in our MN 15 – MN 17 material; it also possesses a narrower upper facet compared to N. milleri. The last lower molar is also often slenderer than in the extant species. Apart from metric characteristics, differences in non-metric traits were observed as well, such as closer position of the teeth row to the ascending ramus, prolonged beanshaped mandibular foramen (which is usually round and small in recent taxa) and strongly pronounced spiculum of the coronoid process in the Plio-Pleistocene form. The cingulum is, in agreement with the recent species, present on both labial and lingual side. In the specimen from Javoříčko XI, it is also present on the lower incisor (unfortunately this specimen is the only one in our collection that possesses the lower incisor), which represents another difference from N. milleri. A faint light orange tooth pigmentation is present in various amounts; in samples of MN 15 – 16 age it is completely absent. The diagnosis (established at the description of Soriculus kubinyii KORMOS, 1934) emphasized “ the bicuspid lower incisor with insignificant cusps and weak pigmentation limited to the apical part of the teeth ”. Regarding the European fossil record, the following features are often mentioned: (i) the lower incisor is short and bicuspid (Reumer 1984, Koufos et al. 2001, Rofes and Cuenca-Bescós 2006, Minwer-Barakat et al. 2010, Angelone et al. 2011, Vasileaidou et al. 2012, Rzebik-Kowalska 2013, Joniak et al. 2017, Moya-Costa et al. 2023) with (ii) a welldeveloped cingulum (Reumer 1984, Dahlmann and Storch 1996, Furió and Angelone 2010, Minwer-Barakat et al. 2010, Rzebik-Kowalska 2013). (iii) Molars possess a rather weak cingulum (Rofes and Cuenca-Bescós 2006, Botka and Mészáros 2017, Joniak et al. 2017), (iv) the entoconid is pronounced (Rofes and Cuenca-Bescós 2006, Minwer-Barakat et al. 2010, Vasileaidou et al. 2012, Botka and Mészáros 2017). (v) Pigment is usually present only slightly, usually limited to tips of trigonid and talonid (Reumer 1984, Popov 2003, Rofes and Cuenca-Bescós 2006, Botka and Mészáros 2017, Moya-Costa et al. 2023). (vi) The terminal part of the coronoid process is round (Popov 2003, Rofes and Cuenca-Bescós 2006, Moya-Costa et al. 2023). (vii) The condylar process carries a wide but low upper facet of cylindrical shape, the lower facet is often bent lingually. The two facets are separated by long and extremely narrow interarticular area (Rofes and Cuenca-Bescós 2006, Rzebik-Kowalska 2013, Moya-Costa et al. 2023). (viii) The horizontal ramus is low (Rofes and Cuenca-Bescós 2006). Compared to it, our material shows a high level of variability. It conforms to the above characters in a narrow upper facet (Včeláre 2 B – MN 15, Včeláre 6 / 1 – MN 17, Včeláre 3 – MN 17, Včeláre 4 E – Q 1), low horizontal ramus (Včeláre 6 / 1 – MN 17, Včeláre 3 – MN 17, Včeláre 3 / 1 – MN 17), narrow tip of coronoid process (Včeláre 6 / 1 – MN 17), significant entoconid (Ivanovce – MN 15, Javoříčko XI – MN 16, Včeláre 3 / 1 – MN 17, Včeláre 4 / 7 – Q 1), and absence or reduction of pigment (Ivanovce – MN 15, Javoříčko XI – MN 16, Včeláre 4 / 7 – Q 1). In contrast, some of our samples possess a high horizontal ramus (Ivanovce – MN 15), wide tip of coronoid process (Ivanovce – MN 15, Včeláre 3 – MN 17, Včeláre 4 E – Q 1) or fuller pigmentation (Včeláre / upper molar / – MN 15, Včeláre 3 / 1 – MN 17).	en	Pažitková, Barbora, Horáček, Ivan (2024): Shrews Of Tribe Nectogalini (Mammalia, Eulipotyphla, Soricidae) In The Fossil Record Of The Czech Republic And Slovakia. Fossil Imprint 80 (2): 190-228, DOI: 10.37520/fi.2024.017
E12187BDFFCCD13DC15188C1D6FCBCD6.taxon	materials_examined	Material. MN 17: Včeláre 3: right md with m 1 – m 3 (as Episoriculus cf. castellarini in Fejfar and Horáček 1983), 4 fragments of left md (3 ramus md, 1 corpus md without dentition). MN 17 / Q 1: Včeláre 3 / 1: 1 part of ramus mandibulae (coronoid process), 1 part of md (corpus mandibulae) with molar tooth. Včeláre 5: fragment of left md (coronoid process). Q 1: Včeláre 4 E: fragment of right md (ramus md). Včeláre 4 A / 7: fragment of left md (corpus md) with m 2 – m 3 (as Episoriculus cf. castellarini in Fejfar and Horáček 1983). Q 1 / Q 2: Včeláre 6 / 7: right md with p 4 – m 1 (alternatively as Neomys newtoni HINTON, 1911).	en	Pažitková, Barbora, Horáček, Ivan (2024): Shrews Of Tribe Nectogalini (Mammalia, Eulipotyphla, Soricidae) In The Fossil Record Of The Czech Republic And Slovakia. Fossil Imprint 80 (2): 190-228, DOI: 10.37520/fi.2024.017
