Skoczenia copernici ( Skoczeń, 1980 )

Rzebik-Kowalska, Barbara, 2014, Review of the Pliocene and Pleistocene Talpidae (Soricomorpha, Mammalia) of Poland, Palaeontologia Electronica (26 A) 26, pp. 1-26 : 17-20

Skoczenia copernici ( Skoczeń, 1980)

Figure 5.1 -8

zoobank.org/ EE3533BF-8577-4EC4-AADD-1643CAB3944D

1980? Geotrypus copernici Skoczeń ; (partim), Skoczeń, p. 414, pls. 3, 4, figs. 2, 3.

1994 “ Geotrypus copernici ” Skoczeń ; (partim), Rzebik-Kowalska, p. 80, 90, 91, 93.

2005 Geotrypus? copernici Skoczeń ; (partim), Rzebik-Kowalska, p. 129, 131.

2009 Scalopoides sp. =? G. copernici Skoczeń ; (partim), Rzebik-Kowalska, p. 8, 24.

Etymology. The genus is named in honour of late professor Stanisław Skoczeń from Kraków ( Poland) who passed away in 2007.

Holotype. Fragment of right mandible with p4-m2, last alveolus of p2 and two alveoli of p3 and of m3 (the same specimen as in the species “ copernici ”, no. MF/1008/17). It is housed in the collection of the ISEAPAS in Kraków.

Type locality. Kadzielnia 1.

Studied localities. Kadzielnia 1, Early Pleistocene (Late Villanyan, MN 17 or Pliocene/Pleistocene boundary); Zamkowa Dolna Cave A, Early Pleistocene (Late Villanyan, MN 17).

Material. Kadzielnia 1. Two fragments of left mandible – one with m1-m2, and one with p4, two toothless fragments of mandibles (left and right), and isolated teeth – two right P4, and two left M1. MNI = 3, catalogue number MF/1008. Zamkowa Dolna Cave A. Six (two right and four left) fragments of mandibles with p4-m2, isolated teeth – five P4 (three right), ten M1 (seven left), seven M2 (four left), five M3 (left), four p4 (three right), ten m1 (eight left), five m3 (four left). MNI = 8, catalogue number MF/1010. All material is housed in the collection of the ISEAPAS in Kraków.

In comparison to Skoczeń (1980) the material in Kadzielnia 1 lacks isolated teeth: one C1, one M2, one p4, four m1 and four m2. In Zamkowa Dolna Cave A one C1, three M3, one m1 and four m3 are missing. Teeth assigned as C1 are present in material from both localities but they are not the canines of moles.

Humeri from Kadzielnia 1 and Zamkowa Dolna Cave A cited by Skoczeń (1980) as “ G ”. copernici cannot be linked together with teeth of “ G ”. copernici as proposed by Skoczeń (1980) and they are tentatively assigned to? Scalopoides sp. (see above).The mandible from Kamyk ( Skoczeń 1980) also cited as belonging to “ G ”. copernici represents T. fossilis Petényi, 1864 ( Rzebik-Kowalska, 2009).

Diagnosis. A large species of Talpini mole with massive teeth, very well-developed parastyles, and clearly divided mesostyles in upper molars. Its dental formula of the mandible is 3i, 1c, 4p, and 3m. The lower canine is small and one-rooted, all lower premolars double-rooted, the p1 enlarged, and p2 and p3 small, almost equal in size, lying slightly obliquely in the mandible with their anterior roots situated more lingually. The lower p4 is long and lower molars diminish in size from m1 to m3 (m1>m2>m3). The horizontal ramus of the mandible tapers anteriorly, and its two mental foramina are situated below the p2/p3 junction and below the posterior root of p4.

Differential diagnosis for the genus. In spite of an unreduced (complete) dental formula in the mandible (22 teeth) and large size as in most Pliocene and Pleistocene European Desmaninae Thomas, 1912 Skoczenia n. g. differs from these by the morphology of upper molars which are devoid of a protoconule (present in Desmaninae ), by a large p1 (small in desmans), by the p4 devoid of a metaconid (present in desmans) and by the oblique cristid of the lower molars, which ends buccally (in Desmaninae situated either lingually against the tip of the metaconid or against the protoconid-metaconid crest) ( Rümke, 1985).

