Type species

Crossopus fissidens Petényi, Reference Petényi1864.

Other species

Beremendia minor Rzebik-Kowalska, Reference Rzebik-Kowalska1976; Beremendia pliocaenica Flynn and Wu, Reference Flynn and Wu1994; Beremendia jiangnanensis Jin et al., Reference Jin, Zhang, Sun and Zheng2009.

Diagnosis

Blarinini-like condyle with a broad interarticular area and anteriorly placed lower facet; heavily pigmented teeth; upper incisor strongly fissident; upper antemolar row with four teeth (A1–A4); M1 and M2 with a developed protoconule and metaloph; lower incisor acuspulate; entocristid of the lower molars are presented.

Occurrence

Early Pliocene to Middle Pleistocene in Europe and early Pliocene to early Pleistocene in Asia.

Comparisons

Beremendia differs from Lunanosorex in: (1) the straight cutting edge of the first lower incisor (Lunanosorex displays one or two denticulations on this edge; Qiu and Storch, Reference Qiu and Storch2000, pl. 3; Jin et al., Reference Jin, Sun and Zhang2007, figs. 2, 3); (2) the shape of the articular surface of the condylar process (Lunanosorex has a longer upper facet; Jin et al., Reference Jin, Sun and Zhang2007, fig. 2); and (3) the longer and lower positioning of the lateral groove of the first upper incisor (Lunanosorex has a groove length of ~1/3 of the root length, and the distance between its lower edge and the root lower edge corresponds to the groove height; Qiu and Storch, Reference Qiu and Storch2000, pl. 3).

Beremendia differs from Peisorex in the number of upper antemolars (Jin and Kawamura, Reference Jin and Kawamura1996a) and in the presence of a metaloph of M1 and M2. However, conditions of the upper teeth of Peisorex pohaiensis Kowalski and Li, Reference Kowalski and Li1963 are only known from one partial skull obtained from the Haimao locality (Jin and Kawamura, Reference Jin and Kawamura1996a), and for the most part specimens are unavailable. Beremendia and Peisorex do not differ in mandibular characteristics due to the presence of overlapping features.

Beremendia differs from Dolinasorex in: (1) the more posterior positioning of the zygomatic process of maxilla (Dolinasorex exhibits an anterior shift in this process to the level of the M2 metastyle; Rofes and Cuenca-Bescós, Reference Rofes and Cuenca-Bescós2009, fig. 3), (2) its uncompressed upper antemolar row (the P4 parastyle in Dolinasorex comes into contact with the posterobuccal corner of A2); and (3) its moderately developed talonid of m3 (the talonid in Dolinasorex is short).

Figure 3.

Teeth and dentary fragments: Beremendia fissidens (Petényi, Reference Petényi1864): (1–3) first upper left incisor from SHM/A (ZIN 104908/1236): (1) buccal view; (2) occlusal view (cut image); (3) lingual view; (4) first upper right incisor from Korotoyak 2 (ZIN 104903/1292); (5, 6) first lower left incisor from KMN (ZIN 104923/1264): (5) medial view; (6) lateral view (cut image); (7) first lower right incisor from ANR/1C (ZIN 104919/1257, i1), medial view; (9) right dentary fragment (cut image) with i1, p4 from SHM/A (ZIN 104908/1222), medial view. Beremendia minor Rzebik-Kowalska, Reference Rzebik-Kowalska1976: (8) right dentary fragment (cut image) with i1–p4 from BRB/C (ZIN 104894), medial view. a1 = first lower antemolar; AAF = area of additional features; p4 = fourth lower premolar; star indicates reversed image. Scale bars = 1 mm.

Remarks

Some authors follow Jin and Kawamura (Reference Jin and Kawamura1996a) in accepting a broad concept for the genus including Chinese forms (e.g., Jin et al., Reference Jin, Zhang, Sun and Zheng2009; Rofes and Cuenca-Bescós, Reference Rofes and Cuenca-Bescós2009; Flynn and Wu, Reference Flynn, Wu, Flynn and Wu2017), which were previously referred to the separate genus Peisorex. Most known fossil material of beremendiins is represented by the presence of the mandible and isolated teeth, which does not permit the determination of the genus. However, we conservatively follow Qiu and Storch (Reference Qiu, Storch, Hoek Ostende, Doukas and Reumer2005) in considering Beremendia and Peisorex as separate valid genera. The validity of the latter is determined at least by the original dental formula (Jin and Kawamura, Reference Jin and Kawamura1996a).

