taxonID	type	description	language	source
9865F809426AFFB7FF34FCCFF2C9F8FA.taxon	materials_examined	Holotype: Right mandibular ramus with p 3 - m 1, I. S. 6001, Malusteni, Romania. Original diagnosis: In Radulesco and Samson (1967: 547). Studied material: Cranium; MVL- 46, MVL- 47, MVL- 48, MVL- 49, MVL- 50, MVL- 51, MVL- 52, MVL- 58, MVL- 59, MVL- 62, MVL- 73 a. Mandible; MVL- 55 (dex i 1 - p 3), MVL- 63 (sin i 1 - m 3), MVL- 64 (sin i 1 - m 2), MVL- 65 (dex p 3 - m 2), MVL- 66 (dex i 1 - m 3), MVL- 67 (dex p 3 - m 2), MVL- 68 (sin p 3 - m 3), MVL- 69 (dex i 1 - m 3), MVL- 70 (dex p 3 - m 3), MVL- 71 (dex p 3 - m 3), MVL- 72 (sin i 1 - m 3) and MVL- 73 b (dex i 1 - m 3 and sin i 1 - m 2). Upper dentition / teeth; MVL- 46 (dex I 1 - M 3 and sin I 1 - M 3), MVL- 47 (dex P 2 - M 3 and sin P 2 - M 3), MVL- 48 (dex I 1 - I 2, P 3 - M 3 and sin I 1 - M 3), MVL- 49 (dex I 1 - I 2, P 3 - M 3 and sin I 1 - M 3), MVL- 50 (dex P 2 - M 2 and P 2 - M 3), MVL- 51 (sin P 2 - M 3), MVL- 52 (dex P 2 - M 3 and sin P 2 - M 3), MVL- 53 (dex P 3 - M 2), MVL- 54 (dex I 1 - I 2 and sin I 1 - I 2), MVL- 56 (sin P 3 - M 2), MVL- 57 (dex P 3 - M 2), MVL- 73 a (dex I 1 - M 3 and sin I 1 - I 2, P 3), MVL- 74 (dex I 1 - I 2), MVL- 76 (I 1), MVL- 80 (I 2), MVL- 81 (I 2), MVL- 85 (I 1), MVL- 153 (I 1). Lower dentition / teeth; MVL- 55 (dex i 1 - p 3), MVL- 63 (sin i 1 - m 3), MVL- 64 (sin i 1 - m 2), MVL- 65 (dex p 3 - m 2), MVL- 66 (dex i 1 - m 3), MVL- 67 (dex p 3 - m 2), MVL- 68 (sin p 3 - m 3), MVL- 69 (dex i 1 - m 3), MVL- 70 (dex p 3 - m 3), MVL- 71 (dex p 3 - m 3), MVL- 72 (sin i 1 - m 3) and MVL- 73 b (dex i 1 - m 3 and sin i 1 - m 2), MVL- 82 (i 1), MVL- 83 (lower molariform), MVL- 84 (lower molariform). Description Seven partlv preserved (MVL- 50, MVL- 51, MVL- 52, MVL- 58, MVL- 59, MVL- 62, and MVL- 73 a) (Fig. 2 E, F, I, J) and four fairlv complete (MVL- 46, MVL- 47, MVL- 48, and MVL- 49) (Fig. 3 A – D; Supporting Information, Models S 1, S 2) crania are available along with several maxillarv parts (NISP: 26), altogether indicating a MNI of 13. Except for the damaged posterior parts of the mandibular ramus and coronoid process, most mandibles are well-preserved. Seven specimens (MVL- 55, MVL- 63, MVL- 64, MVL- 66, MVL- 69, MVL- 72, and MVL- 73 b) maintain the bodv (corpus), the diastema, and most of the teeth (Fig. 4 A – F; Supporting Information, Models S 3, S 4), whereas four specimens are more damaged and lack either the diastema or the dentition. Onlv in MVL- 68, some areas of the masseteric fossa are recovered (Fig. 4 G). MVL- 73 b is the onlv specimen with the two hemimandibles intact. A total of 16 mandibles have been preserved, representing at least eight individuals. Cranial measurements are provided in Supporting Information, Table S 1. Skull Nasals: Te complete nasal bones can be observed onlv in MVL- 47 in excellent condition (Fig. 2 D); the imprint of the nasals is preserved in the majoritv of the rest of the specimens, while the posterior portions are preserved in MVL- 50 and in a bad condition in MVL- 52. Te nasals are broadest posteriorlv to their narrowest point at their straight suture with the frontal bones and taper anteriorlv at the anterior nasal aperture. Te nasals form the upper part of the muzzle, which overall is short and wide. Te suture between the nasal and frontal bones forms a forward-pointing ‘ V’ that extends slightlv bevond the zvgomatic shoulders and reaches approximatelv the level of the P 4 – M 1 limit. In MVL- 47, the dorsal sides of the nasals are somewhat convex over most of their anteroposterior length, except near the base of the frontal process, where thev flaten. A thin and verv prominent posterodorsal process of the premaxilla contacts the nasal bones and separates them from the maxilla (Fig. 2 D). A field of neurovascular foramina is located at the lateral border of the dorsal surface of each nasal, arranged along the premaxilla’s nasal process. Apart from MVL- 47, similar fenestrations are visible in the preserved nasals of MVL- 46 and MVL- 50 (Fig. 3 B). Premaxilla: Te paired premaxillae are relativelv long and bear both pairs of upper incisors. Tev create the anterolateral portions of the rostrum and contribute to the anterior part of the palate. In MVL- 46 – MVL- 49, and MVL- 73 a, both premaxillae are almost fullv preserved. Te premaxillae of the remaining specimens have been severelv damaged. Te premaxilla can be split into two parts, the corpus (bodv) and three processes: a nasal process, a palatine process, and a maxillarv process. Both pairs of incisor alveoli are situated on the premaxillarv corpus. Te alveolus for the anterior upper incisor (I 1) is proportionallv larger and located more anteriorlv and dorsallv than the alveolus for the posterior upper incisor (I 2). Troughout the whole anteroposterior length of the premaxillarv corpus, both alveoli are posteriorlv extending and dorsallv curving. Dorsallv located on the premaxillarv corpus are multiple small neurovascular foramina present on its lateral surface. Tese foramina can be seen as a significant fenestration, albeit of a much smaller size than those described above for the nasals. Furthermore, in lateral view, at the premaxilla – maxilla suture of MVL- 46, MVL- 47, and MVL- 73 a, a significantlv larger, narrower, and longer fenestration appears. In ventral view, the length of the premaxilla occupies approximatelv half of the diastema. Te two premaxillae are robust, nonparallel, and form an angle of approximatelv 35 o with the first incisor roots at their apex. In dorsal view, the nasal process of the premaxilla extends posterodorsallv from the corpus. It possesses a rather sharp ventral edge, and it is anteroposteriorlv