Protadelomys maximini ( Escarguel, 1998 )
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publication ID |
https://doi.org/10.1186/s13358-022-00245-3 |
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persistent identifier |
https://treatment.plazi.org/id/497F1B1D-FFF0-DF6E-C739-4181FA37FDF9 |
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treatment provided by |
Felipe |
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scientific name |
Protadelomys maximini ( Escarguel, 1998 ) |
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? Protadelomys maximini ( Escarguel, 1998)
Holotype. M2 (or M1?); SMX1-28 .
Type locality. Saint-Maximin “Grand Chantier” = SMX 1 (Gard, France).
Other localities. Loci SMX 2 (surface sampling) and SMX 3 (Grand chantier, gallery entrance, alveolus), from Saint-Maximin (Gard); middle Eocene, MP 13/14.
Original diagnosis. Escarguel, 1998: 372–373; translated from French. “ Size similar to P. alsaticus . Low crowned cheek teeth, with low ridges and cusps. Sinus weak or absent on upper molars. Small hypocone; reduced conules, especially the paraconule; small mesostyle; mesoloph much reduced or absent; reduced protoloph and metaloph, especially at the level of the connection with the lingual cusps. Lower molars with: reduced anterolophid occupying a buccal position; very weak metalophid and hypolophid, often separated from buccal cuspids; mesoconid and hypoconulid very underdeveloped, even absent; posterolophid relatively little reduced ”.
Emended diagnosis A species of? Protadelomys , about the same size as? P. alsaticus , but with slightly longer dp4, wider DP4 and significantly larger M1; larger than? P. nievesae .
Enamel of the teeth generally ornamented with weak and low extra-ridges and granules.
Upper teeth. DP4 with anteroloph occupying half-width of the mesial edge and protoloph attached to the junction anteroloph-mesial end of the preprotocrista; Anteroflexus generally wide, protruding and short buccolingually. Protoloph high and continuous, with a small paraconule either distinct or indistinct from the anterostyle. Small hypocone, rarely lingual to the protocone. P4 as length as and slightly wider than the DP4. On molars, complex mesostyle area; anterostyle, pre- and post- protocristae, protocone, thick endoloph and small hypocone aligned mesiodistally; mesoloph short when present; para- and meta-conule often crestiform; anterostyle as strong as the hypocone, symmetrical with respect of the protocone; one or two lingual metalophules attached to the hypocone; m3 with metacone slightly prominent; paraconule protruding and metaconule crestiform.
Lower teeth. All the lophids and ridges are thin. On dp4, anteroconid absent, and anterocingulid variably present; protoconid small, and lower than the metaconid; no strong mesiodistal ridge descending in the mesoflexid. On p4, weak, nearly indistinct protoconid. On molars, the metaconid is the higher cuspid, with a long postmetacristid, often joining a preentocristid; the metalophulid I is generally complete; the mesoconid is small and slightly swollen, often bearing mesolophid or ectomesolophid, Te entolophid is always very low, and often discontinuous and incomplete; it is rarely connected to the postmesoconid or to the prehypocristid. Hypoconulid weak. Floor of the mesoflexid relatively flat.
? P. maximini differs from:
-? P. alsaticus in showing: less bulbous main cusps; thinner lophs, lophids and ridges, generally more numerous but weaker and lower extra-ridges and granules; less rough and less wrinkled crown outskirts; relatively longer (mesiodistally) and flatter mesoflexus and mesoflexids; slightly more mesial protoconid on dp4; protoconid absent or indistinct from the metaconid on p4; reduced hypoconulid; upper molars with longer postparacrista and premetacrista, longer and higher endoloph, smaller hypocone, weaker paraconule, metaconule and mesostyle.
-? P. nievesae in showing: the DP4 with a more mesiodistally stretched protocone and hypocone, a longer pre- and post- protocristae and hypocristae, a more reduced hypocone, and lower crown; the upper molars with less bulged cusps and conules that are distinct from the lophs, a higher endoloph, and the sinus nearly absent; the dp4 with a flat basin; the p4 with an underdeveloped protoconid; the lower molars with a more discontinuous and low entolophid.
Material and measurements. (Additional file 6: S6; Tables 6, 8; Figs. 43 View Fig , 44 View Fig ).