E12187BDFFCCD13DC15188C1D6FCBCD6.taxon	description	Description and comparison. Smaller size, more gracile teeth than Asoriculus gibberodon. Slight pigmentation of teeth, limited to the apical parts of the crown present in all specimens. Upper sigmoid notch not as shallow as in Asoriculus gibberodon, CdL in samples from Včeláre 3 / 1 Q which and) those NM Rec and (OTUs milleri Neomys compared extant of of range samples variation to into,) castellarini falling (cases age of 1 Q – percentages 17 MN – late %, of) those class to character age 17 MN particular – 15 MN for of obtained samples.) data <(Asoriculus metric them of of basic margin (comparisons cases variation of biometric number lower – * n under of. Summary newtoni fall or. N)> (. of 10 it Table sample exceed and Včeláre 6 / 7 significantly exceeds variance span of A. gibberodon, as former corresponds to smallest individual of N. milleri, and latter to mean value of this species. Coronoid process leans in mesial direction, similarly to Neomys. We were able to obtain biometric comparative information for 88 mandibular measurements (n = 3 – 12) of A. castellarini and A. gibberodon, for which the Shapiro-Wilk test showed normal distribution in 80 variables. Deviance from this state (p = 0.003 – 0.05) was observed in the following variables: Cr 3, Cr 5, RM 1, RM 2, mdLm 2, Cd 1 Cd 10, TrLm 1, TalLm 3. In five of them, values of the Q 1 and MN specimens differ distinctly: Cr 3, Cr 5, TalLm 3 (Q 1> MN) and RM 1, RM 2 (Q 1 <MN). The difference was also indicated by the results of PCA 4. Until the late MN 17, our material is homogenous and comprised the original form A. gibberodon, and similarly to the European fossil record (Rzebik-Kowalska 1998, 2002), this taxon represents the vast majority of our material. In contrast to our expectations, our samples did not follow the trend of enlarging body size throughout the MN 15 – Q 1 period. In most of the measurements, specimens from MN 15 exceeded the samples from MN 16 and MN 17 and reached values similar to the Q 1 material. Yet when the MN 15 specimens were excluded, the expected trend is confirmed. Similarly to our results, Popov (2003) also reported bimodality while comparing large sample of A. gibberodon from MN 17 Varshets, Bulgaria with other localities (MN 14 – Q 1). As expected, our results showed a smaller size of Asoriculus in comparison with Q 2 and Q 4 Neomys, as well as extant N. milleri. Sometimes an overlap with N. milleri was recorded. However, some samples exceeded the maximum of N. milleri. A wide terminal part of the coronoid process was observed in samples from Ivanovce (MN 15), Včeláre 3 (MN 17) and Včeláre 4 E (Q 1). The specimen from Ivanovce also exceeds the recent species in other aspects (pW, RmL), yet the distance between tip of upper facet and centre of upper sigmoid notch (lingual view, CdL) is shorter. This trend is observed in another 9 specimens (out of 11), and thus could be considered a significant distinguishing criterion between Asoriculus and Neomys. Similarly to Neomys, the width of the talonid of the first two lower molars exceeds the width of the trigonid, yet the m 3 in Asoriculus is more reduced (comp. smaller length and width of talonid). The sample from Včeláre 3 / 1 (the earliest Q 1) possesses a combination of characteristics mentioned above. There is not an enhanced gap between the molar row and ascending ramus, the coronoid process points almost straight upwards (without leaning forwards, as is typical for Neomys) and its spiculum is pronounced. The talonid of the last molar is small; the entoconid is relatively low in all molars. In terms of metric variables, this specimen is one of the largest, especially regarding the CdL. It corresponds well to the diagnosis and redescriptions of A. castellarini by van der Meulen (1973). In contrast, the specimen from Včeláre 6 / 7 (late Q 1) shares more characteristics with Neomys than Asoriculus. Even though the spiculum is pronounced, and according to the position of alveoli, the last molar was positioned closely to the ascending ramus, based on the value of