elongated, dorsallv curved, and sub-triangular in shape. Te nasal process terminates anterior to the apex of the frontal process of the nasals, as seen in the available specimens. Troughout its entire anteroposterior length, the medial surface of the nasal process of the premaxilla is in contact with the lateral process of the premaxilla. Te maxillarv process of the frontal, which divides the premaxilla from the maxilla in dorsal view, is in contact with the ventrolateral side of the nasal process. In ventral view, the palatine process of the premaxilla is rarelv preserved. It can onlv be seen in MVL- 46, MVL- 47, and MVL- 73 a, extending posteriorlv from the anteromedial surface of the premaxillarv corpus. It is slender mediolaterallv and elongated anteroposteriorlv. Te anterior roof of the palatine process is located approximatelv 2 mm behind the alveoli of I 2 in occlusal view. Te posterior area has not been preserved (Fig. 2 C). Maxilla: In most of the specimens, the maxilla is well preserved (Fig. 3 E, F, I). Τhe maxillarv corpus exhibits a roughlv trihedral shape. Te lateral wall of the maxillarv corpus is perforated bv numerous small openings. Tese fenestrations can be observed on the anteroventral and dorsal surfaces of the maxillarv corpus. All specimens show a large, sub-circular infraorbital foramen on the lateral side of the maxillarv corpus (Fig. 2 B). Te anterior-ventral part of the maxillarv corpus has a narrow fissure that runs rostrocaudallv. Tis fissure is surrounded bv a series of long ridges and troughs that run from the front to the back. Tese ridges and vallevs fit tightlv with a series of similar structures on the back and front of the maxillarv process of the premaxilla. Te facial tuberositv is not pronounced. Te palatine process of the maxilla is preserved in all specimens except MVL- 51, which lacks its right side. Te incisive foramen is long, extending posteriorlv to the limit of P 2. Two rounded notches are formed at the posterior end of the incisive foramen, and thev extend posteriorlv to the mid-level of the paired P 2, divided bv a not that sharp mesial palatal spine. Palatine: Several parts of the palatine bone, forming the posterior portion of the hard palate, are preserved, enabling a detailed description of its anatomv. In MVL- 46 – MVL- 48, the palatine is completelv preserved (Figs 2 C, 3 A – F). It is onlv partiallv retained (Fig. 3 J) or abraded in the rest of the specimens. In each hemipalatine, a lateral perpendicular lamina (portion) and a medial horizontal lamina can be distinguished. Te later forms almost the whole surface of the palatine and approximatelv the posterior part of each side of the hard palate. Te palate appears to be relativelv high, forming a deep concavitv between the maxillarv bones. In ventral view, the walls seem to have a slight slope, and the margins seem to narrow from top to botom. Te anterior margin is located at the level of the anterior part of P 4, and the posterior margin bears a verv notable nasal spine, mediallv. Te sub-circular major palatine foramen is located near the maxillo-palatine suture, anterolaterallv. In four palates, the major palatine foramina open within the palatine while in other two thev are placed on the palatine – maxilla suture. Posteriorlv and posteromediallv to the incisive foramen, the palatine surface has several smaller additional foramina. An additional prominent foramen on the level of the anterior portions of M 2 is seen in MVL- 46 (Fig. 3 B). Te presphenoid and vomer are completelv damaged and hence cannot be available for description. Zygomatic and squamosal: In several specimens (e. g. MVL- 47 – MVL- 49) both of the zvgomatic bones are preserved; most of the rest retain onlv the right or the lef part intact. Anteriorlv, the maxilla’s zvgomatic process and the zvgomatic are extensivelv united to form a distinct ‘ V-shaped’ structure that is positioned horizontallv on the bone (Fig. 2 B). Te zvgomatic is the broadest anteriorlv and mediolaterallv, and forms a basket-handle tvpe of arch morphologv. Its dorsoventral height graduallv increases posteriorlv until the middle of the bone and decreases abruptlv posteriorlv to its end. Te zvgomatic has a concavitv in both the lateral and medial sides, with the former creating a prominent zvgomatic fossa. Te internal lateral surface of the middle cranial fossa is formed bv the squamosal, which resides in the lateral region of the braincase. It can be observed in all studied specimens, except MVL- 73 a. It consists of an anterolaterallv protruding zvgomatic process and the squama of the squamosal. Te cranial portion of the craniomandibular joint is tightlv connected to the posterior region of the zvgomatic process. Te zvgomatic process of the squamosal is not as broad as the zvgomatic arch and is positioned above the posterior margin of the zvgoma, providing an essentiallv trihedral morphologv with a conspicuous oblique line extending anteriorlv in lateral view. Lastlv, in most of the specimens, the squamosal and its sutures with the frontal and parietal are traceable. Frontal: MVL- 47 – MVL- 51 exhibit complete frontals; thev are partiallv preserved (MVL- 52), seriouslv damaged (MVL- 46), or even missing (MVL- 73 a) in the rest of the specimens. Te frontal bone can be separated into an orbital portion and the frontal squama, which are divided from each other bv the supraorbital rim (infraorbital margin). Te anterior part of the cranial roof and the posterior part of the dorsal wall of the nasal cavitv are formed bv the frontal squama. Te dorsal surface seems to be relativelv convex on most specimens, vet in MVL- 49 and MVL- 50 it is more flatened, probablv due to post-mortem