Upper teeth: Probably due to different taphonomic conditions, the DP4 from Saint-Maximin are numerous (14), while they are rare in Bouxwiller. If their length variation (1.30–1.58) is close to that observed in Bouxwiller (1.30– 1.53), where several species are distinguished; their width (1.30 to 1.58) is somewhat larger (1.28–1.45). Te M 1 varies in size from 1.51 × 1.62 mm, to 1.94 × 1.88 mm, and the M2 from 1.63 × 1. 65 mm to 1.92, to 2.03 mm (i.e., the type) ( Table 6A, Additional file 6: S6). Only two M3 ( SMX1-291 and SMX2-226 ) are well preserved, and a few are broken, probably due to the overrepresentation of juveniles in the Saint-Maximin locality ( Escarguel, 1998, Fig. 5–11 View Fig View Fig View Fig View Fig View Fig View Fig View Fig , 12 View Fig ) .
Lower teeth: Fourteen dp4 are present in the sample from Saint-Maximin. Tey appear slightly larger than the dp4 of? P. alsaticus from Bouxwiller ( Tables 4A, 6A). Te size of the p4 of? P. maximini is only slightly smaller than that of? P. alsaticus from Bouxwiller ( Tables 4A and 6A). It is larger than that of? P. nievesae from Casa Ramon (Additional file 7: S7, Table 7). Tere are 16 m 1, 12 m 2 and 8 m 3, and their size is not much different from that of? P. alsaticus ( Fig. 44 View Fig ), but they are significantly larger than those of? P. nievesae from Casa Ramon ( Table 8).
Description
Upper teeth
DP4. ( Fig. 45a View Fig to i). Probably due to different taphonomic conditions, the DP4 from Saint-Maximin are numerous (14), while they are rare in Bouxwiller. Like for? P. alsaticus from Bouxwiller, their features display a strong variation, which we consider as intraspecific (see below, morphotypes).
Te largest DP4 are morphologically homogeneous, with an anteroloph occupying half-width of the mesial edge and a protoloph that is attached to the junction anteroloph-mesial end of the preprotocrista (= endoloph in Escarguel, 1998). Te anteroflexus is relatively wide, protruding, and short buccolingually. One DP4 (SMX1- 253: Fig. 45a View Fig ) has a discontinuous anteroloph, with a short stretched parastyle, separated from the anterostyle, which is prolonged in a short lingual ridge; its anteroflexus is clearly narrower than that of other DP4. Te postparacrista of SMX1-253 is high and ends in a swelling, which is reminiscent of a premesostyle, followed buccodistally by a strong mesostyle. Te short premetacrista is separated from the latter by a shallow valley. Te protoloph is high and continuous; its lingual end turns mesially to join the anteroloph at the level of the anterostyle (= protostyle in Escarguel, 1998), which is indistinct from the paraconule. A tiny paraconule is sometimes distinct on the protoloph (SMX1-254: Fig. 45b View Fig , 259, 260: Fig. 45 g View Fig , 264; SMX2-216: Fig. 46 h View Fig , 217: Fig. 45j View Fig ). On the unworn SMX1-255: Fig. 45c View Fig and weakly worn SMX1-256: Fig. 45d View Fig , the paraconule is strong and indistinct from the anterostyle. Te protoloph is discontinuous on SMX1- 255. On the latter specimen, like on SMX1-260, the hypocone is not displaced lingually to the protocone as in all other specimens; therefore, the arms of these two lingual cusps are more mesiodistal than oblique. On SMX1- 258 and 259, a low connection of the protoloph with the preprotocrista is present more distally than in the specimens described above. Te two arms of the protocone are here aligned obliquely, and the conical hypocone does not display any arm.