Accordingly, remains obtained from the Chiachiashan (= Jiajiashan) type locality and Haimao Chinese early Pleistocene (Jin and Kawamura, Reference Jin and Kawamura1996a) locality are identified as those of Peisorex pohaiensis. The generic position of the Chinese form ‘pliocaenicus’ (Flynn and Wu, Reference Flynn and Wu1994) remains problematic because its skull structure remains unknown. Before the emergence of new fossil data, this taxon was considered a part of Beremendia (Flynn and Wu, Reference Flynn, Wu, Flynn and Wu2017). The description of the enigmatic ‘Nectogalinia altaica’ is based on a poor description of damaged hemimandible fragments (Gureev, Reference Gureev1979), and Nectogalinia should be placed with Beremendia until new data become available. Thus, Beremendia represents a ‘wastebasket’ taxon for any Beremendia-like remains of the mandible (without the first incisor) or of isolated teeth.

According to Reumer (Reference Reumer1984), both Beremendia fissidens and Beremendia minor are identical in morphology and differ only in size with the latter being considerably smaller. Other species of Beremendia sensu lato taken from Chinese sites (e.g., Peisorex pohaiensis) are of a similar size to Beremendia fissidens or are markedly larger than European species (e.g., Beremendia pliocaenica and Beremendia jiangnanensis; see S8–S12). Thus, fossil remains from any Asian locality generally represented by hemimandible fragments and isolated teeth without clear diagnostic features can be classified morphometrically into one of three size groups: small shrews likely corresponding to Beremendia minor; medium-sized shrews likely corresponding to Beremendia fissidens (not including P. pohaiensis, see ‘Comparisons’ above); and large shrews likely corresponding to Beremendia pliocaenica and Beremendia jiangnanensis. Botka and Mészáros (Reference Botka and Mészáros2014) have described features of the m3 talonid to distinguish Beremendia fissidens and Beremendia minor from the early Pleistocene Somssich Hill 2 locality in Hungary. These differences were not observed in materials from Siberia, Kazakhstan, and Mongolia (detailed comparisons given below). Therefore, original materials used in the present work were metrically separated into small and medium-sized shrews (S13–S14).

Type material

Syntypes, V 61.1585 (HNHM), one skull fragment, two left maxilla fragments, two right maxilla fragments, 13 left hemimandibles, 15 right hemimandibles, one condylar process, eight isolated first lower incisors; Beremend 2, Villány Hills, Hungary; late Pliocene, MN 16 (Botka and Mészáros, Reference Botka and Mészáros2014).

Diagnosis

Mandibular condyle with a broad interarticular area and anteriorly placed lower facet; heavily pigmented teeth; heavily fissident upper incisor; upper antemolar row with four teeth (A1–A4); M1 and M2 with developed protoconule and metaloph; lower incisor acuspulate; entocristid of lower molars presented.

Description

Most remains from Siberian, Kazakh, and Mongolian localities belong to medium-sized shrews (S13–S14). Rare rostral parts of skulls from SHM/A (ZIN 104908/1227, /1228) and BRB/C (ZIN 104909) exhibit the dental formula 1.4.1.3/1.1.1.3, corresponding to Beremendia sensu stricto (to Beremendia fissidens in this case).

Dentition

The teeth are heavily pigmented. The first upper incisor includes a large crown and a notably smaller and moderately curved root. The crown apex is distinctly fissident, and both tips are pigmented. The buccal cingulum is well developed along the margins of the talon with triangular thickening occurring at the talon base; crown margins are positioned above the talon base level without cingulum. The talon is complex: the buccal part forms a hatchet-like vertical cutting ridge, and a distinct lingual part includes a moderately developed medial cusp that extends along the lingual side of the talon as a developed, vertical, cingulum-like, short torus (‘lingual torus’). The pigmented talon cutting ridge is positioned on a relatively unworn tooth from SHM/A (ZIN 104908/1228). The root is of equal thickness along its full length. The lateral side of the root includes a deep triangular groove (‘lateral groove’) that extends along half of the root (unlike in Lunanosorex); the groove is positioned close to the lower edge of the root (Fig. 3.1–3.4).

Only four fragments are known of the premaxilla-maxilla with unicuspid antemolar teeth A1–A4 or their alveoli (see S3). The relative sizes of the antemolars are A1 > A2 > A3>>A4. The first and second upper antemolars (A1, A2) are elongated in occlusal view with a marked posterobuccal shifting of the crown relative to their longitudinal axes. The third upper antemolar is half the size of A2 and more rounded with the same posterobuccal shifting of the crown. A1–A3 with well-developed medial longitudinal ridges from anterior to posterior margins passing through the main cusp and dividing the crown into smaller lingual and larger buccal halves (S15, fig. 1). Together with the incisor talon vertical ridge, medial ridges of A1–A3 form a common longitudinal cutting crest within the anterior part of the upper tooth row (‘complex cutting ridge’). The main cusps of the antemolars are pigmented. The fourth upper antemolar is vestigial, rounded in shape without distinct crown structures, and visibly pigmented. The first three antemolars display well-developed ectocingula and entocingula.