deformation resulting in slight dorsoventral compaction. Piting is present on the dorsal surface of the frontal squama. Te orbital portion of the frontal bone is concave mediallv and stronglv arched laterallv (Fig. 2 A, B). In dorsal view, the supraorbital process is verv prominent, thorn-like, and extends ventrallv to form part of the orbital wall. Te lacrimal, squamosal, parietal, and posterior surfaces of the frontal process of the maxilla are all abuted bv the orbital section of the frontal anteriorlv. Te lacrimal is positioned on the orbit’s anterior wall. Due to poor preservation, the lacrimal, as well as the orbits’ oblique dorsolateral view, cannot be described. Parietal: Te posterior ceiling of the skull is formed bv the parietal pair, with the supraoccipital and interparietal adding a small basin and a thin strip in the posterior margin, respectivelv, and the squamosal contributing a narrow strip in the lateral and anterior – lateral edges. MVL- 46, MVL- 49, and MVL- 51 have the paired parietal preserved in good condition (Fig. 3 A, C); MVL- 47 and MVL- 50 have onlv the lef parietal intact. Te piting continues and is visible in most specimens, especiallv at the lateral suture with the frontal and squamosal. However, it is less prominent in MVL- 46, MVL- 48, and MVL- 50 (Figs 2 D, 3 A, E). Te braincase seems to be globular, although in some specimens a slight deformation of the cranium can be observed. It is noteworthv that the parietal width of MVL- 48 is significantlv larger than that of the other specimens, due to taphonomv deformation (Fig. 2 A). Te posterior suture between the parietals is raised as a low sagital crest in MVL- 46 and MVL- 51 (Fig. 3 G, H). Te interparietal and supraoccipital regions are damaged in all specimens. Numerous small foramina on the external surfaces of the braincase bones can be observed in all specimens, especiallv in MVL- 47, MVL- 49, and MVL- 51, where piting is verv prominent (Figs 2 D, 3 A, C). Basisphenoid, pterygoid, presphenoid, and alisphenoid: Te presphenoid is connected to the basal sphenoid and is located on the ventral side of the cranium. Te presphenoid bone forms the ventral and anterior boundaries of the optic foramen. Onlv in MVL- 47 and MVL- 48 is the presphenoid partiallv preserved (Fig. 2 C). It is verv thin and ventrodorsallv concave. Positioned anterior to the basioccipital and posterior to the presphenoid, the basisphenoid is situated near the base of the middle cranial fossa. In ventral view, the basisphenoid forms an isosceles triangular shape, as seen in MVL- 47 and MVL- 48 (Fig. 2 C). On the top medial side of the basisphenoid, the small, vet prominent pituitarv foramen can be observed in MVL- 47 and MVL- 48. Anterolaterallv to the basisphenoid, the paired ptervgoids are extending, onlv poorlv preserved in MVL- 48. Te alisphenoids, firmlv united with the basisphenoid ventrallv, extend into the caudal orbital wall and continue to the caudal borders of the palatine bone, where thev form the ptervgoid process, which is damaged in all specimens. Ectotympanic: Most cranial specimens preserve the ectotvmpanic, while MVL- 58 and MVL- 59 are isolated auditorv bullae (Figs 2 A, B, 3 I, F). Te external acoustic meatus has the shape of a tube that leads within the auditorv bulla and is oriented posterolaterallv. Its external opening is large and ovalshaped. Te auditorv bullae are round, inflated, and have a smooth surface. Te front end is more rounded than its posterior counterpart, which is somewhat more pointed. Tev are angled slightlv toward the centre of the skull. Te petrosal, which occupies the posterodorsal region of the skull and is bordered anteriorlv bv the ectotvmpanic, is poorlv preserved in all specimens. Occipital complex (basioccipital, exoccipital, and supraoccipital): At the back of the cranium, the occipital bone forms a bonv ring around the foramen magnum. Because of its general fragilitv, and likelv taphonomic agents, the occipital can onlv be seen to some extent in MVL- 46, MVL- 48, and MVL- 51 (Figs 2 C, 3 I). Te occipital comprises four portions: the basioccipital, exoccipitals, and supraoccipital. Te posterior section of the cranial base is occupied bv the basilar portion (basioccipital). It is onlv observable in MVL- 48, which is in a good state, and partiallv in MVL- 47, which is quite damaged. Anteroposteriorlv, it occurs as a pair of lobes, narrower anteriorlv and slightlv broader posteriorlv, approaching the foramen magnum. Te paired lateral sections (exoccipitals), mainlv seen in MVL- 48, comprise the lateral borders of the foramen magnum and support the main portions of the occipital condvles. Te exoccipital creates a thin paracondvlar process lateral to the dorsal part of the occipital condvle. Tis process continues ventrolaterallv, forming a prominent ‘ V-shaped’ morphologv with the occipital condvles. Te dorsolateral margins of the foramen magnum are formed bv the supraoccipital. Onlv in MVL- 51 are fragments of the supraoccipital visible vet somewhat poorlv preserved; therefore, no anatomical description can be offered. When viewed ventrallv, in MVL- 47, MVL- 48, and MVL- 51, the foramen magnum has a subcircular appearance that is somewhat compressed anteroposteriorlv. MVL- 48 exhibits a somewhat larger foramen magnum, with wider-spaced occipital condvles. Mandible: Te diastema is relativelv long and slender, somewhat longer than the alveolar length of p 3 – m 3 (Fig. 4 A – F). Te alveolus for i 1 is located anteriorlv on the mandibular bodv. Apart from the m 3 alveolus, which is noticeablv smaller, the posterior dorsal side of the mandibular bodv includes alveoli for p 3 – p 4 and m 1 – m 3. Tese alveoli are approximatelv comparable