taxonID	type	description	language	source
497F1B1DFFB5DF2FC48346A0FC79F957.taxon	materials_examined	Holotype. Ef 1002: right P 4 – M 3. Type locality. Egerkingen (Swiss Jura), Huppersand?; middle Eocene, MP 14. Original diagnosis. Stehlin & Schaub, 1951 (translation from German). Upper teeth subtriangular to quadrangular; pseudypoconus (= hypocone) still directly connected to the protocone; sinus absent; conules well present; transverse crests of lower molars low or incomplete; no indication of a mesoconid.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFB5DF2FC48346A0FC79F957.taxon	discussion	Remarks about the diagnosis of Protadelomys cartieri. Hartenberger defined the genus Protadelomys for the species cartieri from Egerkingen previously referred to the genus Adelomys (type: right P 4 – M 3; Fig. 25, in Stehlin & Schaub, 1951), then to Masillamys (Taler, 1966). Stehlin and Schaub (1951) provided some dental features in their diagnosis, (e. g., the lack of the mesoconid on lower teeth), which were not taken into account in the definition of the species P. cartieri by the following authors (Escarguel, 1998; Hartenberger, 1969; Peláez-Campomanes, 1995). Emended diagnosis. Species of Protadelomys with slender lower jaw. Upper teeth with buccal protoloph and metaloph slightly lower than the paracone and metacone; buccal metalophule I variably present; buccal metalophule II free or fused to the metaconule; lingual protoloph connecting protocone and lingual metaloph absent or very low and connecting hypocone; short high endoloph; sinus shallow and short linguobuccally; paraconule smaller than the metaconule. On lower p 4, protoconid weak and stretched in the protoconid ridge; on lower molars, metalophulid I and entolophid are low, complete or interrupted buccally; no indication of a mesoconid.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFB5DF2FC48346A0FC79F957.taxon	diagnosis	Differential diagnosis. Protadelomys cartieri differs from P. lugdunensis: — in its relatively slender dentary; on lower teeth, in the presence of a thicker protoconid on p 4, in the absence of mesoconid and anterolophulid, and in its weak antesinusid and anterocingulid; on upper teeth, it differs in its narrower P 4 and smaller M 2.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFB5DF2FC48346A0FC79F957.taxon	materials_examined	Material from Egerkingen and Measurements S 1; Tables 1, 8; Fig. 3 A few lower molars are too worn (Ek 68, 44, 75, 76) or damaged (Ek 73) to be identified with certainty. Te length and width of upper teeth, except that of P 4 (Fig. 3, Table 1) are poorly correlated, whereas these dimensions are better correlated on the lower teeth, except p 4. Tese p 4 are far smaller than m 1, which are smaller than m 2. Te m 3 are longer but not wider than m 1 – m 2. Te M 1 are generally longer than M 2 (not on the type, on which both have same length. Te M 3 are narrower and as long as M 1 – 2.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFB5DF2FC48346A0FC79F957.taxon	description	Description. Figures 4, 5, 6, 7, 8, 9 and 10 Skull. Te specimen Ek 245, described and figured in Hartenberger (1969: 38 – 41) is now badly damaged. Te remaining parts include a fragment with left P 4 and M 1. Te casts of the two dental rows with P 4 to M 3 are still available. A fragment of the left maxillary He 14, bearing M 1 – M 2, shows the base of the hystricomorphous infra-orbitary foramen. Upper teeth Description of the type Ef 1002 (Fig. 4 a). Te P 4 is smaller than the M 1 – 2 (Fig. 3 A; Additional file 1: S 1). It is triangular since the hypocone is nearly absent. A weak parastyle connects to the preparacrista, which swells at the level of an anterostyle. Te parastyle is lingually connected to a distinct anteroloph. Tere is no distinct paraconule. Te paracone is bulged, prolonged in a short protoloph oriented anteriorly to the anteroloph. Te enamel of the mesoflexus is smooth. A low ridge and mesostyle a little higher than this ridge terminates the mesoflexus buccally. Te metacone and the paracone are at the same level. A short buccal part of the metalophule II ends at the base of the metaconule, which is rounded and strong. Tis metaconule connects to the postprotocrista through a well-distinct lingual metalophule. Te buccal metalophule I is absent. Te posteroloph is long and ends against the postprotocrista. Te hypocone is indistinct. Te curved parastyle is longer and stronger on M 1 than on M 2. Te anteroloph ends at the anterostyle. A short antesinus marks the anterostyle at its junction with the preprotocrista, which is more distinct on M 1 than on M 2. Te paracone and metacone are swollen, little prominent, and prolonged lingually in complete transverse buccal protoloph and metalophule II. Paraconule and metaconule are slightly protruding, the paraconule being smaller than the metaconule. Protocone and hypocone are equal on M 1, the hypocone being a little smaller than the protocone on M 2. Te protoloph connects to the protocone. Te metalophule joins the hypocone slightly mesially to its center. On the three molars, a distinct mesostyle extends lingually in a short mesoloph. A short premetacrista is present on M 1 – 2. Te posteroloph is long; it extends from the post-hypocrista to the base of the metacone, without merging with the latter. Te M 3 is badly preserved, with the cusps and lophs less visible than on the M 1 – 2. It is triangular and lacks a distinct hypocone. Other teeth P 4. Te teeth are triangular on the type Ef 1002 and on Ek H 006 (Fig. 6 c), due to the absence of hypocone. Te hypocone can, however, be small [Ek 48, Ek 123, (Fig. 5 a, d) Ek 601 (Fig. 6 b)], more developed [Ek 245 (Fig. 5, b, c), Ek H 005 (Fig. 6 a)], or as robust as the protocone (Ek 35). Te size of the P 4 is independent of the presence / absence of the hypocone. Te anteroloph is long (holotype Ef 1002, and EK H 005, EK H 006, Ek 35, Ek 245), short (Ek 123, EK 601), or absent (EK 48). Paracone and metacone are swollen and similar in size. Te paraconule is generally absent, and can be absent, or indistinguishable from the anterostyle on Ek 245, at the level of the connection with the protoloph. Te paracone is isolated, without buccal protoloph, the protoloph being reduced to its low lingual ridge on Ek H 006 (Fig. 6 c); it is more generally short and directed or connected to the anteroloph at the anterostyle level; it is transverse and joins the anterior arm of the protocone on Ek H 005 (Fig. 6 a). Te mesoflexus is narrow, buccally edged by a more or less swollen mesostyle; the latter being lengthened mesiodistally (Ek 245), or swollen (3 / 8). In Ek 48, Ek H 006, and Ek 123, the mesostyle is connected to a mesoloph, which is short or nearly reaching the metaconule level. On Ek 123, a short postparacrista connects the paracone with the mesostyle. On four teeth (Ek 35, Ek 245 (left and right), Ef 1002), the mesostyle is twofold; it is unique on the others. Te paraconule is absent (Ek 48, Ek 35) or pulled forward, merging with the anterostyle. Te buccal metalophule I is reduced or absent. Te metaconule, swollen, is in line with the metacone through the metalophule II. Te lingual metalophule is oriented towards the posterior part of the protocone, joining it only on two specimens (Holotype and Ek 245). Te P 4 Ek 245 shows a short sinus, a metaconule connected to the hypocone and to the postprotocrista. Furthermore, there is a weak ridge marking the buccal metalophule I, several wrinkles around the metaconule, in the mesoflexus and the posteroflexus. On Ek 35, there is a sinus separating protocone and hypocone; a short thin remnant of the lingual protoloph connected with the protocone is visible, separated from the thick buccal protoloph, which turns forwards. M 1 and M 2. Te molars display close dimensions, M 2 being a little shorter than M 1, and M 3 much narrower (Table 1). Te M 2 differs from the M 1 in their narrower and shorter distal area: the hypocone is more buccal and a little smaller than the protocone, whereas the protocone and hypocone are equal in size on M 1. Te lingual flank of the crowns is higher than the buccal one. Te cusps are bulbous and their apex is acute on juveniles. In adults, the buccal cusps merge within the thick buccal parts of the protoloph and metaloph. Tese lophs are slightly less high than these cusps. Te lingual parts of these lophs are very low or absent. Lingually, the protocone merges within its pre- and post- cristae, which are aligned mesiodistally; it is the same for the hypocone that displays shorter and weaker anterior and posterior arms. Te postprotocrista prolongs through the endoloph. Te endoloph is short and high. Te sinus is barely marked; its height is less than one-third of the crown height, so that it is often indistinguishable on worn teeth. M 1 are easily distinguished from M 2 due to their strong parastyle, which lengthens the buccal edge of the tooth as opposed to the lingual edge and gives it a trapezoidal outline. Te protocone and hypocone are closer on M 1 than on M 2. On M 2, the parastyle elongates as an extension of the anteroloph; it is sometimes swollen [Eg 596 (Fig. 7 a), Ek 123 (Fig. 5 a), holotype (Fig. 4 a), and Ek H 006 (Fig. 6 c)]. Moreover, M 2 mostly show a metacone lingual to the paracone and the hypocone more buccal than the protocone, both making the posterior lobe of the tooth narrower buccolingually than the anterior lobe, and the posteroloph shorter. Te anteroloph is lower and thinner than the anterostyle and the preprotocrista. Te paraconule, merged in the protoloph, can be variably stretched distomesially, more or less swollen. Te protoloph ensures the connection from the paracone to the middle of the protocone, this connection being weak on unworn or weakly worn teeth. Te mesostyle is always present and can be constituted by one or two (mesial and / or distal) small low cusps (e. g., Fig. 5 c) (9 / 29). It is nearly in line with the paracone and metacone, e. g., on the holotype and EK H 005 (Fig. 3 a, 6 a), but it is more buccal on Ek 45 (Fig. 7 d). A short premetacrista can be present (9 / 29) and exceptionally, a postparacrista. A mesoloph is often observed (20 / 29), reaching the level of the metaconule only in three specimens. On Ek 124, the top of the arched metacone is doubled (Fig. 7 f 1, f 3). Te metaconule is strong and bulged, a little lower than the metacone; its position is slightly mesial to that of the metacone and hypocone. Te buccal metalophule I is variably present (19 / 29), sometimes highly reduced; it is lower and thinner than the buccal metalophule II and rarely connects the metaconule (4 / 29). Te buccal metalophule II can be free or join the metaconule (12 / 29). Te ridge connecting the metaconule and the hypocone is very low, therefore these two cusps appear separated on unworn teeth. Te enamel surface is generally smooth. Some rare low wrinkles are present at the bottom of the flexi (10 / 29) (Fig. 5 c, 6 a, 7 g, h). Te unworn specimen Ek 87 (Fig. 7 h) displays unusual features, which might be related to wear differences. Te protocone and hypocone are a little closer than on the other molars, both being acute. Te mesostyle is slender, without mesoloph. Te lingual parts of the protoloph and metaloph are absent; therefore, the paraconule and the twinned metaconule are isolated. Te buccal metalophule II is short and transverse. Te paraconule and metaconule are isolated. M 3. Te strongly reduced hypocone sometimes separates from the protocone (4 / 7) with a shallow sinus. On weakly worn teeth, this sinus communicates with the opposite flexus (2 / 4) (Fig. 7 j, k). Te paracone represents the only salient cusp, on unworn teeth. Te metacone is weakly prominent from the buccal ridge it forms with the mesostylar area. Te paraconule and metaconule are present, but low. Te hypocone is isolated on one weakly worn tooth (Eg 597) and appears connected to the postprotocrista on more worn teeth. Te posteroloph usually connects to the metacone and the hypocone. Dentary. Te body of the mandible Eg 587 (Fig. 8 a) is slender than that of Em 21 [? P. cf. lugdunensis (Fig. 8 b)]. Te diastema of Eg 587 is longer and the radius of curvature of the incisor larger than that of Em 21. Te mental foramen opens anteriorly to the p 4, at the level of the posterior part of the diastema. Te masseteric crest is strong and starts ventral to the mesial root of m 1. Lower teeth (Figs. 9, 10). Among the observed materials from Egerkingen, a few specimens (Ek 67, 69, 70, 76, 81, Ek 248, Ek H 002, and Ek H 002 - 4) display the dental morphological features of the lower jaw of Protadelomys cartieri figured by Stehlin and Schaub (1951, Eg 587) such as the absent mesoconid and the broken distal ectolophid. Te mandible Ek 248 is slightly larger than Eg 587, but the size ratio between the different dental loci is similar (see lines connecting p 4, m 1, m 2, and m 3 in both specimens, in Fig. 3 B). dp 4. Only three teeth can be identified as dp 4 based on the low crown and of the morphology of cusps and ridges, and we can condidently refer Ek 81 to Protadelomys cartieri. It differs from the dp 4 Ek 80 in its wider and shorter crown. Te metaconid is mesiomedian. Te low protoconid is more arcuate than in Ek 80 (Fig. 10 c). Te low ectocingulid is present and long. Te strong bulbous hypoconulid is more buccal, connected to the entolophid, making it oblique posteriorly. Te posterolophid is long. Te dp 4 Ek 80 is slender and less bulbous than Ek 81 (Fig. 9 d). We cautiously refer this dp 4 to P. cartieri. Two mesiodistal extra-ridges develop from the distolingual side of its metaconid to the talonid basin. Its entolophid, similarly poswardly oriented and linked to the hypoconulid, is shorter. Its posterolophid is very short lingually to the hypoconulid. p 4. Te p 4 are smaller than the m 1 and m 2 (Table 1, Fig. 3 B, Additional file 1: S 1). A weak and low anteroconid is present only on Ek H 002 (Fig. 9 a). Te metaconid, the highest cusp, shows two symmetrical cristids, the postmetacristid lingually and the premetacristid buccally, the latter being curved towards the protoconid area. Present on EK H 002 and Ek 127, the protoconid is almost absent on Eg 587 (Fig. 4 b), Ek 248, H 002 - 4, and Ek 67 (Fig. 9). A low ectocingulid is visible on Ek 67 only (Fig. 9 e). Te postprotocristid angles at its end, bearing a short spur or a swelling, considered here as a kind of “ premesoconid ”, as there is neither a distinct mesoconid nor a mesolophid or an ectomesolophid. It connects to the short mesiodistal ridge descending from the metaconid on Ek H 002 - 4; it is free on the others and reduces to a low wrinkle on Ek 67. Te ectolophid interrupts distal to the premesoconid and before its connection with the entolophid and the prehypocristid. Te entolophid is not continuous showing breaks at is center, close to the mesial end of the prehypocristid. On Ek 67, the entolophid connects also with the hypoconulid. Te hypoconulid is usually bulbous. Te posterolophid is short and reduced; it is absent on Ek 67. Te two p 4 (Ek 67, 127) are variants of P. cartieri. Tey differ from the one described above in their more lengthened mesiodistally postprotocristid, and Ek 127 (Fig. 9 f) shows a swelling at the protoconid level, when its sinusid is less deep buccolingually than on Ek 67. Te ectolophid is very short and interrupted shallowly at the hypoconid level. Tere is no prehypocristid on Ek 127, but the hypoconid area appears worn here. Te lingual part of the entolophid is present on Ek 127 only, the entolophid being reduced to a short low ridge on Ek 67. Te post-hypocristid is long and the hypoconulid is moderately swollen. Te posterolophid is thin and join the entoconid on Ek 127, closing the posterosynclinid lingually. m 1 – m 2. Tese teeth display a bulbous and low anteroconid, which individualizes from the protoconid flank on unworn and weakly worn molars. A short anterocingulid can be present, and the anteroflexid is narrower on m 2. A very short anterolophid directs lingually from the apex of the anteroconid. It is separated from the metalophulid I by a shallow anteroflexid, which is open buccally and lingually (Fig. 9 a, b). An anterolophulid is distinct on the worn m 1 of Eg 587, or on the unworn Ek 70 (Fig. 9 g). On Ek 248, the anteroflexid closes buccally, whereas it remains open on Eg 587. Te lingual and buccal parts of the metalophulid I does not connect on m 2 and m 3; therefore, the anteroflexid communicates with the basin. On the other teeth, a low and postwardly angled metalophulid I extends from the metaconid to the apex of the protoconid. Te thick postprotocristid slightly swells at its distal end, and rarely bears a premesoconid spur on m 2 (Ek H 002 and H 002 - 4, Ek 70). Te mesial ectolophid and the mesoconid cannot be distinguished from the oblique postprotocristid, except on the weakly worn m 1 of EK 250 (Fig. 10 a 1). Te distal ectolophid, very short and low, appears interrupted on the occlusal surface. Very low and rare extra-ridges are present on the posterior slope of the metalophulid I to the basin (Figs. 9 a, b; 10 a 2). Te distal part of the ectolophid is much lower than the occlusal surface, and only a short crest joins the very low prehypocristid. Te entolophid is almost straight and only slightly angled. It is interrupted at its buccal part, before joining the prehypocristid. Te entoconulid is variably present and slightly swollen. Te salient hypoconulid separates shallow notches from the post-hypocristid and from the posterolophid on the less worn specimen (m 1 of Ek H 002). Te short posterolophid does not reach the summit of the entoconid leaving the posteroflexid opened lingually. Te weakly worn m 2 Ek 70 differs from Ek H 002, H 002 - 4, and Eg 587 in having a longer buccal anterocingulid, a strong anterolophulid, and a weak ectomesostylid. Ek H 002 is not the specimen figured pl. I, Fig. 2 under this number in Hartenberger, 1969. We noticed that the latter actually corresponds to Ek 250 (NHMB collections). m 3. One weakly worn m 3 of the tooth row Eg 587 (Fig. 4 b), one on Ek 248 (Fig. 10 a) and two isolated much worn m 3 (Ek 68, 76) are available. Te anteroconid is small, not connected to the premetacristid, bearing a short buccal cingulid. A few low and short extra-ridges are visible in the basin. Te thick postprotocristid ends in a short lingual spur. Te very short ectolophid shows the break observed on m 1 – m 2.