The fourth upper premolar (P4) is subtrapezoidal with a large, well-developed, pigmented, and anteriorly shifted parastyle connected with the paracone base at a well-distinguished parastylar crest. A visible preprotoconal crest extends along the shallow, concave anterior edge of the tooth crown positioned between the parastyle and protocone; together with the postprotoconal crest, it clearly draws contours of the protoconal part of the trigone basin. The tip of the protocone is pigmented. The hypocone condition reveals two morphotypes. Morphotype I includes no hypocone (Fig. 4.1–4.2); the ridge of the hypoconal flange begins at the elevated part with a gradual decline in height. Morphotype II includes a bulged hypocone on the hypoconal flange edge (Fig. 4.4–4.6). The posterior emargination of P4 is well developed. The parastyle is half the height of the paracone in lateral view; the upper outline in lateral view is slightly undulating with visible signs of mesial inflection; the ectocingulum is well developed and broad along the parastyle-paracone bases and becomes narrower with length before reaching the end of the upper edge of the tooth crown (Fig. 4.3).

Figure 4.

Skull fragments, Beremendia fissidens (Petényi, Reference Petényi1864): (1–3) right maxilla fragment with P4 from SHM/A (ZIN 104908/1230); (1) occlusal view (cut image); (2) explanatory drawing; (3) P4 (cut image), lateral view; (4, 5) left maxilla fragment with P4–M1 from SHM/A (ZIN 104908/1229): (4) occlusal view; (5) explanatory drawing; (6) fragment of lingual half of P4 (cut image) of specimen ZIN 104908/1229, occlusal view. M1 = first upper molar. Scale bars = 1 mm.

The first and second upper molars (M1, M2) differ in size from front to back; both teeth are subquadrate in outline, but M1 includes a trigon that is narrower than its talon (means: AW(_M1) = 2.47 mm vs. PW(_M1) = 2.47 mm; see S13), whereas M2 includes a wider trigon (AW(_M2) = 2.72 mm vs. PW(_M2) = 2.30 mm; see S13). The parastyle of M1 is conical; the paracone is 2/3 of the metacone height; the mesostyle is not bifid and displays a rounded, typically unpigmented tip; the metastyle represents the end of the postmetaconal crest without any bulges or cusplets. Sharp, long, well-pigmented crests connect all cones and styles of the buccal side. The buccal outline of the tooth in occlusal view is slightly concave at the mesostyle level. The protocone is long and broad at its base; the preprotoconal crest forms a well-distinguished protoconule, forming a bulge within the crest. The protoconule bulge and crestal bend form a visible extension of the anterior tooth outline before reaching the protocone level. Simultaneously, the end of the postprotoconal crest is connected to a developed metaloph forming a straight ridge (Fig. 5.3, 5.4). All crests pass through the protoconule, protocone, and metaloph, clearly separating the deep protoconal basin. The hypocone is a weakly distinguished cusp and is conical in the basal extension of the hypoconal flange. The lingual outline of the tooth is slightly stepped due to a shallow notch positioned in front of the hypoconal flange (Morphotype I; see Fig. 5.1, 5.2); it also includes a developed notch in front of the hypoconal flange (Morphotype II; see Fig. 5.5, 5.6). The posterior emargination of M1 is well developed. Cingula are absent, aside from the short and weak entocingulum under the protocone (Fig. 5.1, 5.2).

Figure 5.

Upper molars: Beremendia fissidens (Petényi, Reference Petényi1864): (1, 2) first upper right molar from BH-103/19 (ZIN 104924/1271): (1) occlusal view, white arrow marks shallow notch of M1 hypoconal flange outline; (2) explanatory drawing; (3, 4) fragment of lingual part of M1 of specimen ZIN 104924/1271: (3) occlusal view (cut image); (4) explanatory drawing; (5, 6) fragment of lingual half of left M1 from SHM/A (ZIN 104908/1229): (5) occlusal view (cut image), white arrow marks developed notch of M1 hypoconal flange outline; (6) explanatory drawing. Beremendia cf. B. fissidens (Petényi, Reference Petényi1864): (7) first upper right molar from ORH/1B (ZIN 104911), occlusal view; (8, 9) fragment of lingual part of M1 of specimen ZIN 104911: (8) occlusal view (cut image); (9) explanatory drawing. Star indicates reversed image. Scale bars = 1 mm.

In general, the second upper molar is similar to the first but differs in: (1) trigon and talon width relations (AW vs. PW; see above and S13); (2) its relatively bulged metastyles of M2; and (3) its more developed, concave buccal outline M2 in occlusal view. The structure of the third upper molar (M3) remains unknown.