A postparacrista, more (SMX1-256, 258; SMX2-216) or less strong, is usually present, while a strong premetacrista is always present. Between the two cristae, two adjacent styles are often present: a premesostyle smaller and lower than the mesostyle, the latter being more buccal and often linked to the premetacrista. A very short or medium-length buccal mesoloph is also present; it can be sometimes duplicated (SMX1-253, 254, 260). Te buccal part of the metaloph (metalophule II) runs parallel to the protoloph, and ends at the level of a strong metaconule (metaconule 2). Te latter can be bulbous (SMX1-253, 254, 258, 259, 260, 264), or more or less stretched in its mesiodistal ridges. Tere is no clear metalophule I, but a distomesial ridge, which can be short (SMX1-256, 259, 260) or vestigial. Te metaconule generally separates from the lingual cusps, except on SMX1-253, 258, 259, and 264 where it is linked to the mid-hypocone via a low ridge. Te posteroloph is low and variably long, from the base of the hypocone to the base of the metacone. Faint wrinkles are visible on weakly worn teeth along the anteroloph and/or the posteroloph, and rare extra-ridges or granules are present in the central basin. One of the largest DP4 (SMX1-264: Fig. 45k View Fig ) shows a long low ridge from the mid-protocone (= protocrista), directed towards the buccal mesoloph; on this specimen, the extra-ridges are more numerous. Similar observations can be made on SMX1-260: Fig. 45g View Fig , which is smaller. Te SMX1-252 is a pristine DP4 reduced to an enamel cap with no dentine and roots; it is particularly small. On this tooth, the unworn lophids and extra-ridges are low and discontinuous. Te paraconule is bulged, whereas, the metaconule is ridge-like. Te anteroloph is long, ending in an anterostyle, which connects the preprotocrista. Te protocone and the smaller hypocone are lingually aligned; they are linked through a well-distinct endoloph. Te posteroloph is very short and well separated from the hypocone. A buccal premesostyle weakly separates from the postparacrista, and connects to the more buccal mesostyle. Tis ridge is symmetrical to the premetacrista, on each side of the mesostyle. A well-distinct short mesoloph descends in the mesoflexus from the mesostyle. A few slender and short extra-ridges occupy the mesoflexus.
P4. ( Fig. 45j View Fig to n). Only five upper premolars are present in the sample, one being far stronger and more quadrate than the three others and showing more robust roots (SMX2-218: Fig. 45j View Fig ). Tis tooth is worn and the anteroloph cannot be distinguished. Te paracone prolongs in a short thick protoloph, which is linked mesially to a very short anteroloph. Te latter is hardly distinguishable and linked to the thick preprotocrista. A low central conule is connected to the distal flank of the protoloph. A thick short postparacrista is visible. A small and shallow valley separates the distal extremity of this postparacrista from the mesostyle, which is closer to the premetacrista mesial extremity. A short buccal mesoloph prolongs the mesostyle lingually. Te premetacrista is thick. Te metaloph is buccal and as short as the protoloph. Its posterior part (metalophule II) links to an almost indistinct posterolophid, which is fused to the buccal extremity of the post-hypocristid. Te hypocone is present, even if shorter than the protocone. A short metalophule I is directed from the buccal metaloph to the metaconule. Te latter is swollen, strong, isolated and slightly mesial when compared to the metaloph. Low ridges and granules are present in the central basin.
Te three others P4 differ from SMX 2-218 in their smaller size and slender roots. Teir shape is more trapezoidal (SMX1-262: Fig. 45l View Fig , and 263: Fig. 45m View Fig ) or rounded (SMX1-265: Fig. 45n View Fig ), with a very small to absent hypocone, longer (although short) anteroloph and posteroloph (except SMX1-263 where the anteroloph is as reduced as on SMX2-218). A strong conule widens the mesial end of the preprotocrista on SMX1-262 and could correspond to a mesially displaced paraconule. A weak and low connection is present between this conule and the protoloph–anteroloph. Te metaconule is absent on SMX1-263 and present on SMX1-262 and 265. On the latter, a narrow and shallow break separates the mesostyle from the postmetacrista.
M1–2.
Description of the type SMX1-281. ( Fig. 46a View Fig ).
Te type is the largest upper molar of the sample (1. 91 × 2. 02 mm; Figure 5–8 View Fig View Fig View Fig View Fig in Escarguel, 1998). It could represent an M2, as the hypocone is slightly less lingual than the protocone, and the metacone less buccal than the paracone. Te anteroloph is long, from the stretched parastyle to its junction with the preprotocrista. Te paracone is conical and bears a mesiodistal postparacrista. Te buccal part of the protoloph is relatively high and transverse buccolingually; it is sinuous at the level of the protruding paraconule, and lowers lingually where it connects to the protocone apex. Te mesostyle is duplicated and more buccal than the paracone and metacone; it separates from the premetacrista and prolongs lingually into a short buccal mesoloph. Te metacone is arched together with its curved premetacrista and the short metalophule II. Te extremity of the latter links with a short and low distomesial ridge, which can represent a kind of metalophule I. It separates from the metaconule by a narrow furrow. Te metaconule is equally strong as the paraconule and lowly connects to the hypocone, through a lingual metalophule. Te protocone is only slightly protruding above its pre- and post-protocristae. Te endoloph is long and thick, in line with the postprotocrista. Te hypocone is clearly smaller than the protocone, and do not display distinct arms. Te posteroloph, short, starts by a swelling, probably a posteroconule, and ends at the base of the metacone flank. Some extra-ridges are visible in the mesial and central flexi, and along the posterior slope of the protoloph. Te outer surface of enamel is rough.