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFB5DF2FC48346A0FC79F957.taxon	materials_examined	Material from Lissieu (Rhône) referred here to Protadelomys cartieri (Stehlin & Schaub, 1951) Description. Among the smallest m 1 and m 3 of the Lissieu assemblage, five lower teeth stand out from the material assigned to P. lugdunensis: one right m 1 (LIS 54, Fig. 11 b), one weakly worn right m 2 (LIS 60, Fig. 11 c), and three strongly worn m 3 (LIS 30, 33, (Fig. 11 d) and 45 – 3). Tese teeth present some similarities with the typical lower teeth of P. cartieri from Egerkingen, based on the characteristics of the ectolophid, the mesial area, and of the relative proportions of the synclinids. p 4. One p 4, LIS 41 (Fig. 11 a), has a size compatible with the p 4 of P. lugdunensis LIS 42, 37 and 40, but its crown is higher. Te anteroconid, which is buccally displaced, is stretched mesiodistally, connects to a short metalophulid, and joins buccally the protoconid ridge. Te mesiodistally stretched protoconid is more distinct on LIS 41 than on the other p 4 from Lissieu, and the postprotocristid is higher than the ectolophid and makes an obtuse angle with the stretched protoconid. Te postprotocristid is stretched along a mesiolingual to distobuccal axis, and bears distobuccally a bulged postmesoconid spur. Te lingual entolophid is distally oriented from the entoconid, and not connected to its buccal part, reduced to a spur. As the protoconid is present and the ectolophid angles at the level of the mesoconid, which is indistinct, this p 4 is referred to as P. cartieri. m 1. On LIS 54 (Fig. 11 b), the low anteroconid separates from the protoconid, and nearly connects the plunging premetacristid. Te metalophulid I is complete, low, and curved distally. Te postprotocristid is almost as high as the protoconid, and its distal end is swollen; no distinct mesial ectolophid is observed. Te ectolophid is very low, interrupted, and reaches the base of the postprotocristid swelling and of the prehypocristid. Te entolophid, swollen at the level at the base of an entoconulid, attaches directly to the hypoconid. Te hypoconulid is present and small. Te posterolophid joins the base of the entoconid. m 2. On LIS 60 (Fig. 11 d), the anteroconid, low, is closer to the metalophulid I (anteroflexid narrower) than on the m 1, the anterolophid is hardly distinct and the premetacristid is absent. Te ectolophid is interrupted and very short and the entolophid distally attaches to a short prehypocristid. m 3. Among the three m 3 (L = 1. 8 to 1. 95 mm) that are smaller than those of P. lugdunensis (L = 2. 2 to 2. 4 mm), only one (LIS 33) is unworn enough to permit the identification of possible relationships between the cusps and crests (Fig. 11 c). However, on the m 3, the mesosynclinid is not elongated relative to the posterosyclinid, as is the case in P. lugdunensis. LIS 33 shows a small anteroconid connected to the mesial metalophulid (premetacristid). Te distal metalophulid I is not visible due to wear; only its thick buccal part is distinguishable and distally bent like on the m 3 of P. lugdunensis. Apart from this case, the postprotocristid is short, located just in front of the mesoconid. Te quite short ectolophid shows the break observed on m 1 – m 2, like in typical specimens of P. cartieri. Material referred to Protadelom y s cartieri (Stehlin & Schaub, 1952) from Laprade Upper teeth M 1 – 2. One M 1 (LAP 259: Fig. 12 a) and two M 2 (LAP 193: Fig. 12 c, 260: Fig. 12 b) are referred to Protadelomys cartieri. Tey differ from P. lugdunensis from the same locality in their smaller size (but this is not statistically significant, owing to the limited sample, Table 8), in their higher crowns, as well as in the following features: the paraconule does not protrude forwards and swells within the protoloph on the M 1, less on the M 2; the metalophule II is short and can be posteriorly directed (LAP 193, 259); the metalophule I is very reduced; the metaconule is closer to the hypocone than to the metaloph and connects at mid-hypocone by a short low ridge; only a small mesostyle on LAP 259 is present, which is a little stronger and prolonged in a short mesoloph on LAP 260 and LAP 193.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFA1DF19C4B946C0FDF1FB39.taxon	description	(Figs. 13, 14, 15, 16, 17, 18 and 19).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFA1DF19C4B946C0FDF1FB39.taxon	materials_examined	Holotype. L 2627: left M 1 – M 2; Fig. 15 a. Type locality. Lissieu (Rhône); middle Eocene, MP 14. Original diagnosis (Hartenberger, 1969); translation from French. “ Proladelomys more evolved than P. cartieri and? P. alsaticus; lower p 4 more important in the tooth row than for these two species; mesoconid always well-developed; on upper teeth sinus nearly reaching the base of the crown ”. Emended diagnosis. Protadelomys larger than? Protadelomys alsaticus and? P. maximini; nearly similar in size as Protadelomys cartieri. P. lugdunensis differs from P. cartieri in the higher and more robust mandible with relatively short diastema and incisor, the longer and oblique mesiobuccal to distolingual postprotocristid encompassing the mesial ectolophid and a more or less strong mesoconid, the reduced distal ectolophid, the asymmetrical sinus, which is a little deeper (reaching half height of the crown). P. lugdunensis differs from? P. alsaticus in displaying less ornamented enamel, more bulbous cusps, simpler ridges and lophids on lower teeth, DP 4 more molarized (hypocone as robust as protocone on the only one known DP 4), P 4 less triangular, buccal mesoloph often longer, higher buccal protoloph and metaloph, and sinus generally marked by a higher groove (but still shallow and narrow). P. lugdunensis differs from? P. maximini and? P. nievesae in its stronger hypocone, and in the composition of the lingual wall of the upper teeth (i. e., association of pre- and post- protocristae, protocone, endoloph, pre- and post-hypocristae, and hypocone), which is not stretched mesiodistally. Material and measurements from Lissieu. (L: Collections of the “ Université de Lyon ”; LIS: collections of the “ Musée des confluences de Lyon ”) (Sup data, S 2). Like for P. cartieri from Egerkingen, we observed that the dimensions of upper molars in Figures 13 A and 14 A are more scattered than those of the lower teeth (Table 2). Te deciduous and premolar teeth are smaller than the molars, both on the lower and upper tooth row. Te M 1 and M 2 are of similar size and the M 3 are clearly narrower than the first two molars. Te lower m 1 are moderately larger than the m 2. Te m 3 are longer than the m 1 and m 2 (see Figs. 13 B, 14 B; Table 2 A). All the teeth are significantly larger than those of typical? P. alsaticus,? P. maximini, and? P. nievesae (Table 8).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFA1DF19C4B946C0FDF1FB39.taxon	description	Description Maxillary. A few maxillary fragments are represented (L 45 and L 45 - 2). Te infraorbital foramen is hystricomorphous. Te premaxilla – maxilla suture is located in front of the mesial root of the zygoma. Upper dentition (Figs. 15, 16). DP 4. Tere is only one isolated upper deciduous premolar (LIS 11, Fig. 15 b); [Te tooth n ° 2629 (Pl. 4 Fig. 1, in Hartenberger, 1969) is likely a P 4, even if it shows a mesial buccal root oblique forwardly. Its hypocone is not lingual and quite smaller than the protocone. Te anteroflexus and anteroloph are reduced and the protoloph connects to the mesial end of the preprotocrista]. Tis DP 4, moderately worn, is molariform and trapezoidal. Te hypocone is as robust as the protocone, and lingually displaced relative to the protocone. Te parastyle, standing at the same level as the paracone, is weakly swollen. Te anteroloph links lingually to the anterostyle, which connects to the short preprotocrista. Te four main cusps are bulbous. Te protoloph attaches towards the middle of the protocone. Te paraconule stretches mesially from the protoloph to the anterostyle. Te mesostyle stands at the level of the lingual flanks of the paracone and metacone; it is buccolingually compressed and stretched mesiodistally. Some isolated wrinkles form a mesoloph. Te protocone and hypocone connects through a high and short endoloph. Te sinus is very weak and low. Te metacone and the paracone are similar in size, like for P. cartieri. Te metalophule I is present but very low. It does not join the metaconule, contrary to the metalophule II, which is thicker and higher. Te metaconule is bulbous and separated from the hypocone. Te post-hypocrista connects to the posteroloph. Te thickening (worn area) observed at their junction is probably the posterocone. Te posteroflexus opens buccally. P 4. (Fig. 15 c to f). Te P 4 are triangular due to the weak development of the hypocone. Te anteroloph is short and the parastyle is weakly swollen; it is free (1 / 4) or connected to the flank of the paracone (3 / 4). Te mesoflexus is narrow, buccally closed by a more or less swollen mesostyle; the latter is lengthened mesiodistally (2 / 4), reduced (1 / 4) or swollen (1 / 4). In L 2629, the mesostyle connects to a