in size. Te dorsal side of the mandibular bodv is somewhat concave and contributes to the ventral half of the diastema, located between the alveoli of the i 1 and p 3. Tere is a mental foramen located between the alveoli approximatelv at the posterior third of the anteroposterior length of the diastema, which is probablv homologous with the mental foramen in leporids located in a similar position (Wible 2007). Te buccal surface of the bodv is richlv fenestrated and, to a lesser degree, the lingual one as well (Fig. 4 A – E). In addition, the lateral surface of the mandibular corpus at the retromolar fossa bears a prominent and round-shaped neurovascular foramen dorsallv. Te lower incisive root protrudes from the lingual side of the bodv and extends posteriorlv below the anterior border of p 3. Lastlv, the ventral side of the mandibular bodv has a noticeable mvlohvoid line for the mvlohvoid muscle insertion. Dentition I 1 and I 2: In total, 13 I 1 (eight in anatomical position and five isolated) and 12 I 2 (10 intact and two isolated) are preserved (Table 1). Te occlusal outline of the upper incisors has an asvmmetrical quadrangular shape that extends anteriorlv in its medial part. Te mesial enamel is relativelv thin with no substantial variation. Te anterior notch is ‘ V-shaped’ and shallow, with wide, open walls and is not filled with cement. It separates the tooth into two parts of approximatelv equal anterior prominence. Te second upper incisors are rounded and mediolaterallv elongated. Encircling the teeth, the enamel is thin. P 2: Twelve P 2 specimens are available (Table 1; Fig. 3 K, L), all of which are still preserved in the maxilla. Te occlusal outline of the teeth is generallv oval. Tev displav three flexus, of which the paraflexus is the deepest, while the hvpoflexus and mesoflexus are shallower or almost absent. When present, the hvpoflexus and mesoflexus are labiallv and mesiobuccallv situated, respectivelv. In all specimens the flexus is filled with cement. Te development of the hvpoflexus ranges from nonexistent (lef side of MVL- 46) to usuallv weak, forming a shallow-narrow notch (MVL- 46 right side, MVL- 47, MVL- 48, MVL- 49, MVL- 51, and MVL- 52) to beter defined and more prominent, forming a less shallow and narrower notch (MVL- 50 lef side, MVL- 73 a). In several specimens (MVL- 46, 50, 73 a) small differences in hvpoflexus development are observed between the right and lef P 2. Te mesoflexus is usuallv shallow and forms a relativelv broad ‘ V-shaped’ notch, weakest in MVL- 49, deeper and wider in MVL- 52 (right side) and MVL- 47, and narrower in MVL- 51, 73 a. Te hvpercone is advanced, achieving the same width development as the lagicone and postcone combined. Te later forms a 45 o angle with the distal hvperloph, whereas the hvpercone has a 25 o to 30 o gradient. Following Fladerer and Reiner (1996) and Čermák (2009), the lagicone is of the B morphotvpe, and the hvpercone is between the IV and V tvpes of enamel structures. Upper molariforms: Te morphologv of the upper molariform teeth (P 3 – M 3) exhibit a folded hvpoflexus crossing more than half of the tooth width and highlv crenulated in most of the specimens, with extensive cement linguallv (Table 1; Fig. 3 K, L). Enamel is ofen denser on the mesial part of the anteroloph and the lingual tips of the posteroloph. Te P 3, P 4, M 1, and M 2 are of comparable size, but the M 3 is considerablv smaller. M 3 lacks a hvpoflexus and is round-shaped. In addition, the separation between the talonid and trigonid is not easilv traceable. Te folded hvpoflexus of M 2 is approximatelv half the width of the tooth crown. Te degree (number and depth) of crenulation decreases from P 3 to M 2, with P 3 and P 4 exhibiting more prominent crenulations. Te P 3 and most of the P 4 trigonids have a narrower width than the talonid. Te situation is reversed at M 1 and M 2, where the trigonid has a larger width than the talonid. In terms of enamel stages, MVL- 49 has the most worn teeth, with nearlv no enamel remaining. MVL- 46 and MVL- 73 a have less damaged enamel than MVL- 49, although thev are much poorer than the remainder of the specimens. Te enamel in the studied material appears to be weaker in the labial sections of the dentition while staving stronger in the buccal parts. Notablv, the wear stage of the enamel differs significantlv between the lef and right maxillae in three specimens. In MVL- 48, MVL- 52, and not as prominentlv in MVL- 50, the lef dentition is more worn than the right. i 1: In total, 10 lower incisors are available from MVL (Table 2). Te i 1 is quadrangular and markedlv prolonged mesiodistallv. Te anterior enamel wall is constantlv thick. In MVL- 64, MVL- 69, and MVL- 72, the occlusal surface is longer mesiodistallv, but not in MVL- 63, MVL- 66, and MVL- 73 b, where the surface is more square-shaped. Te i 1 does not extend to below the p 3. p 3: Among the 12 p 3 (Table 2), onlv one is isolated and most are well-preserved. Te tooth shaf is prismatic with a rhombic occlusal outline. It is quadrangular and prolonged mesiodistallv. Te tooth can be divided into two main parts. Te first is the anterior loph, the trigonid, which is divided into four lobes (conids), and the second is the posterior loph, the talonid, which consists of one lobe prolonged labiallv. Te later is approximatelv one-third of the width of the trigonid. Te two lobes are connected bv a dentine isthmus that is overall quite wide and short. Te development of the anteroflexid (AR) shows variations among the specimens. It can be distinguished in three main morphologies: a shallow and narrow ‘ V-shaped’ AR (MVL- 55, MVL- 64, MVL- 67, MVL- 71, and MVL- 73 b sin) (Fig. 4 J); a narrow ‘ U-shaped’ AR, less shallow than the previous tvpe and quite wider (MVL- 63, MVL- 66, MVL- 69, MVL- 70, and MVL- 73 b dex) (Fig. 4 H, I); and a wide, deep ‘ U-shaped’ AR (MVL- 65) (Fig. 4 K). MVL- 72 has a bilobal anteroconid with dual anteroflexids that are wide and short (Fig. 4 L). Te hvpoflexid (PER) and the mesoflexid (PIR) are the deepest flexids, dividing the tooth into the trigonid and talonid parts, buccallv and linguallv, respectivelv. In most of the specimens, the two flexids are of equal development. However, in MVL- 69, MVL- 72, and MVL- 73 b, it is clear that the PIR is not as deep as PER and shorter. Te PIR is nearlv half the length of PER in MVL- 72 and quite shallow. In MVL- 69 and MVL- 73 b, the PIR is approximatelv two-thirds of PER length. Te angle of the hvpoflexid shows variations in the studied material, exhibiting a slope of about 45 o towards the talonid. Te gradient is more prominent in MVL- 73 b and lower in MVL- 55, MVL- 66, MVL- 69, and MVL- 71. Te protoflexid (AER) also exhibits a varietv of different developments in MVL. It can be distinguished in two main morphotvpes. Te first one shows a shallow and wide ‘ V-shaped’ AER (shown in MVL- 63, MVL- 65, MVL- 70, MVL- 71, and MVL- 73 b dex) and the second a narrow and shallow ‘ L-shaped’ AER (seen in MVL- 55, MVL- 66, MVL- 67, MVL- 69, MVL- 72, and MVL- 73 b sin). Lastlv, the hvpoconid is relativelv massive and rectangular. Te buccal side of the hvpoconid shows a small, rounded notch with verv thin enamel. Te PR 1 / A 1 or Alilepus patern (i. e. no mesofossetid / presence of anteroflexid) is the onlv one recorded (100 %; N = 12). Following Fladerer and Reiner (1996) and Čermák (2009), the hvpoconid is predominantlv of the ‘ c morphotvpe’ (MVL- 63, MVL- 65, MVL- 66, MVL- 69, and MVL- 71), more rarelv of the ‘ a morphotvpe’ (MVL- 55) or ‘ b morphotvpe’ (MVL- 70 and MVL- 72), whereas also an in-between stage of ‘ morphotvpes b – c’ is found (MVL- 66 and MVL- 73 b). Te hvpoflexid is of the ‘ A morphotvpe’ in MVL- 71, the ‘ C morphotvpe’ in MVL- 65, MVL- 66, MVL- 69, and MVL- 73 b, and the ‘ D morphotvpe’ in MVL- 55, MVL- 70, and MVL- 72. An in-between stage of the ‘ C and D morphotvpe’ is shown in MVL- 63. Lower molariforms: Te two cvlindrical shafs united bv a small bridge at the lingual side characterize the morphologv of all teeth, which is tvpical of the great majoritv of leporid species (Table 2; Fig. 4 H). In contrast to the p 3, the rest of the lower molariforms (p 4, m 1, m 2, and m 3) in Lagomorpha are rather consistent and do not displav anv remarkable traits. In comparison with the talonid, the trigonid is larger (both in length and width). Te mesiobuccal corner of the trigonid is small and smooth. Mesiolabiallv, the labial portion of the talonid is convex. Te isthmus joining the trigonid and talonid is lengthv and narrow. Te flexid between the trigonid and talonid is smooth, and the enamel on the labial portion of the trigonid’s distal edge is extremelv thick. Te p 3, p 4, and m 1 are of comparable size; however, the m 2 is considerablv smaller. In addition, the m 3 is cvlindrical-shaped, and the talonid portion is round, as opposed to the cvlindrical shape of the remainder of the lower molariforms. No crenulations are noticeable. Regarding the overall tooth wear, the enamel is more damaged or even completelv absent on the lingual side of the molariform teeth, as in the upper dentition. Te most worn molariforms are those of MVL- 65 and MVL- 67 coupled with a less worn p 3 than the remainder of the teeth. Comparisons Our comparison is focused on the lower third premolar (p 3), which is considered to be one of the most diagnostic anatomical elements on leporid taxonomv. Te second upper premolar also plavs an important taxonomic role but less so than the p 3 (e. g. Radulesco and Samson 1967, Erbajeva and Angermann 1983, Averianov and Tesakov 1997, López Martínez et al. 2007). Te cranial anatomv of leporids, especiallv fossil ones, has not been comprehensivelv documented. Cranial findings are uncommon in the fossil record, and when thev do occur, thev mostlv comprise of fragments of the skull (e. g. Dietrich 1942, Averianov and Tesakov 1997, Asher et al. 2005, Şen and Geraads 2023). Hence, we compare the cranium of the MVL taxon with extant leporid species, as well as with available fossil crania of related leporids. Dental comparison: Te p 3 of Hypolagus Dice, 1917 lacks an anteroflexid and a mesoflexid (A 0 / PR 0 tvpe), thus differing from the studied material (Čermák 2009, Čermák and Wagner 2013). Pliopentalagus Gureev and Konkova, 1964 has a three-folded trigonid p 3 (Alilepus patern) like the MVL sample, but the anterior enamel of its talonid is highlv crenulated (Tomida and Jin 2005) in contrast to the much simpler enamel of our sample. In addition, the enamel of the mesial edge of the talonid on the m 2 is also crenulated in Pliopentalagus in contrast to the material from MVL (Tomida and Jin 2005). While Alilepus possesses a mesoflexid or a mesofossetid on p 3 (A 0 / PR 1 – 2 tvpe) (Čermák and Wagner 2013, Čermák et al. 2015), it lacks an anteroflexid, which contrasts with the MVL morphologv. Different from MVL, the p 3 of Serengetilagus is rounded-shaped (or crescentlike), lacks anterior and posterior lingual flexids (AIR, PIR, respectivelv), the lingual anteroconid is weaker than the buccal one and the anteroflexid is weak to absent. In addition to that, it is reported that the upper molariform teeth from both African species of Serengetilagus show a well-crenulated enamel along the distal wall of the hvpoflexus (López-Martínez et al. 2007). Te morphologv of the folded hvpoflexus in MVL is much simpler and occurs primarlv on the mesial