The first lower incisor is straight without denticles on the cutting edge and with a tapered tip bent upward (Fig. 3.6); the tip is heavily pigmented. Longitudinal axes of the incisor crown and root are continuous; the root length (from the lower outline) is ~1/3 of the crown length. The medial side of the incisor root includes a long (~1/2 of the root length), deep groove. The crown medial groove is long and well developed. The posteromedial area of the crown (‘Area of Additional Features’ or AAF) presents two different conditions: Morphotype I with smooth AAF without any folds and grooves (Fig. 3.5, 3.7) and Morphotype II with a groove (Fig. 3.9). The ectocingulid is very short and distinguishable only from the upper part of the crown base from the anterior part of a1 to the level of p4.

The first lower antemolar (a1) is relatively small and moderately procumbent; its crown overlaps with the incisor; the main (protoconid) cusp is heavily anteriorly shifted; the buccal arm of the posterocristid extends posterolaterally from the protoconid along with a surface ending with a cuspule adjacent to the fourth premolar. The posterolingual basin is shallow and delineated. The ectocingulid is well developed, narrow at the base of the protoconid, and gradually broadens toward the posterolateral end of the crown; entocingulids on the two investigated teeth of SHM/A (ZIN 104908/1221, /1223) are rounded, and their shapes and degrees of development remain unknown. The size and shape of the root are also unknown. The main cusp and part of the posterocristid are pigmented.

The fourth lower premolar (p4) is soricine-like in shape with a strong anterior shifting of the high protoconid and a well-developed, short, high posterocristid buccal arm ending at the developed additional cusp (‘cusp’); the posterocristid is slightly buccally shifted from the longitudinal axis of the tooth; an additional cusp is situated farther away before reaching the posterobuccal corner of the tooth (Fig. 6.1, 6.2). The posterolingual basin is shallow and slightly inclined toward the lingual side. In general, the crown is asymmetrical in shape with a shorter lingual half and with a longer, strongly exaenodont buccal half of the tooth. The size and shape of the roots are also unknown. The ectocingulid and entocingulid are well developed. The main cusp and buccal side of the posterocristid are heavily pigmented.

Figure 6.

Dentary fragments: Beremendia fissidens (Petényi, Reference Petényi1864): (1, 2) fragment of the right dentary from SHM/A (ZIN 104908/1222): (1) medial view (cut image); (2) explanatory drawing; (3) fragment (cut image) of the right dentary from SBH (ZIN 104893/02), occlusal view; (5) fragment (cut image) of the right dentary from SBH (ZIN 104893/01), lateral view; (6) third lower right molar (cut image) from SBH (ZIN 104893/02), occlusal view; (8, 11) third lower right molar from SBH (ZIN 104893/02): (8) lateral view; (11) explanatory drawing of lingual side of the talonid, white arrow marks gap of entocristide. (4) Nectogalinia altaica Gureev, 1979, holotype from RZD (GIN 664/201), diagrammatic image of m1 and m2 of occlusal view. (7, 9, 10) Beremendia minor Rzebik-Kowalska, Reference Rzebik-Kowalska1976, third lower right molar from BRB/C (ZIN 104894): (7) occlusal view (cut image); (9) explanatory drawing of lingual side of the talonid; (10) lingual view (cut image). a = line indicating longitudinal axis of tooth crown; a1 = first lower antemolar; b = left-turned arrow indicating anterior tilt of protocristid of m1 relative to a; c = right-turned arrow indicating weak tilt of protocristid of m2 relative to a; d = height of m3 entocristid; e = height of entocingulid of m3 relative to d; i1 = first lower incisor; p4 = fourth lower premolar; star indicates reversed image. Scale bars = 1 mm.

The lower molars are graded in size: the first molar is the largest, the second is slightly smaller, and the third is even smaller (Fig. 6.3). Teeth m1 and m2 differ in their absolute dimensions (means: L_(m1) = 2.58 mm vs. L_(m2) = 2.26 mm) and include trigonids of comparable lengths: the trigonid of m1 is ~70–71% the total length of the tooth, whereas that of m2 is ~68–69% the total length. The trigonid of m1 is narrower than that of m2 (S13). The paraconid of m1 is anteriorly shifted, and the protoconid is large and anteriorly shifted; hence, the protocristid is slightly inclined relative to the longitudinal axis of the tooth crown. The metaconid forms a small cusp with its tip extending higher than the paraconid tip. The hypoconid forms a large cusp. The well-defined hypolophid ends without a bulge or cusplet and with a distinct but narrow gap between its base and that of the entoconid (Fig. 6.3). The entoconid is a well-developed conical cusp that reaches the base of the metaconid; its height from the upper margins of the entocingulid is twice that (or slightly more) of the entocingulid. The precingulid, ectocingulid, posterocingulid, and entocingulid are well developed as narrow, clearly delineated belts; the lower tooth outline undulates with visible inflections at the protoconid level on both the buccal and lingual sides. The main conids, cristids, and trigonid basin are pigmented.