mesoloph, which nearly reaches the metaconule level. In the other cases, the mesoloph is absent or reduced to rare wrinkles. Te paraconule is absent. Te protoloph connects to the preprotocrista (3 / 4) or the anteroloph (L 2629). Te metalophule I is reduced or absent. Te metaconule, swollen, connects to the protocone (L 45 - 2) or the postprotocrista (L 2629). It aligns with the metacone through the metalophule II. Both are oriented towards the posterior part of the protocone, joining only on the most worn specimen (LIS 13). Te hypocone is small (L 45 - 1 and 2, L 2629, LIS 13) or indistinct (LIS 10), connecting the posteroloph. Te latter does not connect to the metaconule, leaving the posteroflexus opened buccally. M 1 and M 2. (Fig. 15 a, f to j; Fig. 16 a to e). Te crowns are unilaterally and moderately hypsodont, like in P. cartieri. Te cusps are bulbous and their apex is acute on juveniles (i. e., weakly or unworn teeth). In adults, the buccal cusps extend within the buccal parts of the protoloph and metaloph, which are slightly less high. In contrast, the lingual parts of the lophs are very low or absent. Lingually, the protocone merge within its pre-and post-protocristae, aligned mesiodistally. Te endoloph prolongs the postprotocrista mesiodistally. Te hypocone shows a similar pattern, but the pre- and post-hypocristae are shorter and weaker. Te endoloph is short and high, but lower than the apexes of the protocone and hypocone. Te sinus is barely marked; its height is less than half the crown height, and it is almost completely absent on worn teeth. Te M 1 can be quite easily distinguished from M 2 due to the strong development of its parastyle, its trapezoidal outline consecutive to the lengthening of its buccal edge compared to the lingual one. On the M 2, this parastyle elongates in an extension of the anteroloph. Moreover, they mostly show a metacone lingual to the paracone, so that the posterior lobe of the tooth appears narrower buccolingually than the anterior lobe, with the posteroloph being shorter than on M 1. Te anteroloph is lower and thinner than the strong anterostyle and the preprotocrista. Te paracone can be rarely divided (1 / 20: L 45 - 1: Fig. 15 g) into a bigger half displaced lingually and a stylar one developed between the end of the parastyle and the mesostyle. Te paraconule attaches to the protoloph and protrudes mesially. Te protoloph ensures the connection from the paracone to the middle of the protocone. Te mesostyle is stretched mesiodistally or bulbous. A low mesoloph reaches the level of the metaconule (9 / 19); it can be shorter (6 / 19) or absent (4 / 19). Te strong metaconule is slightly lower than the metacone; its position is slightly mesial to that of the metacone and hypocone. Only one specimen (LIS 5) displays a duplicated metaconule. Te metalophule I is always present, sometimes much reduced, lower and thinner than the metalophule II. Te latter is generally free and rarely joins the metaconule (2 / 20). Te metaconule weakly connects to the hypocone. Te enamel surface is generally smooth. Some rare low wrinkles are visible at the bottom of the flexi (LIS 1, 2, 8, 26); in one case (L 45 - 1), they are present as surface folds of the lingual flanks of the protocone and hypocone. M 3. (Fig. 16 f to h). Te hypocone is strongly reduced. It often separates from the protocone (3 / 4) with a shallow sinus. On weakly worn teeth, this sinus communicates with the opposite flexus (2 / 4). Te paracone constitutes the only salient cusp. Te metacone is only slightly higher than the buccal ridge it forms with the mesostylar area. (See figure on next page.) Fig. 15 Upper teeth of Protadelomys lugdunensis Hartenberger, 1969 from Lissieu (Rhône). a L 2627, left M 1 – M 2, Holotype; a 1, occlusal view; a 2, lingual aspect. b LIS 11, left DP 4; b 1, occlusal view; b 2, lingual aspect; b 3, buccal aspect. c: L 2629, left P 4; c 1, occlusal view; c 2, lingual aspect; c 3, buccal aspect. d LIS 13, right P 4, occlusal view. e LIS 10, right P 4; e 1, occlusal view; e 2, lingual aspect; e 3, buccal aspect. f LIS 45 – 2, left P 4 – M 1; f 1, occlusal view; f 2, buccal aspect of P 4; f 3, lingual aspect of P 4. g L 45 – 1, left M 1; g 1, occlusal view; g 2, lingual aspect; g 3, buccal aspect. h LIS 14, left M 1, occlusal view. i: LIS 6, left M 1, worn; i 1, occlusal view; i 2, lingual aspect; i 3, buccal aspect. j: LIS 22, right M 1, strongly worn; j 1, occlusal view; j 2, lingual aspect; j 3, buccal aspect. Scale bar: 1 mm Te paraconule and metaconule are present, although low. Te posteroloph can be either distinct from the metaconule (1 / 4) or connected to it (3 / 4). Dentary. (Fig. 17). Te body of the mandible is robust and high (L 44). Te diastema and incisor are short. Te mental foramen opens close to the mesial root of p 4. Te masseteric crest is strong and begins below the mesial root of m 1. Lower dentition. (Figs. 18, 19). dp 4. Only one tooth (LIS 40; Fig. 18 b), smaller than the p 4, could be identified as a dp 4. Its posterior buccal root is inclined distally. Te metaconid is mesiolingual. Te enamel has weak and rare wrinkles along the lingual flank of the long postmetacristid. A short mesiodistal anterolophid attaches to the metaconid. A bulged anteroconid is present buccally. A short transverse metalophulid I connects the metaconid to the protoconid, from which a low mesiodistal ridge descends in the basin. Te lophid constituted by the stretched protoconid, the postprotocristid and the mesial part of the ectolophid is weakly oblique, almost mesiodistal. Te short distal part of the ectolophid links the anterior arm of the hypoconid with the entolophid. Te latter makes an angle oriented distally at the level of a small entoconulid. Te hypoconid is higher than the hypoconulid, which is swollen. Te long posterolophid reaches the mesiolingual edge of the entoconid. A small ectostylid is present at the base of the sinusid. p 4. Te p 4 is slightly shorter than m 1, and clearly narrower (Table 2 A). An anterior tooth is listed as a dp 4 in the collection from Lissieu (LIS 47). However, the height and robustness of the crown and roots rather evoke a p 4. Four isolated p 4 (LIS 42, 37, 39, and 41) are similar in size with p 4 included in dental rows (L 2625, L 2628, and LIS 44). However, the crown of LIS 41 is slightly higher than the others are and displays some peculiar features, which enable us to reattribute it to P. cartieri (see below). LIS 32, LIS 37, and L 2625 (Fig. 18 d, e, f) show neither an anteroconid nor an anterolophid. Te metaconid is the only cuspid of the mesial border of the crown, which is then narrower than the other p 4. On L 2628 (Fig. 18 a), the anteroconid is buccal and isolated; a short buccal cingulid descends from the mesial end of the preprotocristid. A short mesiobuccal anterolophid is visible on L 44 (Fig. 18 e). Te metaconid is the highest cusp of the tooth; it connects to the protoconid, which is low, weak and stretched in a mesiodistal protocristid including its pre- and post-protocristae. A very low ectocingulid sometimes interrupts an asymmetrical sinusid (LIS 32, 37, and 44). On the lingual side, the postmetacristid descends to the lingual notch. A low crestid extends from the metaconid apex to the mesoflexid; it can be continuous or not and composed of two to three very low crestules. Te mesostylid is absent. Te ectolophid is very short and distally interrupted superficially, separating the postprotocristid from the prehypocristid (LIS 37 and L 44). On L 2628, the entolophid reduced to its lingual half is posteriorly oriented, connected to the mesial end of the prehypocristid. On the others, the entolophid is complete from the entoconid to the junction prehypocristid – ectolophid. Te posterolophid is well separated from the entoconid, so that the posterosynclinid opens lingually. m 1. Te m 1 are smaller than the m 2, the protoconid – metaconid width is lower than the hypoconid – entoconid (See figure on next page.) Fig. 18 Lower teeth of Protadelomys lugdunensis Hartenberger, 1969, from Lissieu (Rhône), a L 2628, right p 4 – m 3; a 1, occlusal view; a 2, buccal aspect. b LIS 40, left dp 4; b 1, occlusal view; b 2, buccal aspect; b 3, lingual aspect. c: LIS 32, right p 4; c 1, buccal aspect; c 2 occlusal view; c 3, lingual aspect. d L 2625, right p 4 – m 3; d 1, occlusal view; d 2, lingual aspect. e L 44, right p 4 – m 2; e 1, occlusal view; e 2, buccal aspect; e 3, lingual aspect. f LIS 37, right p 4; f 1, occlusal view; f 2, buccal aspect; f 3, lingual aspect. Scale bar = 1 mm width. Te lingual side is only slightly shallower than the buccal side. Te anteroconid is individualized on unworn or weakly worn teeth (LIS 67 and 66; Fig. 19 a,), and the anterolophid is short. Te lingual anterocingulid and the anterosinusid are weakly marked or absent. Te anterolophulid is rarely visible, mainly on worn teeth, low and connects the anterior flank of the protoconid to the anteroconid. Te metaconid represents the highest cusp, from which the postmetacristid steeply descends. Te metalophulid I is generally angled, its lingual and buccal parts distally oriented; the latter is connected to the protoconid apex. On