side. Lepus Linnaeus, 1758 and Oryctolagus Lilljeborg, 1874, have the leporine tvpe of p 3 but other than the MVL taxon thev displav a deep PER and no / weaker PIR, together with ofen stronglv crenulated enamel (A 1 / PR 3 tvpe) (Hibbard 1963, López-Martínez 1980, Callou 1997). Te dental features, and especiallv the p 3 morphologv of the studied leporid material from MVL, fit perfectlv those of the genus Trischizolagus with which it shares the following diagnostic features: (i) the presence of a three-folded trigonid in p 3 (Alilepus patern); (ii) the rhombic outline of p 3 with a broad lingual anteroconid and a well-developed AR; and (iii) the simple and uncrenulated enamel patern of the lower and upper check teeth (Averianov and Tesakov 1997, Čermák and Wagner 2013). Čermák and Wagner (2013) provided a comprehensive review of Trischizolagus and recognized six species: T. crusafonti (Janvier and Montenat 1971); T. dumitrescuae; T. gambariani (Melik-Adamvan 1986); T. maritsae de Bruijn et al., 1970; T. mirificus Qiu and Storch, 2000; T. nihewanensis (Cai, 1989); and T. raynali (Geraads 1994). Şen et al. (2024) confirmed with new data the debatable generic atribution of the later species (e. g. Şen and Erbajeva 1995, Averianov and Tesakov 1997, López-Martínez et al. 2007, Čermák and Wagner 2013), while Şen and Geraads (2023) added recentlv one more species, T. meridionalis Şen and Geraads, 2023 from Morocco.	en	Kalaitzi, Christina N., Kostopoulos, Dimitris S. (2025): Craniodental anatomv of Late Ruscinian Trischizolagus (Leporidae: Lagomorpha) from Megalo Emvolon (Tessaloniki, Greece). Zoological Journal of the Linnean Society 204 (2), DOI: 10.1093/zoolinnean/zlaf067, URL: https://doi.org/10.1093/zoolinnean/zlaf067
9865F809426AFFB7FF34FCCFF2C9F8FA.taxon	description	Te p 3 on the holotvpe mandible of T. dumitrescuae (I. S. 6001) from Malusteni, Romania, is characterized bv an A 1 / Pa 1 / PR 1 p 3 morphologv, which is consistent with the MVL specimens (Radulesco and Samson 1967, 1995, Averianov 1995, Čermák and Wagner 2013). Te anteroflexid (AR) of I. S. 6001 shows a comparable degree of development to that observed in the specimens from MVL, as evidenced bv the sketches presented bv Radulesco and Samson (1967) and Crespo et al. (2023). In both the tvpe specimen and MVL (especiallv in MVL- 69, MVL- 72, and MVL- 73), the AR divides the anteroconid into two nearlv equal halves. Te AIR is relativelv narrow and shallow, whereas the AER is wider and deeper in both the MVL and I. S. 6001. Te PER and PIR of I. S. 6001 p 3 are compatible with those from MVL. In addition, the trapezoidal shape of the talonid remains consistent in both the Romanian tvpe specimen and the majoritv of the MVL specimens, albeit some MVL toothrows displav reduced enamel on the buccal side of the hvpoconid and lack of the characteristic notch possiblv due to the wear stage of the enamel. Additional specimens of T. dumitrescuae recovered from Malusteni exhibit a comparable size (Fig. 5) and morphologv of p 3 with both the holotvpe and the MVL sample. Te PR 2 patern has been observed in a single specimen from Malusteni (I. S. 6008) in contrast to the more common PR 1. Based on the measurements provided bv Crespo et al. (2023), the p 3 size range of the species at this Romanian localitv overlaps with the size range recorded for the MVL sample. Regarding the mandible, the length of the diastema is onlv available for one specimen (I. S. 6008); it is measured at 19.5 mm (Radulesco and Samson 1967), exceeding the measured range of 15.56 to 18.01 mm (N = 8) in the MVL material. Te mandibular fenestration is in both the MVL specimens and those from Malusteni prominent on the buccal side of the mandibular corpus at the level of the premolars and in the lower region where the mental foramen is located. Two p 3 specimens (I. S. 6006 and I. S. 6007) of T. dumitrescuae are reported from Berești (Romania). I. S. 6007 is notablv smaller than the MVL specimens, while I. S. 6006 falls within the lower size range of MVL. According to the illustrations provided bv Radulesco and Samson (1967), the morphologv of these two specimens is similar to that of MVL. As noted in the revision of Malusteni and Berești bv Crespo et al. (2023), a significant number of specimens were lost, including the second upper premolar. No distinct size variations were provided for the upper premolars and molars, with onlv a size range for the upper molariforms (P 4, M 1, and M 2) available (Crespo et al. 2023). Nevertheless, the size range of the upper molariforms from MVL generallv aligns with those from both Berești and Malusteni. Averianov and Tesakov (1997) described a rich material of Trischizolagus from Moldova and Ukraine, the vast majoritv of which was atributed to T. dumitrescuae. Unfortunatelv, the available information is poor. As in the MVL leporid, the East European sample shows the rhombic-rectangular p 3 outline, which is accompanied bv a relativelv short and wide anteroconid in all cases. Te PR 1 patern is the predominant, as reported bv Averianov and Tesakov (1997) and Čermák and Wagner (2013). Tese traits are consistent with the MVL leporid. Čermák and Wagner (2013) documented two records of T. dumitrescuae from the Beremend 39 site in Hungarv (MN 15 b). Te first specimen comprises a lower mandible encompassing p 3 to m 2. It displavs the PR 2 patern on p 3, in contrast to the MVL specimens but it exhibits a bilobal AR that is similar to that observed in MVL- 72. In terms of phvsical dimensions, the Hungarian specimen has a large p 3, close to the length measurements of MVL- 69 and MVL- 70 but slightlv narrower. Te remaining molariforms fall within the size range of the MVL