CdL and the high coronoid process pointing dorsally, it could alternatively be assigned to Neomys newtoni. Apparently, Asoriculus gibberodon differs from Neomys newtoni significantly by the shape of the lower premolar (massive unicuspid with pronounced distal prolonging of the labial cingulum in Asoriculus, and wide bicuspid with significant occlusion edges of the cusps in Neomys). Unfortunately, not a single specimen from all of the European Q 1 fossil record possesses this particular tooth, except for those from Atapuerca, Trinchera del Elephante, Spain (Rofes and Cuenca-Bescos 2006), in which it corresponds clearly to the state in A. gibberodon, while at the same time, it differs from it by a dorsally tapered coronoid process and distinctly pronounced high position of the upper articulation condyle.	en	Pažitková, Barbora, Horáček, Ivan (2024): Shrews Of Tribe Nectogalini (Mammalia, Eulipotyphla, Soricidae) In The Fossil Record Of The Czech Republic And Slovakia. Fossil Imprint 80 (2): 190-228, DOI: 10.37520/fi.2024.017
E12187BDFFCCD13DC15188C1D6FCBCD6.taxon	discussion	Comments. Unfortunately, our material is not sufficient to test the relevance of the hypothesis that A. castellarini represents a transient form between A. gibberodon and N. newtoni, nor to prove the trend of enlarging body size during the period of MN 17 – Q 1. A similar issue stands behind the sceptical view on the validity of other forms described from various European Pliocene and Early Pleistocene sites: Soriculus kubinyii KORMOS, 1934, Episoriculus tornensis JÁNOSSY, 1973 and Episoriculus borsodensis JÁNOSSY, 1973. Considering the wide variability of Asoriculus gibberodon, these taxa are usually synonymised with this form (Reumer 1984, Rzebik-Kowalska 1998). Nevertheless, at least considering the span of variation among our OTU IV specimens, the tendencies conforming to expected transitional state between the Asoriculus and Neomys phenotypes seems to be clearly marked. Hence, we tentatively propose, as a provisional solution, to separate the late MN 17 – Q 1 populations as a distinct taxonomic unit, for which a prior name A. castellarini (PASA, 1947) is available.	en	Pažitková, Barbora, Horáček, Ivan (2024): Shrews Of Tribe Nectogalini (Mammalia, Eulipotyphla, Soricidae) In The Fossil Record Of The Czech Republic And Slovakia. Fossil Imprint 80 (2): 190-228, DOI: 10.37520/fi.2024.017
E12187BDFFD1D13DC2708FB1D38AB66B.taxon	description	In addition to the above performed comparisons, we analysed the phenotype relations between the studied mid-European taxa and a set of items representing the extant Oriental clades, namely Soriculus nigrescens, Episoriculus macrurus, E. leucops, Chimarrogale platycephalus (TEMMINCK, 1842), and Anourosorex sp. (Anourosoricini ANDERSON, 1879, formerly part of Neomyini MATSCHIE, 1909). In contrast to the latter two genera, which exhibit distinct differences, the representatives of Soriculus and Episoriculus reveal obvious similarities in cranial and dental characters to the European Pliocene gibberodon, for which that species was included in these extant genera (e. g., Ellermann and Morrison-Scott 1951, Repenning 1967), until Hutterer (1994) demonstrated the differences validating the concept of an independent genus Asoriculus, proposed by Kretzoi (1962). We examined the samples of these extant species using the biometric technique applied in other OTUs, and quantified the differences in form of Euclidean distances of 42 metric variables represented in the majority of the OTUs (unfortunately except for OTU V), normalized by filtering off their size differences, and further evaluated by PCA and cluster analyses. The comparison of frequency distributions for basic dental dimensions in particular OTUs is summarized in Text-fig. 9. The results of multivariate analyses demonstrating similarity relations among particular OTUs are visualized in Text-figs. 10 and 11. They show close relations among the W-Palearctic clades (Asoriculus-Neomys) and distinct differences from both Episoriculus and Soriculus samples. The results thus strongly support the stand of Kretzoi (1962) and Hutterer (1994) against alternative synonymisation of Asoriculus, Soriculus and Episoriculus (e. g., Ellermann and Morrison-Scott 1951, Repenning 1967). The case of Macroneomys