Gaillardinae Hutchison, 1968 – aquatic desman-like moles from North America – also differ from Skoczenia n. g. by morphology of their mandibles in which a posterior mental formen is situated below the posterior root of m1 (in Skoczenia n. g. it is situated below the posterior root of p4), and by lower molars in which the oblique cristid reaches the lingual side and the metastylids ( Hutchison, 1968). The teeth of Skoczenia n. g. are devoid of metastylids and the oblique cristid is short and only reaches the buccal (m1) or middle sides of the teeth (m2, m3).

Large fossil Suleimaninae Van den Hoek Ostende, 2001 from Anatolia ( Turkey) with reduced dental formulae (only two upper molars present in jaw, Van den Hoek Ostende, 2001) cannot be taken into consideration because of the several very prominent M3s present among the specimens of Skoczenia n. g. from Kadzielnia 1 and Zamkowa Dolna Cave A.

The large size, complete lower dentition (22) and the hypertrophy of p1 with two roots (which can be concluded from mandible no. MF/1010/56) exclude Skoczenia n. g. from the Scalopinae [both Condylurini ( Condylura ) and Scalopini ( Parascalops , Scalopus , Scapanus , Scapanulus, Hugueneya , Proscapanus , Leptoscaptor , Yanshuella , Yunoscaptor , Scalopoides , Mydecodon, Scaptonoscaptor , Proscalops , and? Domninoides )]. Scalopinae , known from Eurasia and North America, are generally small, they have a reduced dental formula, their p1 is small with one root or this tooth is absent. If the dental formula of some genera of Scalopinae is unreduced and p1 has two roots as in Condylura or in Scaptonoscaptor, the first form is much smaller than Skoczenia n. g., its p1 is small and the lower molars have metastylids, and the second, Scaptonoscaptor, although large, also has a small p1 and small metastylids absent in Skoczenia n. g. Hugueneya has a two-rooted p1 as in Skoczenia n. g., but it is not very large and moreover its m1 is smaller than the m2 whereas in Skoczenia n. g. the m1 is visibly larger than the m2 ( Van Zyll de Jong, 1983; Wilson, 1960; Hutchison, 1968; Storch and Qiu, 1983, 1991; Van den Hoek Ostende, 1989; Ziegler, 2003).

The small shrew-moles - Uropsilinae Dobson, 1883 from Eurasia and North America include genera such as Uropsilus Milne-Edwards, 1871 , Mystipterus Hall, 1930 , Desmanella Engesser, 1972 , Theratiskos Van den Hoek Ostende, 2001 , Mygatalpa Schreuder, 1940 , Nasillus Thomas, 1911 , and Rhynchonax Thomas, 1912 , with small or absent p1s are also quite different from Skoczenia n. g. and can be easily excluded as its relatives.

Skoczenia n. g. also differs from the old (Oligocene – early Miocene) extinct American moles of the subfamily Proscalopinae K.M. Reed, 1961 raised by Barnosky (1981) to family ( Proscalopidae K.M. Reed, 1961 ) level by having an unreduced dental formula (in Proscalopidae only one species, Proscalops tertius K.M. Reed, 1961 , has an unreduced formula, Barnosky, 1981), by the presence of bifid mesostyles in upper molars, by the absence of metastylids in lower molars, by enlarged p1 and by m1>m2.

The subfamily Talpinae consists of tribes Urotrichini Dobson, 1883 , Scaptonychini Van Valen, 1967 , Neurotrichini , and Talpini .

Eurasiatic Urotrichini [( Urotrichus Temminck, 1841 = Dymecodon True, 1886 ), Myxomygale Filhol, 1890 , Paratalpa Lavocat, 1951 , Tenuibrachiatum Ziegler, 2003 , Urotrichus (Scaptonyx) dolichochir ( Gaillard, 1899) ] differ from Skoczenia n. g. by small size and reduced dental formula, i.e., 36 – 42 teeth. Only Myxomygale has a full set of teeth but its p1 is small, one-rooted, its m2>m1, and upper and lower molars have distinct cingula ( Ziegler, 1990). The enigmatic Urotrichus (? Scaptonyx ) dolichochir Gaillard, 1899 described on the grounds of the humerus is also smaller and quite different in morphology.

Asiatic Scaptonychini with one genus Scaptonyx Milne-Edwards, 1872 (Recent form – S. fusicaudatus Milne-Edwards, 1872 ) can also be excluded. This form is large and it has a double-rooted p1 as Skoczenia n. g., but the dental formula of its mandible is reduced (3,1,3,3, Gureev, 1979), and its lower molars have metastylids which are absent in Skoczenia n. g.