weakly worn teeth, the enamel surface of the mesoflexid bears short mesiodistal wrinkles and granules (LIS 67, 68, and 72). Exceptionally, these wrinkles merge into longer mesiodistal ridges, which are connected to the metalophulid I (L 2628, Fig. 18 a). Te postprotocristid is thick and oblique; it extends to a more swollen area that corresponds to the mesoconid. Both are discernible on slightly worn teeth (LIS 53, 55, 67, and 68, Fig. 19 a, c, h), while they appear in continuity in a long oblique lophid on more worn teeth. Some teeth have a bulbous and low anteroconid and the buccal cingulid is reduced to absent (LIS 68, 69, and 71); but their size and other features are not different of the other teeth of P. lugdunensis. A short ridge on its lingual side indicates the mesoconid area; there is no well-individualized mesolophid. Some teeth have an ectomesolophid connected to an ectostylid (LIS 55 and L 2628), while other specimens only display an ectostylid more or less swollen (L 2625, LIS 53, 68, and perhaps on badly preserved LIS 70). Te distal part of the ectolophid is very short and low. It appears interrupted on unworn or weakly worn teeth (LIS 53, 68, and 70). Te distal part of the ectolophid ensures the connection with the mesial end of the short prehypocristid and the buccal part of the entolophid. Tis buccal part is lower than the lingual one and it is partly missing on two teeth (LIS 67 and 68); both parts make a distally oriented angle. Te lingual half of the entolophid is swollen at its base, his swelling being reminiscent of an entoconulid. Te post-hypocristid is connected to the hypoconulid; it is always well defined and higher than the posterolophid, which is short. If the lingual border of the posterosynclinid is rather high, the posterolophid itself does not merge with the entoconid. Tis cusp is lower than the hypoconid on weakly worn teeth, but the lingual side is higher than the buccal one on worn teeth: wear affects more the buccal half of the tooth than its lingual counterpart does. m 2. (Fig. 19 e to i) Te m 2 differ from the m 1 in showing a larger size, a rectangular outline due to the equal protoconid – metaconid and hypoconid – entoconid widths, a longer anterolophid and sometimes a longer anterocingulid, as well as different connections with the metalophulids. Te anteroconid is less bulbous than on the m 1; the anterolophulid is lingual, often linking the anterolophid with the lingual metalophulid I, thereby forming a longer antesinusid. Other features are more similar between m 1 and m 2: secondary wrinkles on LIS 42, 59, 66, and 69 (Fig. 18); long low mesiodistal crestids and ectomesostylid on L 2628; the presence of an ectostylid on LIS 42, 56, 57. As wear increases, the posterolophid can reach the apex of the entoconid (LIS 51, 59; L 2625 and L 2628), even on a moderately worn m 2 (LIS 42). m 3. Te m 3 are clearly longer than the m 2 (x about 1. 15), due to the lengthening of the mesosynclinid area, they are as wide as to slightly wider as the m 1 (Table 2 A). Te posterior area is narrower than the anterior one. Te lingual metalophulid I is more mesial, it is thus closer to the anterolophid (L 2628, LIS 29; Fig. 18 a, 19 k) with which it is often connected (LIS 46, 39; Fig. 19 l). Te connection between the lingual metalophulid I and the buccal metalophulid I can be disrupted, or reduced to weak short ridges Te anterolophulid is buccal when present (LIS 29, 36, 39). Te low mesiodistal ridges arising from the buccal metalophulid I extremity are more frequent and more visible on weakly worn teeth (LIS 29, 36, 39, and L 2628). Te entoconid is reduced, which puts the metaconid in relief. Te entolophid makes an angle pointing posteriorly at the level of the entoconulid, when the latter is present (LIS 49, 46). Te buccal part of the entolophid is sometimes absent (L 2625 and LIS 46). On L 2628, the apex of the angle defined by the lingual and buccal parts of the entolophid connects to the hypoconulid by a thin mesiodistal ridge; ditto for LIS 46 but the entolophid is very incomplete on this tooth. Te hypoconulid is bulged while the posterolophid is lower, short, and does not reach the entoconid.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFA1DF19C4B946C0FDF1FB39.taxon	materials_examined	Material referred to Protadelomys lugdunensis Hartenberger, 1969 from Egerkingen	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFA1DF19C4B946C0FDF1FB39.taxon	discussion	Remarks Together with the mandible Em 21 (Fig. 8 b, Fig. 20 b) and a few isolated molars, we tentatively refer the specimen Ek 592 (right M 1 – M 2), identified as? Protadelomys in Hartenberger (1969) to? P. lugdunensis. Te ratio between M 1 and M 2 width is different from that of the holotype of cartieri in which M 1 and M 2 have same length, whereas here the M 2 is well shorter than the M 1 (Fig. 19 a), the cusps more acute and higher, and looks like the holotype of P. lugdunensis from Lissieu, LIS 2627. Among the lower teeth from Lissieu, we detected similar variation as that observed in the lower teeth of specimens previously identified as P. cartieri from Egerkingen (Hartenberger, 1969), which led us to find both species in Egerkingen and Lissieu (see below and discussion). P. cartieri is more abundant than P. lugdunensis in Egerkingen, whereas the opposite is true in Lissieu.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFA1DF19C4B946C0FDF1FB39.taxon	materials_examined	Material referred. Em 21, right lower jaw, with p 4 – m 3; Ek 71, weakly worn right m 2, Ek 126, right m 1, Ek 72, right m 3; EK 592, right M 1 – M 2; all in Fig. 20. Locality. Egerkingen? α and fissure indet., upper middle Eocene (MP 14?).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFA1DF19C4B946C0FDF1FB39.taxon	description	Description Upper teeth Ek 592, M 1 – M 2, (Fig. 20 a) the paracone and metacone are relatively high and acute. Te parastyle is present and swollen on M 1, less on M 2. Te anteroloph extends between the parastyle and the anterostyle. Te paraconule, a little less robust than the metaconule, is protruding from the protoloph, the latter being sinuous at this level. Te protoloph connects lingually at the center of the protocone. Te preprotocrista, the protocone, and the postprotocrista are aligned mesiodistally, the protocrista being relatively short and slightly separated from the anterostyle by a shallow notch. A strong mesostyle is present, linked to a thin low ridge, as a lingual mesoloph. A few low and thin extra-ridges are visible and restricted in the center of the mesoflexus. A clear but low metalophule I connects the metacone and the strong metaconule, which is mesially protruding. Te buccal part of the metalophule II, more distal, is short and high. A deep notch separates the metalophule II from the metaconule, which is free from a very short lingual spur of the metaloph visible at the level of the mid-hypocone. Te endoloph is relatively low compare to the hypocone. A shallow, narrow, and short sinus is marked between the two lingual main cusps. Te hypocone is nearly as robust as the protocone. Te posteroloph is long on M 1, reaching the base of the mesiobuccal corner of the metacone. It is much shorter on M 2, on which it stops at the level of the lingual end of the buccal metalophule II. Lower jaw with p 4 to m 3. Te body of the mandible Em 21 (Fig. 8 b) is wider and higher than that of Protadelomys cartieri Eg 587 (Fig. 8 a). Te diastema is shorter and the radius of curvature of the incisor is smaller. Te mental foramen opens more distally, below the anterior root of the p 4. Te masseteric crest is strong and begins below the mesial root of m 1. Te incisor is thicker. p 4. (Fig. 20 b) Te metaconid is mesiolingual and a small anteroconid is mesiobuccal, both separated by a well-marked notch. A short metalophulid connects the metaconid and a low incipient protoconid. Te posprotocristid is long and oblique, bearing a short lingual premesoconid spur. In the continuity of the posprotocristid, the ectolophid bears a stretched mesoconid, associated to a short mesolophid and ectomesolophid. Te asymmetrical sinusid (ectoflexid) is limited buccally, at its base, by a strong ectostylid. Te distal part of the ectolophid connects to the mesial end of the prehypocristid. A buccal short lophid joins more mesially this ectolophid; it corresponds to the buccal part of the entolophid of the molars. On p 4, the lingual part of the entolophid turns postwardly to the junction hypoconulid – posterolophid. Te latter is short and does not join the entoconid. Molars. m 1 is a little wider than p 4, and shorter and narrower than m 2 and m 3 (Additional file 1: S 1, Fig. 3, Table 1 B). Te anteroconid extends buccally through a clear anterocingulid; it lingually joins the mesial branch of the lingual metalophulid (premetacristid). Another lingual branch, close to the former, turns posteriorly and joins the transverse buccal metalophulid I (Fig. 20 b 1). Te anteroflexid is defined between the anteroconid and the elements constitutive of the metalophulid I; it is limited buccally by an anterolophulid. Te metaconid is clearly higher than the entoconid, resulting in a steep postmetacristid (Fig. 20 b 3). One short mesiodistal spur descends in the basin located at the buccal end of the lingual metalophulid I, the enamel of which bears low and weak extra-ridges. Te postprotocristid, the ectolophid and mesoconid are in line obliquely. Te mesoconid displays two short spurs, one lingual (mesolophid) and one buccal (ectomesolophid). Like on p 4, the strong ectostylid limits the sinusid buccally. Te entolophid, complete, connects to the junction ectolophid – prehypocristid. An entoconulid is more visible on m 2 than on m 1 and m 3. Te hypoconulid is bulged at the end of the post-hypocristid, and the posterolophid is short, distinct from the entoconid, leaving the posteroflexid opened lingually. Other lower teeth. Te m 1 Ek 126, m 2 Ek 71 and the m 3 Ek 72 (Fig. 20 c, d, e) shares some features with Em 21, like the reduced or absent lingual metalophulid I, the long oblique postprotocristid bearing a more or less distinct mesoconid, an ectomesolophid, the posteroflexid closed lingually, the long postmetacristid. However, the entolophid is not connected buccally, and these teeth do not display an ectostylid. On the m? 1 Ek 126, the anteroconid is connected to the preprotocristid, closing the anteroflexid buccally, whereas it is stretched and free and developed more lingually and buccally on the others. Te buccal metalophulid I is transverse on Ek 71 and Ek 72, it is oblique distobuccally on Ek 126. Te mesoconid is barely distinguishable; it corresponds to the ectolophid area located at the end of the thick oblique posprotocristid. A small mesoconid seems to be present on the m 3.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFA1DF19C4B946C0FDF1FB39.taxon	materials_examined	Material referred to Protadelomys lugdunensis (Hartenberger, 1969) from Laprade (Tarn-et-Garonne, Quercy) List of material and measurements: Additional file 3: S 3; Te size of the teeth from Laprade does not differ significantly from that of the teeth of Protadelomys lugdunensis from Lissieu (Tables 2, 3, 8; Fig. 21).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFA1DF19C4B946C0FDF1FB39.taxon	discussion	Remarks Sudre et al. (1990) have identified the larger theridomorphs from Laprade as Protadelomys cf. lugdunensis. A smaller species was referred to Elfomys nov. sp. (Marandat et. al .. 1993; Comte et al., 2012). Here, we reassess these species and also distinguish a few teeth that could be referred to P. cartieri, and possibly another species differing from both P. lugdunensis and P. cartieri. Te co-occurrence of P. cartieri and P. lugdunensis has been reported above, both from Egerkingen and Lissieu.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFA1DF19C4B946C0FDF1FB39.taxon	description	Description Maxillary. Two maxillary fragments are present in the collection (Fig. 22). LAP 224 is an incomplete right hemi-maxillary, showing the alveolus of DP 3; DP 4 is broken and a fragment of an erupting P 4 is present in the mesiobuccal alveolus; only the nearly unworn M 1 is present associated with damaged alveoli of M 2 and M 3. Te mesial end of the zygomatic process of the maxilla is preserved, which enables to characterize the wide opening of the infra-orbitary foramen in frontal view. In palatal view, the maxillary part of the palate is flat and wide, showing the edge for the insertion of the superficial masseter between the level of the alveolus of DP 3 and the premaxillary – maxillary suture. Medial to that, there is a trace of the distal closure of the anterior palatine foramen (f. p. a.). Posteriorly, the palate is damaged, but the maxilla – palatine suture can be distinguished at the level between M 1 and M 2. LAP 83 is a fragmentary left maxilla, broken at the level of the mesial alveoli of M 2. On this specimen, a longer part of the zygomatic arch is preserved, showing the ridge of the insertion of the anterior deep masseter, and the posterior edge of the f. p. a .. LAP 83 shows the alveoli of P 4 and a remnant of the alveolus of D 3. Te only molar preserved, the M 1, is more worn than those of LAP 224. Upper teeth DP 4. Only one unworn DP 4 is present in the material from Laprade (LAP 191; Fig. 23 a). It is molariform, very similar to LIS 11 from Lissieu. Because it is unworn, the anterostyle is superficially separated from the short preprotocrista. Te paraconule is displaced mesially from the protoloph. Te mesostyle weakly protrudes buccally; it is located closer to the metacone than to the paracone and associated to a weak premetacrista. A very low ridge is reminiscent of the presence of a mesoloph. Te metalophule I is weak, low, and does not join the metaconule, unlike the metalophule II, which is thicker and higher. A minute additional conule is present between the bulbous metaconule and the metacone. Te hypocone, more lingual than the protocone, is separated from the posteroloph by a shallow notch. P 4. Tere are four well preserved and two damaged P 4. A first group of three P 4 is very similar to the P 4 known from Lissieu. Tey (LAP 84, (Fig. 23 d), 192 (Fig. 23 b), 223, L 45 – 2) are characterized by a weak development of the hypocone. Te parastyle is absent. On a weakly worn P 4 (LAP 192), the anterostyle is slightly swollen, separated from the protocone and connected to the short anteroloph. Teir mesostyle is swollen and not connected to a mesoloph. Some low wrinkles are present in the mesoflexus; they can be in line with the mesostyle forming a kind of mesoloph (LAP 84, Fig. 23 d), or directed towards the metaconule (LP 223) or the protoloph (LAP 192). Te protoloph does not reach the protocone on LAP 84, 192, and 269; LAP 223 is worn at this level. Te metalophule I is absent. Te metaconule is strong and isolated on unworn teeth (LAP 84, 192 and 269), or connected to the metacone and the hypocone on worn teeth (LAP 223). Te two other specimens, weakly worn (LAP 235: Fig. 23 c, and LAP 272) display different features. Te anteroloph is much reduced and low. A short distomesial crest (anterolophule) links the buccal part of the protoloph to the anteroloph on LAP 235 (Fig. 23 c). On both teeth, the protoloph connects to the preprotocrista. Te protoloph makes an angle pointing anteriorly at the middle of the occlusal surface of the crown. Te mesostyle is twinned, the main part being prolonged in a mesoloph directed towards middle of the mesoflexus. Te strong bulbous metaconule shows a short buccal ridge, ending free on LAP 235, and extended to the metacone mesial flank on LAP 272. On the latter, there is also a short lingual ridge, which is oblique mesially. Tese ridges may constitute remnants of the metalophule I. Te short metalophule II is distally oriented. Te P 4 L 45 - 2 from Lissieu shows a similar orientation of the short branch of the metalophule II, but the other features resemble more those of the first group of P 4. M 1 and M 2. (Fig. 23 e to m) Teir features are very close to those of P. lugdunensis from Lissieu. Te paracone is never divided on the M 1 – 2 (contra in a few specimens from Lissieu: e. g., L 45 - 1). Te paraconule is developed mesially to the protoloph. Te latter ensures the connection from the paracone to the middle of the protocone, which is bulbous at its base and pinched at its upper part from the preprotocrista to the postprotocrista. Te low mesoloph reaches the level of the metaconule (2 / 19); it is often shorter (15 / 19) and rarely absent (2 / 19). Te strong metaconule is swollen, a little lower than the metacone; it is slightly mesial to the metacone and hypocone. One specimen (LAP 196) displays a duplicated metaconule. Te buccal part of the metalophule I is absent or reduced to a low and short ridge. Te metaconule is sometimes more mesial; the metalophule II is directed distally from the metacone, but does not reach the posteroloph. Among these teeth, three (LAP 85, 86, and 203) are a little smaller than the abovementioned specimen, but shows very similar features, with a metalophule I being well present for LAP 85 and 203. M 3. Te hypocone is strongly reduced, separated from the protocone (LAP 206: Fig. 23 n) by a shallow sinus or connected by a low and short curved endoloph (LAP 25: Fig. 23 o). Dentary. (Fig. 24) Te body of the mandible (LAP 246) is robust and dorso-ventrally deep, as in typical P. lugdunensis. Even if the mandible is fragmentary, the preservation of a posterior part of the symphysis indicates that the diastema is short. Te mental foramen opens a little anterior to the mesial root of p 4 as for typical P. lugdunensis. Te masseteric crest is strong and begins below the mesial root of m 1 as for P. lugdunensis from Lissieu and P. cartier i from Egerkingen. Lower teeth. Te lower p 4, m 1, and m 2 are significantly larger than those of P. lugdunensis from Lissieu (Fig. 21 B; Tables 2, 3, 8). dp 4. (Fig. 25 a, b, c) Tree specimens are deciduous teeth on the bases of the low crown, low profiles and shape of cusps, low ridges and diverging roots. We attributed these teeth to P. lugdunensis, even if dp 4 are poorly documented in Lissieu, because the crown is higher than that of P. cartieri. Te metaconid is mesiolingual and constitutes the highest cusp. Te protoconid is low but well distinct from the metaconid due