In general, the morphology of the second lower molar is similar to that of the first with two main differences: (1) having relatively comparable m2 trigonid (TRW) and talonid (TAW) widths (means: TRW = 1.30 mm vs. TAW = 1.31 mm; see S13) and (2) the nearly perpendicular positioning of the protocristid of m2 relative to the longitudinal axis of the crown (Fig. 6.4).

The lower third molar (m3) includes a developed trigonid and a smaller talonid; the talonid is narrow with a shallow, rounded basin; the borders of the basin are formed by a continuously ‘oblique cristid–hypolophid–entocristid’ without a distinct hypoconid or entoconid; the entocristid does not reach the metaconid base (Fig. 6.6), and the talonid basin is opened by a narrow gap (Fig. 6.11). Only the precingulid and ectocingulid are well defined (Fig. 6.8); the others are weak and smooth. The buccal sides of the trigonid cusps and cristids and the buccal side of the talonid are pigmented.

Skull fragments

Fragments of a rostral part of the skull were collected from two Mongolian localities: SHM/A (N = 4) and BRB/C (N = 1) (see S3). The maxillary fragments with preserved infraorbital canal areas include a canal with wide lateral walls; the anterior margins of the wall are rounded and situated above the P4-M1 junction (S15, Fig. 1); the posterior margin is wider than the anterior margin and is also rounded and situated above the M2 metacone cusp (or is shifted slightly posteriorly) (see S15, Fig. 1). The lower part of the wall within the upper molar roots is strongly sculptured, including convexities and concavities within the bone above and between the roots; in the inferoposterior corner of the wall there is a well-distinguished undulating row of attached muscle (Blarina-like condition; likely M. levator nasi inferior; Nikolskaya, Reference Nikolskaya1965). Its lacrimal foramen is positioned at the M1 mesostyle and lies at the anterior third of the wall of the infraorbital canal. The relatively small zygomatic process of the maxilla of SHM/A (ZIN 104908/1234) extends laterally; its tip is an acute, obliquely oriented, short crest without any bulges situated behind the M2-M3 junction; the process is half the length of the postmetacrista of M2. The preserved fragment of nasal and frontal bones of BRB/C (ZIN 104909/1241) includes a relatively large, paired frontal foramina (Sorex-like condition); the foramina are located immediately behind the border of the cribriform plate.

Mandible

The mandibular body is robust and wide (high); its ventral margin includes a weak ledge situated within the center of the m1 crown; due to the presence of this ledge the dentary includes a gradually tapered contour from the anterior end to the posterior end. Moreover, the ventral margin includes two digastric muscle attachments with the first taking the form of a small, smooth spicule (slightly bulged facet) positioned under the end of m3 and shifted buccally from the vertical plane; the second includes a weak longitudinal facet positioned between the end of the symphysis (level of m1 talonid) and the spicule (Fig. 7.1). The mental foramen is positioned within a slightly tilted groove under the hypoconid of m1 (from the anterior end to the hypoconid). The mandibular ramus is moderately long and large; its anterior margin is tilted posteriorly and bends in the upper part so that the tip of the coronoid process is heavily anteriorly tilted. The coronoid process includes a well-developed coronoid spicule and two well-delineated external fossae for the temporal muscle (‘external fossa’) positioned anterior and posterior to the spicule (Fig. 7.3, 7.4); the internal fossa of the temporal muscle (‘internal fossa’) is deep and clearly delineated. The base of the condylar process in medial view includes a wide and shallow subtriangular fossa with a moderately distinguished pterygoid spicule. The condylar process is distinct; the posterior subtriangular surface includes a broad interarticular area; the upper articular facet is short and tilted; the lower facet is longer than the upper facet and lacks a free lateral end due to the positioning of the dorsal margin of the angular process (i.e., the facet rests against the base of the angular process; see Fig. 7.2). The angular process is short and robust with an obtusely pointed, medially bent tip. The mandibular foramen is large, connects to the internal fossa, and is positioned at the toothrow baseline.

Figure 7.