specimens, albeit in the higher part of the variation. Te dimensions of both the diastema and the alveolar region are significantlv larger in the Hungarian specimens (21.11 mm in Beremend 39) compared to those from MVL (15.56 – 18.01 mm), while the length of the alveoli region ranges from 15.13 to 16.78 mm in MVL versus 18.2 mm in the Hungarian specimen. Te second Hungarian specimen has been atributed to cf. Trischizolagus dumitrescuae and comprises a single fragment of a lef mandibular bodv bearing p 3 – p 4. Te patern of p 3 corresponds to that of Alilepus as in MVL. Te anteroconid exhibits a broad morphologv, defined bv an undulating enamel patern similar to that of MVL- 65. Te previouslv described specimens of T. dumitrescuae from the Megalo Emvolon- 1 site (MEV) are also being compared with the newlv discovered ones of MVL. Te four available specimens, MEV- 1001 (lower lef and right upper maxillae with P 2 – M 3), MEV- 1002 (lef mandibular fragment with p 4 – m 2), MEV- 1003 (lower lef mandibular fragment of p 3 – m 3), and MEV- 1004 (lower lef incisor), fullv match those described in the present studv on the P 2 and p 3 morphologv and overall size. Apart from the poor preservation, the PIR of MEV- 1003 p 3 seems to be linguallv closed while retaining the appearance of a fold. Although none of the MVL specimens have a closed mesoflexid, in MVL- 70 and MVL- 73 b it is near to shuting. As in MVL- 73 b, the AIR of MEV- 1003 p 3 is quite broad and open. According to Fladerer and Reiner (1996) and Čermák (2009), the hvpoconid of MEV- 1003 is either of an ‘ e’ or ‘ d morphotvpe’, and the hvpoflexid shows an ‘ A’ or ‘ C morphotvpe’. Although none of the MVL specimens show a similar hvpoconid to MEV- 1003, its hvpoflexid is like that in MVL- 65, MVL- 66, MVL- 69, MVL- 71, and MVL- 73 b. Cranial comparison: A cranial comparison with extant leporid taxa is provided in the Supporting Information, Appendix S 1. Alilepus hibbardi White 1991 (IMNH 38695) is known for a complete skull recovered from the Late Miocene of North America (White 1991). It exhibits cranial dimensions close to those of MVL. Nevertheless, the premaxilla is shorter, and the degree of fenestration observed in the maxilla and premaxilla is less advanced than in MVL. White’s (1991) illustrations indicate that the zvgomatic arches exhibit a similar level of inclination as those observed in MVL. Nevertheless, when viewed laterallv, the posterior end of the zvgomatic arches is situated at a higher position in A. hibbardi relative to that of MVL, resulting in comparativelv smaller eve sockets. Te anterior portion of the nasals is wider in A. hibbardi and the supraorbital process of the frontal bone shows a greater development being more rounded and laterallv extended, therebv leading to a posterior flexid that is wider and deeper in comparison to the thorn-like state seen in MVL. Te palate of Alilepus shares a similar relative length to that of MVL (index I 4, Supporting Information, Fig. S 1). On the other hand, compared to MVL, A. hibbardi has narrower incisive foramen (index I 2) and a shorter upper diastema (index I 6, Supporting Information, Fig. S 1). On the mandible, the diastema is shorter in A. hibbardi (index I 10, Supporting Information, Fig. S 1) and more concave than in MVL (White 1991). Hypolagus balearicus Quintana et al., 2010 from the Earlv Pliocene localitv of Ses Fontanelles (Eivissa) (Balearic Islands, Western Mediterranean Sea) (Cardona and Moncunill-Solé 2014), is one of the few Eurasian fossil species known bv its skull. Although poorlv preserved and lacking its dentition, the much smaller cranium of H. balearicus also differs from the MVL one in the narrower palate and choanae, and the more laterallv elongated root of the zvgomatic process of the maxilla (Cardona and Moncunill-Solé 2014). According to the descriptions and illustrations bv Dietrich (1942), the muzzle region of Serengetilagus praecapensis Dietricht, 1941 cranium from the Middle Pliocene of Laetoli, Tanzania, appears to be comparable to that of MVL in dorsal view. Nevertheless, when viewed laterallv, the rostrum of S. praecapensis appears to occupv a significantlv greater surface than in MVL and its muzzle top area is more arched. In addition, the maxillarv fenestration in S. praecapensis seems to be more prominent than in MVL, but limited to the posterior part of the maxilla, once again in contrast to the MVL cranium in which the premaxilla shows prominent piting. Te supraorbital process of S. praecapensis shares the same development as in MVL. Te zvgoma of S. praecapensis is higher positioned compared to MVL and exhibits a more angular lower portion as it shows a noticeable change in slope halfwav along its length, resulting in a steeper angle towards the posterior end. Tis contrasts sharplv with the zvgoma of the MVL cranium, which maintains a consistent slope throughout its entire length, lacking anv mid-arch angular transition. Te incisive foramen of S. praecapensis is wider than that of MVL (index I 2, Supporting Information, Fig. S 1) and the palate is shorter (index I 4, Supporting Information, Fig. S 1). In addition, the upper diastema of S. praecapensis is longer than in MVL (López-Martínez et al. 2007, Şen and Geraads 2023). It must be noted that the ratio of choanae breadth to palatal length for S. praecapensis has not been indicated bv Dietrich (1942), but according to the notes of López-Martínez et al. (2007) it would be larger than 1. Measurements provided from Şen and Geraads (2023) suggest a ratio about 1.3; it is close to 1 in MVL. Lastlv, the lower diastema of S. praecapensis is shorter than in MVL. Few cranial remains of S. tchadensis López-Martínez et al., 2007 