(See figure on next page.)
Fig. 42 View Fig Teeth of? Protadelomys alsaticus from Cuzal (Quercy). a CUZ183, left DP4; occlusal view. b CUZ176,left DP4; occlusal view. c: CUZ177, left
P4; occlusal view. d CUZ188,right P4–M1; occlusal view. e CUZ189, left P4–M1; occlusal view. f CUZ155, left M1; occlusal view. g CUZ156, right
M1; occlusal view. h CUZ182, left M1; occlusal view. i CUZ178, right M1; occlusal view.: CUZ155, left M1; occlusal view. g, right M1; occlusal view.
h CUZ182, left M1; occlusal view. i CUZ178, right M1; occlusal view. j: CUZ186, left M1; occlusal view. k: CUZ154,right M2; occlusal view. l: CUZ179, right M2; occlusal view. m: CUZ180, right M3; occlusal view. n: CUZ187,left dp4; n1, occlusal view; n2, buccal aspect. o: CUZ161, right dp4; occlusal view. p: CUZ152, left p4; p1, occlusal view; p2, buccal aspect. q: CUZ164, right m1–m2–m3; occlusal view. r: CUZ168,right m1; occlusal view. s: CUZ163, left m1; occlusal view. t: CUZ153, left m1; occlusal view. u: CUZ167, right m1; occlusal view. v: CUZ172,right m2; occlusal view. w: CUZ169, left m2; occlusal view. X: CUZ173, left m3; occlusal view. Scale bar= 1 mm
Other M1–M2. ( Fig. 46b View Fig to o).
Te molars (SMX1- 269, 270, 271, 272, 275, 278, 279, 282, 284, 285, 286, 288; SMX2- 220, 221, 222, 223, 224) have a well-ornamented enamel, as well as well-defined lophs, which are sharp and often thin. Te posterior part of the M2 is shorter than the anterior part, the hypocone and metacone being closer than on M1.
On these molars, the antero- and mesoflexi are wide, filled with low granules and/or ridges. Te paraconule and metaconule are more often crestiform than bulged. Te parastyle is stretched. Te anteroloph is long and wrinkled distally. Te protoloph is thin and angles at the level of the paraconule, which is protruding, and then connects to the preprotocrista or to the anterostyle. Tis anterostyle, as seen on weakly or unworn molars, is nearly as strong as the hypocone. Te paraconule can connect mesially to the anteroloph via a small ridge (SMX2-223). Te anterostyle, the protocone, the pre- and post-protocristae, the endoloph, and the hypocone are mesiodistally aligned, the anterostyle and hypocone being symmetrical with respect to the protocone. Te mesostyle area is relatively complex: a mesostyle is present; it is more or less shifted buccally, bulged or stretched mesiodistally, and framed by swellings of the distal extremity of the thin postparacrista and the mesial extremity of the strong premetacrista. Te three swellings can prolong lingually into low and short ridges; the medial swelling can even do so as a lingual mesoloph. Premetacrista + metacone + metaloph make an arcuate loph (thin on unworn or weakly worn teeth), which is usually separated from the metaconule area. Tis area is complex and can be filled with several conules and ridges (e.g., on SMX1-275: Fig. 46f View Fig 3 View Fig ). One or two (e.g., SMX1- 275) lingual metalophules attach to the hypocone. Te endoloph is thick and high. On SMX1-278: Fig. 46h, a View Fig low entostyle is located below the endoloph. Te posteroloph is short and rarely reaches the distolingual corner of the metacone (SMX1-271: Fig. 46e View Fig ).
Some M1 (SMX1-273) and M2 (SMX1-276, 280), similar in size, share features, such as the extra-ridges in the synclines, with the teeth described above. However, they also display some morphological variations. Te M1 show a more swollen parastyle, anterostyle, and posteroconule, as well as thicker lophs, especially the protoloph. As such, the synclines appear narrower and the granules and ridges are less numerous. Tis arrangement occurs in the M2 SMX1-276: Fig. 46o View Fig , which however displays numerous extra-ridges. Te M2 SMX1-280, ( Fig. 5–5 View Fig , in Escarguel, 1998: Fig. 46m View Fig ) is slightly different, with its longer and transverse buccal metaloph, which is parallel to the protoloph, its bulged paraconule and metaconule, and its mesocone developed on the endoloph.