to the presence of a shallow mesiobuccal notch. A very short and low anterolophid (LAP 208 — 209) or anteroconid (LAP 207) limits this notch mesially. Te sinusid is asymmetrical and short buccolingually. One slender long ridge descends from the premetacristid in the trigonid basin: it nearly reaches the bottom of the mesosynclinid. Te hypoconid, entoconid, and hypoconulid display similar height, but the latter is weakly swollen. Te entolophid is continuous, transverse, and connected to the junction ectolophid – prehypocristid. Te ectolophid is low when compared to the hypoconid. Te posterolophid is longer than that of P. cartieri (Ek 80). p 4. (Fig. 24 a 1; Fig. 25 d to g) p 4 are represented by four isolated teeth (LAP 210, 212, 251, and 256) and one damaged on the mandible LAP 246. Te p 4 are shorter than m 1 (Table 3). Two p 4 have a reduced buccal anteroconid. Te metaconid is strong and mesiolingual. Te protoconid is poorly developed and merged in the protocristid. Te enamel shows low wrinkles along the protocristid and / or the median crestid. Te ectolophid is weakly swollen within the mesoconid. An ectocingulid is present and long, joining the protocristid to the buccal edge of the sinusid (LAP 212: Fig. 25 f). Tere is no mesostylid. Te ectolophid is very short and interrupted shallowly, and sometimes separates the ends of the postprotocristid from the prehypocristid (LAP 251: Fig. 25 g; LAP 256: Fig. 25 e). Te orientation and position of the entolophid vary. On LAP 251, from its lingual part, a short ridge is posteriorly oriented to the hypoconulid while another ridge is directed anteriorly to the ectolophid, near the mesoconid. On LAP 256, the lingual part of the entolophid is longer and oblique postwards to the hypoconulid. A narrow, shallow slit separates this lingual part of the entolophid from its very short buccal part. On LAP 256, the low entolophid joins the extremity of the prehypocristid. Te posterolophid reaches the entoconid posterior flank, closing the posterosynclinid lingually. m 1. Te m 1 are smaller than the m 2 (Fig. 21 B; Table 3) the protoconid – metaconid width is lower than the hypoconid – entoconid width. As often seen on weakly worn teeth, the lingual side is as high as the buccal side. Te anteroconid is bulbous on unworn to weakly worn teeth and the anterolophid is short. Te lingual anterocingulid and the anterosinusid are weak or absent. Te anterolophulid is low, rarely visible, mainly on worn teeth on which it connects the anterior flank of the protoconid to the anteroconid. On LAP 87 (Fig. 26 e), a metastylid is distinct on the postmetacristid. Te buccal and lingual parts of metalophulid I often make an obtuse angle open mesially; the lingual part is weak to absent (LAP 87), distally oriented (LAP 215, 254: Fig. 25 h, LAP 217: Fig. 25 i, LAP 218: Fig. 26 j) or mesially (LAP 216: Fig. 25 k, LAP 220); the buccal part is connected to the center of the protoconid. On weakly worn teeth, the enamel surface of the mesoflexid bears short wrinkles and granules. Te postprocristid is oblique and extends to the mesoconid, which is swollen. Te lingual side of the mesoconid is rarely prolonged by a short edge on its lingual side. Buccally, the ectomesolophid is absent. Te short ectolophid connects together to the mesial extremity of the short prehypocristid and to the buccal end of the entolophid, which is complete. Te entolophid weakly angles at the junction of its lingual and buccal parts. Te entoconulid and the hypoconulid are more or less swollen. Te posterolophid is short. m 2. (Fig. 26 a to d) Teir morphology is similar to that of P. lugdunensis from Lissieu, apart from their size, which is larger, and their rectangular outline due to the equal protoconid – metaconid and hypoconid – entoconid widths. Te m 2 differ from m 1 in their longer anterolophid. When present, the anterolophulid has a more lingual position, often joining the anterolophid to the lingual metalophulid I, thereby forming a longer antesinusid. On LAP 249 (Fig. 26 b), the metalophulid I is not angulated, but rectilinear and continuous from the metaconid to the protoconid; on this unworn tooth, the wrinkles and granules are numerous, but still very low. Te posterolophid is always short. m 3. (Fig. 26 f, g, h) Te m 3 are longer and narrower than the m 2 (Table 3 A). Te lingual metalophulid I is mesially oriented and joins the anterolophid. On unworn LAP 233 the buccal metalophulid I is forked, (Fig. 26 g) a part oriented distally, like on the other m 3, and a short edge oriented mesially. Te anteroconid is swollen on LAP 233 and LAP 230; it is stretched as a longer anterolophid on the other teeth. Te connection between the lingual and the buccal metalophulids is broken. Granules are rare on the enamel surface. Te entolophid is very low; its buccal part is also very low or absent (LAP 232, 233, 252: Fig. 26 h, and 253: Fig. 26 g). Te hypoconulid is poorly inflated. Te posterolophid is short and does not reach the entoconid.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF95DF19C48347CBFA9BFD19.taxon	description	As the materiel of P. lugdunensis from Egerkingen most of the lower teeth of P. lugdunensis from Lissieu display obliquely aligned postprotocristid – ectolophid. Te main difference with the lower teeth of P. cartieri, as defined by Stehlin and Schaub (1951), corresponds to the absence of a mesoconid associated to a disruption of the distal ectolophid distally in P. cartieri. Te P 4 are only slightly less reduced in P. lugdunensis than in P. cartieri, and the m 3 are only slightly longer. We have not confirmed that the lower p 4 of P. lugdunensis from Laprade are less molarized than those of P. lugdunensis as previously considered (Sudre et al., 1990: 20). Tey have similar length and the protoconid is weak as in typical P. lugdunensis from Lissieu. Te P 4 are not significantly narrower than typical P. lugdunensis (Table 8), but similar in size to P. cartieri. Extra-ridges and granules are variably present as for the teeth from Lissieu, and the M 3 are not significantly smaller than the M 3 from Lissieu (Tables 2, 3, 8). Tese features cannot support the idea that the teeth from Laprade, are more “ primitive ” than those from Lissieu (Sudre et. al.: 20), bearing in mind their limited number. It would rather be the opposite, some dental categories (p 4, m 1, m 2) in Laprade being actually significantly larger than specimens from Lissieu (Table 8), while dental size increase was often observed through the evolutionary history of these lineages (e. g., Escarguel, 1998; Vianey-Liaud & Marivaux, 2017). In addition with the differential development of extra-ridges and granules, most of the morphological differences observed between P. lugdunensis and? P. alsaticus involve the lower teeth: in P. lugdunensis, the cusps seem more bulged, the entolophid is more continuous, the postprotocristid is fused to the mesoconid area (no distinct pre, meso and postmesoconid elements).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF9ADF14C73943E6FD5EF899.taxon	discussion	Remarks On the bases of the observation of “ two upper jaws ” of the Naturhistorisches Museum Basel, Hartenberger (1969, p. 46) wrote that the M 2 are stronger than the M 1, their posteroloph being longer on M 2. For this reason, he assigned the specimen BUX 67 – 2 (Te holotype) to an M 1. However, on this specimen, the metacone – hypocone width is reduced compared to the paracone – protocone width, and the posteroloph is short, like for the M 2 of the species of Protadelomys (P. cartieri or P. lugdunensis) and more generally of the basal Teridomorpha. Te holotype rather represents an M 2. Moreover, the size distribution (L x w) of the M 1 overlaps that of M 2 (Fig. 27 A and Table 4), which indicates that M 2 are not typically larger than M 1. Furthermore, the two specimens from the NHMB (Buchs 660) are for one an upper jaw with M 1 – M 3 and the other one a lower jaw with p 4 – m 2. Tey are not “ two upper jaws with P 4 – M 1 – M 2 ” (Hartenberger, 1969: 46).