brachygnathus (Q 2 / Q 3 border, 11 European sites) occupying the most distant position will be discussed elsewhere. European radiation is characterized by successive enlarging of body size and arrangements of horizontal ramus (including strengthening of the distal position of coronoid process) in the sequence AsorQ 1 - NeomQ 2, later followed by divergence to the morphotypes of Neomys milleri and Neomys fodiens during the Middle Pleistocene. Episoriculus macrurus BLANDFORD, 1888 is smaller in comparison with the extant European species, and there is occasional overlap with Neomys milleri. Its size is comparable to Asoriculus or Q 2 Neomys, while Episoriculus leucops HORSFIELD, 1855 is approximately of Neomys fodiens size (or exceeding it in some dimensions – e. g., pL, Cd 1 Cd 2). Therefore, it exceeds all the fossil forms in terms of size, similarly to Soriculus nigrescens. Overall, even though we witnessed some overlap in particular measurements (esp. E. macrurus with Asoriculus), they differ considerably in combinations of characters. We noticed several characteristics that are shared by all examined Asian species: (i) the upper sigmoid notch is shallow and round, (ii) the coronoid process is robust, especially in Soriculus but with the exception of E. macrurus, (iii) the entoconid is long (especially in Soriculus), but in Episoriculus it is not very significant, (iv) there are only three unicuspid teeth in the rostral part of the upper dentition (however, Soriculus carries a tiny residual Z 4 on the palatal edge of the P, (v) the first two upper molars possess a peculiarly shaped hypocone with strengthened cingulum, which is notably distanced from protocone. Even though we do not have access to the upper dentition of Asoriculus, according to Reumer (1984), this is a trait clearly distinguishing Asoriculus from Episoriculus and Soriculus.	en	Pažitková, Barbora, Horáček, Ivan (2024): Shrews Of Tribe Nectogalini (Mammalia, Eulipotyphla, Soricidae) In The Fossil Record Of The Czech Republic And Slovakia. Fossil Imprint 80 (2): 190-228, DOI: 10.37520/fi.2024.017
E12187BDFFD0D13FC1518967D304B931.taxon	description	Apart from N. newtoni, further three fossil species of the genus Neomys were reported from biozones Q 2 and Q 3: N. intermedius BRUNER, 1952, N. hintoni ZAITSEV et BARYSHNIKOV, 2002, and N. browni HINTON, 1911. Neomys intermedius is reported from two localities in Germany (Breitenberghöhle – Q 3 (Bruner 1957), Markgrabenhöhle – Q 3 (Bruner 1952 )) and followed by an uninformative description that is not sufficient for exact diagnosis. Neomys hintoni is reported from three localities from Caucasus: Treugolnaya Cave – Q 3 (Zaitsev a Baryshnikov 2002), Mezmaiskaya Cave – Q 3 (Zaitsev a Osipova 2004), Haykadzor – Q 2 (Tesakov et al. 2019). Even though the description is detailed and exactly defines the traits of this taxon, its taxonomic relevance should be further confronted with detailed information on phenotype variation of related species and dynamics of their local divergences, which are unfortunately still not available in a proper extent. The same can be said about Neomys browni, reported from two localities in the United Kingdom (Grays Thurrock – Q 3 (Hinton 1911), Cudmore – Q 3 (Roe et al. 2009 )), and diagnosed by a narrow and low coronoid process. With regards to a broad phenotype variation (also demonstrated in the present paper) as a characteristic feature of N. newtoni, we tend to consider N. hintoni, N. intermedius as the local forms of Neomys newtoni, alternatively Neomys milleri (pertinent in the case of N. browni). Numerous hypotheses regarding history and range dynamics of Neomys were proposed based on methods of molecular phylogenetics and phylogeography, palaeontology and morphological similarities. Molecular phylogenetics establishes the interval of divergence of the extant species between 0.4 and 1.6 Ma. According to Igea et al. (2015), Neomys fodiens was the first species to diverge (around 1.22 Ma), being the sister group of the rest of the species. Colonization of Europe during the Early Pleistocene followed by rapid expansion to northern and central European regions is presumed by Rzebik-Kowalska (1998) and Castiglia et al. (2007). According to Igea et al. (2015), Neomys teres separated from the common ancestor of