M3. ( Fig. 46p View Fig ). Te metacone is present but little prominent from the arcuate cingulum encompassing the premetacrista+ posteroloph + low hypocone + protocone + pre- and post-protocristae + endoloph, to the anteroloph. Te only break on this “pericingulum” is located between the parastyle and the paracone. Te paraconule is protruding and the metaconule is crestiform. Low granules and extra-ridges fill the basin.
LoWer teeth
dp4. ( Fig. 47a View Fig to g). Teir morphology is relatively homogeneous, but a few features are only present in the smallest dp4, which are here attributed to? Protadelomys cf. maximini morphotype 2.
Te two roots diverge, as usual on dp4. Te protoconid is present, small, and lower than the metaconid. A mesial notch separates the two cusps mesially (except on SMX1- 203), but a short metalophulid I can be present. Te latter is absent on SMX1-201 and SMX2-205, and interrupted on SMX1-206. A short anterocingulid descends from the protoconid on some teeth (SMX2-204, 205: Fig. 47e, f View Fig ). On all the dp4s, the postmetacristid is high and sharp. No strong mesiodistal ridge descends from the metalophulid I to the center of the basin; only one isolated short ridge is visible (SMX1-203, 206: Fig. 47b, d View Fig ). Te long postprotocristid is nearly mesiodistal, aligned with the lower ectolophid; the latter bears a small salient mesoconid. On some teeth, the hypoconid displays a spur at its mesial base ( Fig. 47b View Fig 1 View Fig , d 2 View Fig ), which is linked to a short ectocingulid that closes the base of the sinusid (SMX1-203, 206). Tis cingulid can be simply a flat and not a ridge (SMX1-201; SMX2-205: Fig. 47a, f View Fig ). Te short prehypocristid descends towards the ectolophid. Te entoconid is small and conical. When present, the entolophid is low, complete (SMX1-203; SMX2-204) or incomplete (SMX1- 202, 204, 206; SMX2-205; SMX3-26). It is connected, or directed, to the junction prehypocristid – ectolophid, or to the hypoconid (SMX1-205). Te posterolophid is high, reaching the distal slope of the entoconid. It bears a weakly salient hypoconulid. In the basin, the granules and small extraridges are rare (SMX1-202, 203; SMX2- 202, 204, 205) to numerous (SMX1-206: Fig. 47d View Fig ).
p 4. ( Fig. 47h View Fig to k). Tere are seven p4 from Saint-Maximin, only four of them can be described in details, the others being too much worn or damaged. On p4, the main cusps are more bulbous and robust than on dp4. Te protoconid is weak, nearly indistinct of a protocristid mesiodistal ridge and lower than the metaconid, which is more mesially placed. Te protoconid is slightly more distinct on one p4 only (SMX1-208). Te metaconid is not much higher than the hypoconid and entoconid. Te posprotocristid is oblique; it joins the very short mesiodistal ectolophid, which bears a swollen mesoconid. A mesiodistal ridge descends from the metaconid, which is only strong on SMX1-208. Te thick and short postmetacristid ends at a notch, which is the lingual opening of the central basin. Te entolophid is discontinuous and low. Extra-ridges are visible in the basin of SMX1-208. A spur on the protoconid distal flank prolongs in an ectostylid on SMX1-210. A short ectomesolophid is noticeable on SMX1-208 and 210.
m1–m2. ( Fig. 48a View Fig to g). Like in Bouxwiller, we observed a large morphological variation among the 34 m 1–m2 from Saint-Maximin, which it is worth comparing to the size variation. For both m1 and m2, the morphological variation is not as important as in? P. alsaticus , but they are half less numerous than in Bouxwiller. Two molars are more “bulbous” and are described as? Protadelomys cf. maximini morphotype 1. Te m2 SMX1-400 is by far the smallest, and is attributed here to? Protadelomys cf maximini morphotype 2, alongside four small other teeth. Terefore, 27 lower molars (m1–m2) are considered as typical? P. maximini .
(See figure on next page.)