Dentary fragments: Beremendia fissidens (Petényi, Reference Petényi1864): (1) diagrammatic image of the right mandible dentary fragment from SHM/A (ZIN 104908/1222), lateral view; (2) left condylar process (cut image) from KMN (ZIN 104923/1263), articular view; (3, 4) right mandibular ramus of specimen from SBH (ZIN 104893/01): (3) lateral view (cut image); (4) explanatory drawing. Beremendia minor Rzebik-Kowalska, Reference Rzebik-Kowalska1976: (5, 6) right mandibular ramus of specimen from BRB/C (ZIN 104894): (5) lateral view (cut image); (6) explanatory drawing; (7) diagrammatic image of the right (reversed image) condylar process from BRB/C (ZIN 104894), articular view; (8) diagrammatic image of the right mandible dentary fragment from BRB/C (ZIN 104894), lateral view; (9) anterior part (cut image) of the right dentary from BRB/C (ZIN 104894), lateral view; (10, 11) lower teeth a1–m1 of the right dentary from BRB/C (ZIN 104894): (10) occlusal view (cut image); (11) explanatory drawing. a = area of a digastric muscle attachment (‘weak longitudinal facet’); a1 = first lower antemolar; b = anterior portion of the lateral temporal fossa; c = a posterior portion of the lateral temporal fossa; i1 = first lower incisor; m1, m2, m3 = first, second, and third lower molars; p4 = fourth lower premolar. Scale bars = 1 mm.

Nectogalinia altaica

The holotype from RZD (GIN 664/201) includes a fragment of the right hemimandible with m1 and m2. The teeth include wide and well-distinguished cingulids and include visible PPt (see S5, fig. 4, table 4). The third lower molar is missing, but the anterior root is present in its alveolus. The lateral wall of the first lower incisor is heavily damaged; the alveoli of a1 and p4 and the posterior alveolus of m3 are filled with solid sand. The symphysial area of the dentary is damaged; the spicule within the ventral margins is markedly expressed; the facet of the digastric muscle is well distinguished along the lower side of the dentary and is shifted posteriorly on the lateral side. Complete coronoid and condylar processes represent the mandibular ramus, but the angular process has broken off (see S15, fig. 2).

Variation

The studied remains of Beremendia fissidens show variations in tooth pigmentation, characteristics, and dimensions.

Comparisons

The remains of Asian Beremendia fissidens investigated here are mostly comparable in size (e.g., the length of m1 and m2) to Beremendia fissidens collected from European sites (S8, S9). However, two fossils—the holotype of Nectogalinia altaica (GIN 664/202) and a specimen collected from the Sibiryachikha Cave (ZIN 104893/01)—include a smaller m1 (Fig. 8, ‘Na_h’ and ‘893/01’). The coefficient of variation (CV, %) for m1 length for the investigated material is 5.03% larger than that of materials collected from European localities: Osztramos 7, 4.37%; Csarnóta 2, 4.88%; Villány 3, 4.10%; Tegelen, 3.71%; and Osztramos 3/2, 4.71% (calculated with S8 measurements). The mandibular dimensions of the Asian fossils are comparable to (or slightly smaller than) their European counterparts (MRH_(Asia) = 5.88–6.84 mm vs. MRH_(Europe) = 5.93–7.48 mm; see S8, S13). Qualitative characteristics of the Asian specimens vary considerably or at least more than those of specimens from European sites (e.g., Botka and Mészáros, Reference Botka and Mészáros2014, figs. 10–12; see S7). The most visible differences are: (1) two morphotypes of hypocone conditions of the fourth upper premolar (European materials include only the second morphotype; Fig. 4); (2) two morphotypes of the lingual margin outline of the first upper molar (European materials include only the second morphotype; Fig. 5); and (3) two morphotypes of AAF of the first lower incisor (European materials include only the first morphotype with a slightly developed, short groove; Fig. 3.7). Furthermore, the European fossils include a weaker or even absent ectocingulid of the first lower incisor than that observed in Asian material.

Figure 8.

Measurements of the m1 and m2 of Beremendiini samples: (1) common graph for eight species: Beremendia fissidens (Petényi, Reference Petényi1864) (Bf); Beremendia jiangnanensis Jin et al., Reference Jin, Zhang, Sun and Zheng2009 (Bj); Beremendia minor Rzebik-Kowalska, Reference Rzebik-Kowalska1976 (Bm); Dolinasorex glyphodon Rofes and Cuenca-Bescós, Reference Rofes and Cuenca-Bescós2009 (Dg); Lunanosorex lii Jin and Kawamura, Reference Jin and Kawamura1996b (Ll); L. qiui Jin, Sun, and Zhang, Reference Jin, Sun and Zhang2007 (Lq); Na = Nectogalinia altaica Gureev, Reference Gureev1979; Peisorex pohaiensis Kowalski and Li, Reference Kowalski and Li1963 (Pph); Beremendia [Peisorex] pliocaenica Flynn and Wu, Reference Flynn and Wu1994 (Ppl); (2) detail of graph with variability area of European samples of Beremendia fissidens (dark gray polygon as in 1), original material of Bf (three- and four-digit numbers), Ll, Pph, and Ppl. Bf-B5 = Beremendia fissidens from Beremend 5 locality (Hungary; late Pleistocene); Bm_RKr = Beremendia minor from Rębielice Królewskie I (Poland; late Pliocene); Bm_SH = Beremendia minor from Somssich Hill 2 locality; h = holotype; h1 = holotype dimensions from original description of Pph (IVPP V2671); h2 = holotype of Pph redimensions by Jin and Kawamura (Reference Jin and Kawamura1996a); h4 = holotype dimensions from original description of Ppl (IVPP V8900); h5 = redimensions of Ppl holotype by Jin and Kawamura (Reference Jin and Kawamura1996a) (see S7); L(m1) = m1 length; L(m2) = m2 length; Ll_HX = L. lii from Houxushan Hill locality (China; late Pliocene); Ll_QP = L. lii from Qipanshan Hill locality (China; late Pliocene); Pph_HAI = P. pohaiensis from Haimao locality (China; early Pleistocene); r = redimensions herein of replicated Pph holotype (see S6). Also see S2.