Although this character (the presence or absence of metastylids) seems unstable because in 14 genera of Scalopini seven ( Parascalops , Scapanulus , Proscapanus , Yunoscaptor , Domninoides , Scalopoides , Scapanoscapter ) have metastylids and seven do not ( Scalopus , Scapanus, Hugueneya , Leptoscaptor , Yanshuella , Mydecodon , and Proscalops ), Scaptonyx additionally differs from Skoczenia n. g. by the lack of a tooth cingulum, a different position of crista obliqua in the lower molars and by a larger m2 than m1.

The fossil “ Scaptonyx ” edwardsi Gaillard, 1899 has teeth and mandible similar to the Recent Scaptonyx ( Gureev, 1979) and it is also different from Skoczenia n. g.

The North American and Eurasian Neurotrichini ( Neurotrichus , Quyania ) are also small shrew-like moles with reduced teeth (mandible dental formula = 3-1-2- 3 in Neurotrichus and 3-1-3- 3 in Quyania ), and thus they also cannot be united with Skoczenia n. g.

Skoczenia n. g. is most similar to representatives of the tribe Talpini . The tribe comprises such genera as Euroscaptor Miller, 1940 , Mogera Pomel, 1848 (= Nesoscaptor Abe, Shiraishi and Arai, 1991 ), Parascaptor Gill, 1875 , Scaptochirus Milne-Edwards, 1867 , Geotrypus Lavocat, 1951 , and Talpa Linnaeus, 1758 . All Recent forms are known from Asia with the exception of Talpa living in Eurasia. They are characterized by a large or very large p1 with two roots as in Skoczenia n. g. They have, however, a reduced dental formula ( Parascaptor and Mogera 42, Scaptochirus and Geotrypus , e.g., G. ehrensteinensis 40 and Nesoscaptor (now included into Mogera ) 38, with the exception of the Recent Euroscaptor (included by some authors in Talpa ) and Talpa which have 44 teeth. However, in comparison to Skoczenia n. g., Euroscaptor is smaller and has a very small p2, much smaller than p3 ( Abe et al., 1991).

Representatives of the genus Mogera have the mental foramen of the mandible situated between the roots of m1 and its m2>m1 which differentiates them from Skoczenia n. gen. Scaptochirus has large molars and lower molars diminishing in size from m1>m2>m3 as in Skoczenia n. g., but contrary to the latter, a very small, one rooted p3 and very narrow pointed coronoid process ( Abe et al., 1991). In contrast to Skoczenia n. g., Parascaptor is rather small, it has p2 and p 3 in one line parallel to the long axis of the mandible and its p2 is very small, considerably smaller than p3.

Besides the dental formula, which in some species is unreduced, Geotrypus differs from Skoczenia n. g. by the lack of a protoconule, metaconule, and metastyle in upper teeth and by its mesostyle only slightly divided or undivided. In addition, its lower molar m1 is always smaller or equal to m2 and its crista obliqua is situated more lingually ( Crochet, 1995; Van den Hoek Ostende, 2001).

Skoczenia n. g. also differs from Recent and fossil species of Talpa . Its teeth are much stronger and contain a cingulum below the hypoflexid of the lower molars and in the posterior walls of the p4 and P4 as well as on the buccal side of M1 above the metacone. Its upper teeth have deeply fissured mesostyles, distinct metaconules, its lower p4 is elongated, lower molars with mesoconids and grooves situated between the oblique cristids and the posterior wall of the trigonids, posterior mental foramen situated below the posterior root of p4 and the tip of the coronoid process narrow and curved posteriorly.

Fossil Talpinae incertae sedis, e.g., Desmanodon Engesser, 1980 known from the Miocene of Europe and Asiatic Turkey, the Early Miocene Nuragha de Bruijn and Rümke, 1974 - endemic to Sardinia and only known from isolated teeth as well as Eotalpa anglica Sigé, Crochet and Insole, 1977 from the Middle – Late Eocene of Wight Island, apart from their different morphology, are much smaller.

Description of holotype. It is represented by a fragment of mandible with p4-m2 and alveoli of m3 as well as two alveoli of p3 and second alveolus of p2. The horizontal ramus of the mandible is heavy and tapered anteriorly. There are two mental foramina, one beneathp2/p3 and the other below the posterior root of p4. The alveoli of the p3 are slightly oblique and its anterior roots are situated slightly lingually.