Fig. 46 View Fig Upper molars of? Protadelomys maximini Escarguel, 1998 from Saint-Maximin (Gard). a SMX 1-281, left M1, Holotype; a1, occlusal view; a2, lingual aspect; a3, buccal aspect. b SMX 1-269, left M1 or 2; b1, occlusal view; b2, buccal aspect. c: SMX 1-287, left M1, enamel cover only; occlusal view. d SMX 1-273, right M1; d1, occlusal view; d2, buccal aspect; d3, lingual aspect. e SMX 1-271, left M1; e1, occlusal view; e2, lingual aspect;e3, buccal aspect. f SMX 1-275, right M1; f1, lingual aspect; f2,, buccal aspect;f3; occlusal view. g SMX 2-224, right M1; g1, occlusal view; g2, lingual aspect; g3: buccal aspect. h SMX 1-278,right M1; h1, occlusal view; h2, lingual aspect; h3: buccal aspect. i SMX 2-221, left M1; i1, occlusal view; i2, lingual aspect; i3, buccal aspect. j: SMX 2-222, right M1 or M2; j1, occlusal view; j2, lingual aspect; j3, buccal aspect. k: SMX 2-220, left M1; k1, lingual aspect; k2, occlusal view; k3, buccal aspect. l: SMX 1-223, left M2; l1, occlusal view;l2, lingual aspect; l3, buccal aspect. m: SMX 1-280, left M2; m1, occlusal view; m2, lingual aspect; m3: buccal aspect. n: SMX 1-268, right M2; occlusal view. o: SMX 1-276, left M2; o1, occlusal view; o2, lingual aspect. p: SMX 1-291, left M3; p1, occlusal view; p2, buccal aspect.Scale bar: 1 mm
On all the teeth, the metaconid represents the higher cusp, followed lingually by the high and acute postmetacristid. Te latter is as long as in? P. alsaticus , and longer than in? P. nievesae . Te mesostylid is absent. Te valley between the postmetacristid and the entoconid is generally obtuse; the postmetacristid joins a preentocristid, nearly closing the valley, on SMX1-230. All the lophids and ridges are thin. Te anteroconid (mainly distinct on m1) and the anterolophid are present but are thinner than in? P alsaticus . Te anteroflexid is generally closed distally by a complete (buccolingual) metalophulid I (SMX1-213, 214, 216, 218, 220, 221, 227, 228, 229, 230, 246, 250; SMX2- 212); it is rarely opened distally when there is no connection between the buccal and the lingual halves of the metalophulid I (SMX2-208, 209, 213, 215; SMX1-232, 238 (= 298 in Escarguel, 1998, Fig. 4–9 View Fig View Fig View Fig View Fig View Fig View Fig ), 241, 248). Te anteroflexid slightly opens buccally on weakly worn teeth, but closes through a buccal elevation of enamel of its floor.
Te buccal half of the metalophulid I always displays the same arrangement and buccolingual orientation, starting always from the apex of the protoconid, and running linguobuccally to midwidth of the tooth. As for? P. alsaticus , the main variation is observed at the level of the lingual metalophulid I. Te latter arises completely mesial, and its buccal end slightly turns distobuccally to join the buccal metalophulid I; it can be more distal, descending from the middle of the metaconid (SMX1-216, 218).
Te postprotocristid is strong, thick distally and oblique mesiobuccal to distolingual; it is always longer than the prehypocristid. Te variation observed along the ectolophid area is less large than in? P. alsaticus . Te mesoconid can occupy most of the ectolophid area. On both sides of the mesoconid, the ectolophid is short. Its mesial and distal parts are low and distinct on weakly worn molars. Te path of this ectolophid+ mesoconid varies less than in? P. alsaticus . Te postprotocristid is thick and swollen at its end, this thickening could represent an incipient “premesoconid”, frequently marked by a lingual spur or ridge (e.g., SMX1- 227, 238, SMX2-209, 212). Te shape of the mesoconid only slightly varies with wear. It is generally small and slightly swollen, and can bear a short ridge towards the mesoflexid (mesolophid) or an ectomesolophid (e.g., SMX1- 227, 230; SMX2-209). A postmesoconid spur is present on some specimens (e.g., SMX2-212, SMX1- 233) and can be linked to extra-ridges oriented towards the entolophid. Te entolophid is always very low, and often discontinuous and incomplete; it is rarely connected to the postmesoconid or to the prehypocristid. Te floor of the mesoflexid is relatively flat. Te thick and long post-hypocristid ends with the hypoconulid, which is weakly or not bulged. Te hypoconulid prolongs in a thinner posterolophid, ending at the level of the entoconid, which is small.
Low extra-ridges, wrinkles, and granules are numerous in the anteroflexid, mesoflexid and posteroflexid. A few of these ornamentations organize in very low oblique ridges in the basin; some converge from the metaconid flank to the mesoflexid.
m3. ( Fig. 48h, i View Fig ). All eight m3 are quite larger than the m1–2. Tey show the same features as the other molars.
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