The lengths of lower molars in the investigated specimens of Beremendia fissidens are larger than those of Beremendia minor (S9; Fig. 8). Qualitative differences observed between Beremendia fissidens and Beremendia minor in the shape of the m3 hypolophid (Beremendia fissidens with “straight posterior margin < m3 talonid basin>”) and high entoconid (Beremendia fissidens; the “entoconid is lower than that of the smaller species”) described by Botka and Mészáros (Reference Botka and Mészáros2014, p. 91) were not observed in the Asian material. Seven specimens with a preserved m3 from BRB/C (ZIN 104894), SHM/A (ZIN 104908/1221–1224), ORH/1A (ZIN 104910), and ANR/1C (ZIN 104918/1256) include a rounded posterior margin of the m3 talonid basin (state of Beremendia minor according to Botka and Mészáros, Reference Botka and Mészáros2014) and an entoconid varying in height depending on the degree of wear. However, Beremendia minor from the Asian material includes a longer entocristid of m3, which forms a complete lingual wall around the trigonid basin; in contrast, Beremendia fissidens from Asia includes a slightly shorter entocristid of m3 and a small gap between its end and the base of the metaconid (not forming a full lingual wall around the basin; Fig. 6.11). However, these aspects of the m3 trigonid shape require additional investigation with more material. In addition, the mandibular ramus of Beremendia minor collected from BRB/C (ZIN 104894) includes a relatively longer coronoid spicule and another configuration of facets for the temporal muscle (M. temporalis, pars ventralis; Cornette et al., Reference Cornette, Tresset and Herrel2015), with an anterior rather than posterior facet; the latter is shifted heavily toward the posterior margin of the coronoid process (Fig. 7.5, 7.6, compare ‘b’ and ‘c’). The posterolingual basin of p4 of Beremendia minor is not shifted lingually unlike that observed for Beremendia fissidens (Fig. 7.10, 7.11). However, these results are strictly based on observations of a single Mongolian specimen, and European Beremendia minor can exhibit other features along the lateral surface of the mandibular ramus.

The examined specimens of Beremendia fissidens are smaller than Beremendia jiangnanensis (S6); the former species possess includes more complex anterior parts of the upper antemolars (Beremendia jiangnanensis includes “no parastyle on A1 and A2” according to Jin et al., Reference Jin, Zhang, Sun and Zheng2009, p. 159).

Beremendia fissidens from Asian localities is slightly smaller than the holotype of Beremendia pliocaenica (Fig. 8, ‘Ppl’; S11). Qualitative features of the mandible/lower teeth for the differentiation of Beremendia fissidens and Beremendia pliocaenica are not known.

Remarks

The examined Asian specimens of Beremendia fissidens exhibit variability in morphological features (e.g., morphotypes of the first lower incisor, fourth upper premolar, and first upper molar). Because all of these features can be found in different parts of the geographic range, we assume the intraspecific nature of such variability. Species diagnosis based on new morphological features is possible only after conducting a comprehensive study of additional Asian materials.

Description

Beremendia cf. B. fissidens includes a single isolated first upper molar from ORH/1B (ZIN104911/1244). This tooth is generally comparable in size and shares features with the M1 of Beremendia fissidens from Asian material. However, this tooth also presents some significantly different features, e.g., (1) an entocingulum developed around the base of the protoconid (Fig. 5.7) and (2) a weakly developed metaloph forming a concave, ‘low’ ridge (Fig. 5.8, 5.9).

Remarks

Given that the weakly developed metaloph of M1 is likely typical of the M1 of Peisorex pohaiensis (see Jin and Kawamura, Reference Jin and Kawamura1996a, fig. 3B) and because such metaloph conditions were not observed in other available materials of Beremendia fissidens, we excluded this specimen from Beremendia fissidens.