have been found from the Late Miocene deposits in the Toros Menalla region of Northern Chad (López-Martínez et al. 2007). From the limited available measurements, S. tchadensis exhibits a longer palate (index I 4, Supporting Information, Fig. S 1) and a shorter upper diastema (index I 6, Supporting Information, Fig. S 1) than MVL. Te breadth of the incisive foramen is similar to that of MVL (index I 2, Supporting Information, Fig. S 1) (Şen and Geraads 2023). According to Averianov (1995), the remains of T. dumitrescuae crania are limited to two small fragments of the frontal and occipital bones from southern Moldova. Regretablv, onlv sketches are accessible, and thus, we base our comparison onlv on the descriptions. Te median portion of the preserved right frontal does not differ from the condition seen in L. europaeus, similarlv to MVL. Averianov (1995) highlights that the foramen magnum displavs a resemblance to that of L. europaeus, wherein it is positioned more superiorlv. Tis description appears to be inconsistent with that of the MVL leporid where the foramen magnum is preserved. Te placement of the foramen magnum in these MVL crania is relativelv more posterior and can onlv be fullv observed from an occipital viewpoint. Averianov (1995) concluded that the auditorv bullae of T. dumitrescuae are presumablv smaller in comparison to extant species and similar in size to that of amami rabbit (Pentalagus). However, the auditorv bullae in the cranial material from MVL are close in size to that of Oryctolagus. Finallv, the mandible of T. dumitrescuae (ZIRS 80457) exhibits a slightlv shorter diastema (index I 10, Supporting Information, Fig. S 1) compared to MVL, whereas the rest of the mandible does not show anv major dissimilarities. Te palatal parts of T. meridionalis were recentlv described bv Şen and Geraads (2023). Like MVL, the posterior end of the incisive foramen produces two rounded grooves that are separated bv a pointed mesial palatal spine and extend posteriorlv to the mid-level of the P 2 s. Te incisive foramen of T. meridionalis is, however, slightlv wider than that of MVL (index I 2, Supporting Information, Fig. S 1). No major palatine foramen is visible in the Moroccan leporid for a comparison with MVL. In T. meridionalis, the palatal length in relation to the breadth of the choanae exhibits significantlv lower values than in MVL. It is also noteworthv that the absence of the posterior spine of the palate in T. meridionalis is a distinguishing characteristic from the MVL specimens and the majoritv of leporids (Şen and Geraads 2023). In addition, the choanae of T. meridionalis is wider than that of MVL and the palate of T. meridionalis is shorter than in MVL (index I 4, Supporting Information, Fig. S 1). Te inter-zvgomatic width in T. meridionalis measures 34.7 mm between the antero-external borders of the zvgomatic arches, falling below the range of the MVL specimens, vet in close proximitv to MVL- 52. Like T. meridionalis, the zvgomatic arches of MVL exhibit a slight posterior divergence. In contrast to the sharp ‘ V-shaped’ structure of the zvgomatic process of the maxilla in MVL, this feature is less developed in T. meridionalis and not as arched and sharp. Lastlv, the available mandibles of T. meridionalis have a slightlv shorter diastema in comparison to that of MVL (index I 10, Supporting Information, Fig. S 1). Finallv, T. raynali from Grote des Rhinocéros exhibits a mandible with a significantlv reduced value for the I 10 index when compared to MVL (Supporting Information, Fig. S 1). Laterallv, the mandibular bodv below the diastema is shallower in T. raynali. Te mental foramen shows similar development in both taxa. Te mandibular bodv of the Moroccan taxon displavs numerous prominent foramina; in MVL, these foramina appear relativelv subtle, expanding in some cases beneath the p 4. Cladistic results: Te cladistic analvsis (Supporting Information, Appendix S 2) provided nine most-parsimonious trees (length: 107; CI: 59; RI: 83) and hence failed to resolve anv relationship apart from the more primitive position of Hypolagus and Alilepus compared to the rest of ingroup taxa, the alreadv known sister-group relationships between North American Syloilagus and Brachylagus (see also: Robinson and Mathee 2005), and the close relationships between Pentalagus and Pliopentalagus (see also: Tomida and Takahasi 2023). Te strict consensus tree is given in the Supporting Information, Figure S 2. As in Averianov (1999), Trischizolagus takes part in a wide polvtomv including extant and extinct Pliocene to recent genera from both the New and the Old World. Although highlv unresolved, in seven out of the nine trees (78 %), Trischizolagus is nested in different basal positions within a group including Serengetilagus, Pronolagus, Pliopentalagus, and Pentalagus ± Romerolagus (TS 2 P ± R group). A deep AIR [character A 15 (1)] on p 3 excludes Romerolagus from the TS 2 P ± R group and supports the monophvlv of the rest. On the other hand, the common presence of PIR [either as a flexid or fossetid; character A 17 Rw (1)] excludes Serengetilagus from this group and links the rest of the genera to the more primitive Alilepus, in distinction to other ingroup taxa. Within the TS 2 P ± R group, the retention of a more primitive P 2 with two re-entrants [character A 9 (1)] discriminates African from non-African taxa.	en	Kalaitzi, Christina N., Kostopoulos, Dimitris S. (2025): Craniodental anatomv of Late Ruscinian Trischizolagus (Leporidae: Lagomorpha) from Megalo Emvolon (Tessaloniki, Greece). Zoological Journal of the Linnean Society 204 (2), DOI: 10.1093/zoolinnean/zlaf067, URL: https://doi.org/10.1093/zoolinnean/zlaf067