N. milleri and N. anomalus 0.56 Ma. N. teres and N. anomalus / milleri could represent two lineages that survived Quaternary climatic oscillations in southern refugia: N. teres in Anatolia and N. anomalus on the Iberian Peninsula (Castiglia et al. 2007). This agrees with one of the hypotheses presented by Kryštufek et al. (2000), who assume that the common ancestor of Neomys milleri and Neomys anomalus crossed the Bosporus and colonized Anatolia during one of the glacials. Part of the population stayed in the region south from the Black Sea and later evolved in Neomys teres. An alternative hypothesis assumes expansion of Neomys fodiens during a cold period southwards across Balkan-Anatolian bridge, followed by a return northward during an interglacial. The authors presume that one population was isolated in the Pontic mountains, where it evolved into N. teres. The population living along the land bridge evolved into the ancestor of N. milleri / teres (Kryštufek et al. 2000). Based on morphologic features, the same authors present a third hypothesis: the possibility of expansion of N. fodiens along the shore of the Black Sea to Caucasus and the Pontic mountains, where as a result of allopatric speciation, it evolves into N. teres. However, this hypothesis does not correspond to the results of molecular genetics, hence Kryštufek et al. (2000) as well as Castiglia et al. (2007) and Igea et al. (2015) consider the possibility of N. fodiens diverging first and being the sister taxon of the rest of the species the most probable. On the contrary, Neomys milleri would be a relatively recent taxon. According to Castiglia et al. (2007), expansion of Neomys fodiens was faster in comparison to Neomys anomalus / milleri, due to N. fodiens possessing more developed adaptations to a challenging environment. Climatic oscillations would have had a more serious impact on the latter species. According to Castiglia et al. (2007), the N. anomalus-milleri clade shows a considerably higher degree of divergence. According to mitochondrial phylogenetics, N. anomalus comprises the population inhabiting the central region of the Iberian Peninsula, and N. milleri the rest of European population, including northeast of the Iberian Peninsula. These two lineages would have diverged 0.4 Ma, with a deviation 0.26 – 0.86 million years – in other words, during the Middle or Late Pleistocene (Igea et al. 2015). According to the authors, it is probable that N. anomalus evolved on the Iberian Peninsula during one of the Middle Pleistocene glacials, while N. milleri colonized the Pyrenees only during the latest Pleistocene or Holocene. The fossil record does not support hypotheses about older origins of N. fodiens and later divergence of N. milleri. The presence of N. fodiens in Europe is not proved until the Middle Pleistocene age (a few records, biozone Q 3); most of the records are dated to the present cycle (biozone Q 4). The hypothesis by Rzebik-Kowalska (1998) assuming colonization of Europe by N. fodiens during the Early Pleistocene still lacks any support from the actual fossil record. Considering its phenotype constitution, N. milleri can undoubtedly be regarded as the form retaining plesiomorhic patterns of the genus (comp. e. g., less reduced distal molars), while N. fodiens exhibits a derived state in most of the phenotype variables. It cannot be ruled out that the increase of N. fodiens size was related to the disappearance of another large shrew, Macroneomys brachygnathus, during early Toringian. The earliest records positively representing Neomys fodiens are reported from the Iberian Peninsula (TD 10 in Atapuerca, 370 ka; Moya-Costa et al. 2023). The respective specimens exceed the limits of our extant samples, particularly with the specimens (from the same layer) identified as “ Neomyini cf. Macroneomys ”. These extremely large phenotypes correspond to the extant Iberian form, Neomys fodiens niethammeri, which shows – against expectancy – only shallow genetic differences from other extant populations of the species (Balmori-de La Puente et al. 2019).	en	Pažitková, Barbora, Horáček, Ivan (2024): Shrews Of Tribe Nectogalini (Mammalia, Eulipotyphla, Soricidae) In The Fossil Record Of The Czech Republic And Slovakia. Fossil Imprint 80 (2): 190-228, DOI: 10.37520/fi.2024.017