Holotype

ZZSiD MF/1513, the fragment of right mandibular ramus with p4–m3 in situ and partly damaged mandibular ramus without an angular process; Rębielice Królewskie I, Poland; late Pliocene, MN 16 (Rzebik-Kowalska, Reference Rzebik-Kowalska1976).

Diagnosis

See Rzebik-Kowalska (Reference Rzebik-Kowalska1976), Reumer (Reference Reumer1984), and ‘Emended diagnosis’ by Botka and Mészáros (Reference Botka and Mészáros2014, p. 92).

Description

The rare material investigated, represented by a complete hemimandible and two dentary remains (see S2, S3), can be referred to as Beremendia minor given its small size (Fig. 8; S13, S14).

Dentition

The first lower incisor is straight without denticles on the cutting edge; in general, its morphology corresponds with that of the lower incisor of Beremendia fissidens. The lingual side includes a moderately developed groove on the AAF (Morphotype II, see Fig. 3.8). Ectocingulids are absent.

The first lower antemolar is small and relatively long: its main cusp (protoconid) is relatively longer than that found in Beremendia fissidens, and so the crown of a1 appears to be more compact. The tooth includes a visible posterolingual basin and broad cingulids. The fourth lower premolar is soricine-like in outline. The posterolingual basin is not tilted or shifted lingually. Cingulids are developed (Fig. 7.9–7.11).

The lower molars differ in size. The dimensions of the lower molars are as follows: m1 lengths range from 1.99–2.37 mm; m2 lengths range from 1.71–2.06 mm. The length of m2 is ~85–86% that of m1. To compare, the relative lengths of m1 and m2 of Beremendia fissidens range from 80–86% (calculated from means, see S8, S9); individual relative values of the investigated Asian specimens range from 83–90% (calculated from values for S14). The trigonid of m1 is ~68% the total length of the tooth, and the trigonid of m2 is 66–68% the total length of the tooth. Thus, m2 is shorter than m1 due to the length of the talonid (at least for the ZIN 104894 specimen). The m1 and m2 differ slightly in width (TRW, TAW; see S13). The third lower molar includes a developed trigonid and a smaller talonid. The longer entocristid of m3 forms the lingual wall of the trigonid basin; the entocristid is slightly longer than the ectocingulid (Fig. 6.9, 6.10). All major tips of the lower teeth are pigmented (S5, fig. 1) except in the case of the talonid cusps and cristids of m3.

Mandible

The mandibular body includes a well-developed spicule along the lower margin and facets for attachment of the digastric muscle (as in Beremendia fissidens; Fig. 7.8). The mental foramen lies anterior to the hypoconid of m1. The mandibular ramus includes a relatively straight anterior margin of the coronoid process. The coronoid process includes a long coronoid spicule; the posterior facet for the attachment of the temporal muscle is narrow and shorter than the anterior facet. The mandibular foramen is large and round. The superior pterygoid fossa within the medial base of the condylar process is broad and delineated; its size is comparable to that of the internal fossa. The superior pterygoid spicule is developed as a sharp, cone-like bulge. The short, massive angular process is fully preserved; its tip is bent upward and medially; both sides bear well-delineated facets: an inferior pterygoid fossa on the medial side is formed as a long, moderately deep depression (M. pterygoideus medialis insertion sensu Dötsch, Reference Dötsch1986) and an ‘inferior masseteric facet’ (M. masseter insertion sensu Dötsch, Reference Dötsch1986).

Variation

Teeth from ANR/1C display PPt. The hemimandible of BRB/C displays reddish VPg. EDX analyses reveal the presence of iron at varying levels of 1.15–25.92% (S5, fig. 1, table 1). Dimensions of the lower molars vary moderately but range within the sizes of m2 of Beremendia minor (see Fig. 8). The ventral margins of the dentaries of BRB/C reveal a well-developed spicule; the ANR/1C specimen (ZIN 104918/1255) with a partly preserved ventral margin lacks a bulging spicule.

Comparisons

Beremendia minor is smaller in size than other species of the genus (see Beremendia fissidens above).

Remarks

In general, the dental and mandibular morphology of the material collected from BRB/C is typical of Beremendia. Features that complicate definitive species identification include: (1) an m3 talonid morphology that does not correspond with features found in European fossils described by Botka and Mészáros (Reference Botka and Mészáros2014), and (2) a strong tendency toward ‘sculpturization’ within the mandibular body (i.e., development of a ventral margin spicule) and mandibular ramus (i.e., development of a coronoid spicule). The paucity of Asian fossils available does not allow us to confirm species diagnoses.