Teeth are massive but unfortunately very worn. The p4 is very long and narrowed in the middle. Its protoconid is situated more or less in the middle of the tooth, the antero-labial side is convex, the postero-labial side is slightly basined, and the lingual side is almost straight. The anterocrista (paracristid) is sharp, the posterocrista very worn so it is unknown if the metaconid was present. The talonid is very short. The parastyle and cingula are lacking.

The m1 is also very long and its trigonid is longer than the talonid. The crista obliqua terminates bucally, below the protoconid, the hypoflexid is shallow and a trace of the cingulum is visible below it, the entoconid is wide, the entostylid rather small, and metastylid and remaining cingula absent.

The m2 is smaller than the m1, and its trigonid and talonid are more or less of the same length. Crista obliqua terminates more lingually, in the middle of the posterior wall of the trigonid (at the level of the protocristid notch), and the hypoflexid is deeper and cingulum below it more prominent. The parastylid and entostylid are present, and the metastylid and cingula are absent.

Description of the remaining material. A detailed description is in Skoczeń (1980). Here some of the most characteristic features are repeated and considering the description of the new genus the specimens were measured once more.

P4 is robust, its paracone is high, rounded anteriorly. The sharp posterior ridge extends from its tip to the posterior cingulum and is terminated by a small cuspule. The protocone is well developed, the parastyle in some specimens is also well developed but in others less so. A feeble cingulum surrounds the tooth with the exception of the protocone. It is more protruding in the buccal than in the lingual side.

The M1 has a very large parastyle, a deeply divided mesostyle, and large protocone and metaconule. A protoconule is lacking. The cingulum is also absent with the exception of the fragment on the buccal side beyond the metacone. The M2 has a large parastyle, large divided mesostyle, large metaconule, and very small protoconule.The M3 is similar to M1 and M2 with a strongly divided mesostyle, large parastyle, and a trace of a protoconule, but the cingulum is absent.

In the other mandible with unworn teeth (p4 - m2), the p4 is also long but its anterocrista is less sharp, talonid slightly longer, and a parastyle or a fragment of anterior cingulum is visible. In m1 a small mesoconid and a notch (between crista obliqua and posterior wall of the trigonid) are present, and the cingulum below the hypoflexid is more developed. Mesoconids and grooves are also visible in the m1 of the mandibles from Zamkowa Dolna Cave A (MF/1010/, 58 – 60).

The most important is mandible no. MN/1010/ 56. It bears a p4, the first alveolus of m1 and alveoli before p4. There are four alveoli of p3 and p2 more or less equal in size and placed slightly obliquely to the long axis of the mandible, with the first root of both teeth deflected to the lingual side. Before them are two large alveoli of p1, and subsequently four for c (the largest of these four) – i1.

Measurements. See Table 4.

Systematic position. Mandible No. 56 from Zamkowa Dolna Cave A and the remaining material from this locality as well as from Kadzielnia 1 are homogeneous and represent the same mole species as the holotype from Kadzielnia 1. The long p4 and the posterior mental foramen of the mandible situated below the posterior root of p4, characters unknown in other genera, discriminate this form (as shown above) from other moles known so far. The large size, 22 teeth in the mandible, double-rooted lower premolars, small lower c, and enlarged p1 allow for an unambiguous assignment to the tribe Talpini . However, the remaining characters do not allow its inclusion into Recent and fossil genera of Talpini .

Within the Talpini an unreduced tooth formula is present only in Euroscaptor and Talpa ; however, the differential diagnosis shows that both forms also differ in morphology from Skoczenia n.g. In this situation the massive teeth and mandibles from Kadzielnia 1 and Zamkowa Dolna Cave A were described as a new genus, and the species “ copernici ” as established by Skoczeń (1980). The humeri previously associated with these teeth by Skoczeń (1980) were transferred to the genus? Scalopoides . It is strange, however, that humeri corresponding in size to the Skoczenia teeth were not found in the material.

Figures

FIGURE 5. Skoczenia copernici from Zamkowa Dolna Cave A (1-6, catalogue numbers MN/1010/3, 10, 18, 28, and 58 respectively) and from Kadzielnia 1 (7-8, catalogue number MF/1008/17). 1 - left P4, 2 - left M1, 3 - left M2 and 4 - left M3 (occlusal view), fragment of right mandible with p4-m1 (holotype), 5 - buccal, 6 - lingual view, fragment of right mandible with p4-m2, 7 - buccal, 8 - lingual view.