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF9ADF14C73943E6FD5EF899.taxon	materials_examined	Holotype. M 2 (not M 1) sup. (Hartenberger, 1969: PI. 2, Fig. 6; this paper: Fig. 31 a). Type locality. Bouxwiller (green marls), France; middle Eocene, MP 13. Original diagnosis (translation from French). “ Protadelomys more primitive than P. cartieri. Upper molars with sinus less pronounced. Paraconule, metaconule, and mesostyle variable in volume. Short lower fourth premolars. Numerous wrinkles. ” New diagnosis. Dental morphology “ intermediate ” between Masillamys and Protadelomys. Unilaterally weakly hypsodont teeth; almost the entire teeth bear more or less low extra-ridges and granules, and enamel surface of the outskirts of the crown (mainly lingual for upper teeth or buccal for lower teeth) displaying faint and oriented ornamentations from the base to the edge of the crowns. Upper teeth: DP 4 a little smaller than P 4 and more molarized, with the hypocone lingually displaced. P 4 with paracone and metacone more bulbous than on DP 4; paraconule mesially displaced and fused with the anterostyle; small hypocone variably present. On molars, the hypocone is slightly smaller, to nearly equal in size, than the protocone. Te M 2 posterior width is shorter than the M 1 posterior width. Te postparacrista and premetacrista variably present. Te mesostyle is always present; the mesoloph sometimes reaches the mediobuccal ridge of the metaconule. A small hypocone is generally present on M 3. Lower teeth: the anteroconid is buccally located on molars; it is more or less lingually stretched, and sometimes joins the linguobuccal premetacristid. Te anterocingulid is absent. Te mesial metalophulid I often corresponds to the premetacristid, which sometimes bifurcates or is parallel to a more distal branch of the metalophulid I. Te latter sometimes turns distally to join the buccal transverse branch of the metalophulid I. Te postprotocristid is thick, long and oblique; its distal end is often swollen with a premesoconid spur. Te short mesiodistal ectolophid bears a small mesoconid. Te ectomesolophid and mesolophid spurs are variably developed or absent; the postmesoconid swelling and / or spur is present and variably developed. Te entolophid lowers buccally, and attaches either to the postmesoconid spur / swelling or to the junction of the distal part of the ectolophid and the prehypocristid, which is short. Te hypoconulid is bulged and sometimes duplicated.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF98DF06C73943E1FEB1FA79.taxon	description	- Protadelomys lugdunensis in the absence of anterocingulid on lower teeth, and the enamel ornamentations more numerous and frequent. -? Protadelomys maximini in the often longer and more frequent mesoloph, the less reduced hypocone, and the thicker lingual wall. - Tardenomys. On upper teeth: In the constant presence of a high endoloph, a thick prehypocrista and well-distinct metaconule, isolated from the hypocone; in the absence of a fully developed mure. On lower teeth: in the less frequent and weaker oblique extra-ridge from the lingual extremity of the buccal metalophulid I, and in the absence of buccal cingulid developed from the anterolophid together with the buccal closing of the anteroflexid.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF98DF06C73943E1FEB1FA79.taxon	materials_examined	Material and measurements: (Additional file 4: S 4, Table 4 A). Numerous isolated teeth (6 DP 4, 11 P 4, 69 M 1 – M 2, 21 M 3; 9 dp 4, 9 p 4, 64 m 1 and m 2, 21 m 3) come from the green marls at the base of the Bouxwiller Quarry. Most of the teeth belong to the species? P. alsaticus here revised. Tey are provisionally attributed to the paraphyletic genus? Protadelomys. Te congruence between the size and some morphological features makes it possible to distinguish several morphotypes within the assemblage from Bouxwiller that are plotted on the Figure 27 (morphotype 1: light violet; morphotype 2. turquoise; morphotype 3: light green). Like for P. cartieri and P. lugdunensis? P. alsaticus displays DP 4, P 4, dp 4 and p 4 smaller than the molars. M 1 and M 2 are similar in size and the M 3 are clearly narrower than the other molars, but are close in length. Te m 1 are moderately smaller than the m 2. Te m 3 are longer than the m 1 and m 2, but less than in P. lugdunensis. Te teeth are a slightly smaller than P. cartieri (Fig. 28; Tables 1, 4), and significantly smaller than P. lugdunensis (Table 8).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF98DF06C73943E1FEB1FA79.taxon	description	Description. Most of the teeth bear more or less low additional ridges and granules, and the enamel surface of the crown displays faint and oriented ornamentations from the base to the edge of the crowns, which are mainly lingual for upper teeth and buccal for lower teeth. Description of the type (BUX 67 - 2) (Fig. 31 a). Te type specimen corresponds to an M 2 that displays a long and thin buccolingual anteroloph, in the extension of a thick and high preprotocrista, swollen at is mesial extremity in an anterostyle. Te bulged paracone and metacone are similar in size. Te buccal part of the protoloph is straight, until its low connection to the paraconule. Te latter is bulged and protruding mesially from the protoloph. Lingually to the paraconule, the very thin and low lingual part of the protoloph weakly connects the apex of the protocone. A thin postparacrista and a thicker premetacrista are present and descend towards but do not fuse with the bulged mesostyle. Te latter prolongs lingually in a buccal mesoloph, for which the distal end faces but does not fuse with the metaconule. Te buccal metalophule II is strong, first parallel to the buccal protoloph, then turns mesially; it joins a high distolingually to distobuccally stretched metaconule, which reaches the center of the tooth. Several extra-ridges descend mesially from the buccal protoloph and the metalophule II, around the mesiobuccal extremity of the metaconule, and on the buccal flank of the association of the protocone and endoloph. Te metalophule I is not distinct among these extraridges. Te hypocone is nearly equal in size to the protocone; both cusps are a little stretched, mesiodistally for the protocone and mesiolingual to distobuccal for the hypocone. Tis stretching of the protocone and hypocone prolongs in their anterior and posterior arms. Te metaconule is stretched and prolonged mesiobuccally to distolingually to join the hypocone via a high lingual metaloph. Te postprotocrista and prehypocrista are short and linked by a thick and high endoloph. Te posteroloph is short and lowly connected to a short post-hypocrista; it begins lingually in a bulged posteroconule, and ends at the base of the metacone, the posteroflexus being opened buccally. Upper teeth. Te teeth of P. alsaticus are unilaterally and weakly hypsodont. Tereby, the occlusal surfaces vary with wear, but not strongly. Te cusps are bulged; the conules and associated ridges are sharp on unworn teeth, but blunt and rounded on worn teeth. Te breaks along the lophs as well as the extra-ridges and granules sometimes disappear with wear. DP 4 (Fig. 29 a to d). Eight teeth correspond to DP 4 in our sample; among them six only are referred to this species, and the two largest (CSBX 2 and 3) to? Protadelomys cf. alsaticus morphotype 3. In all cases, DP 4 are smaller than P 4 (Fig. 27 A; Table 4 A). Teir contour is triangular due to the presence of a small hypocone. Te parastyle is present and the anteroloph relatively long and low. Te anteroloph links with the preprotocrista. Te anterostyle is not clearly distinct, due to wear. Te paraconule is smaller than the metaconule and slightly protruding from the protoloph, the latter being weakly linked to the preprotocrista. Te mesostyle is slightly more buccal than the paracone and metacone; it is closer to the metacone and is separated or connected to a short mesoloph. Te metalophule I is sometimes absent, often present but short, low, discontinuous, directed or connected to the mediobuccal crest of the metaconule. Te metaconule is strong, isolated from the hypocone, and weakly connected to the postprotocristid. Te hypocone is conical and much smaller than the protocone; it is lingual when compared to the protocone. Te endoloph is very short and high. Te sinus is absent or barely marked. Te posteroloph is generally short and swollen at the level of the posteroconule. Te hypocone – posteroloph connection is either low and short, or weak or absent. P 4 (Fig. 29 e to l). Tirteen teeth correspond to P 4. On a length x width diagram (Fig. 27), two groups are identifiable. Te first gathers the longest and narrowest teeth (CSBX 5, 6, 7, 8, 9), and the second the shortest and longest ones. However, their morphological variation is partly independent of this grouping. Moreover, their lengths and widths show a coefficient of variation v compatible with a single population (Table 4 A) (Legendre & Vianey-Liaud, 1986). Among the largest P 4, CSBX 8 is morphologically simpler and can be referred to? Protadelomys cf. alsaticus morphotype 1; CSBX 12 displays a sharp and strong preparacrista and a mesoflexus closed buccally, it can be referred to? Protadelomys cf. alsaticus morphotype 3. All the other P 4 are attributed to? Protadelomys alsaticus. All P 4 are triangular with a reduced hypocone (12 / 13). Only CSBX 9 displays a hypocone nearly as strong as the protocone; it is more squared, with parallel anteroloph and posteroloph but its other features are similar to those of the other P 4 (Fig. 29 i). Te endoloph is very short to absent, and the sinus is generally absent or barely marked. Te paracone is isolated (9 / 13) or lowly connected to the paraconule (4 / 13). Te anteroloph is very short and slender. Te paraconule is generally indistinct from the anterostyle, as a strong mesiolingual conule, lower than the paracone and protocone. Te mesostyle prolongs in a mesoloph; on CSBX 6 (Fig. 29 f) the buccal openings of the mesoflexus, on either sides of the mesostyle, are blocked by short ectocingulids. A premetacrista is generally present. Te metalophule I is short and weak. Te metaconule is strong, isolated from the lingual cusp on the less worn P 4 and, on the others, it is connected either to the postprotocrista by a very low ridge on CSBX 7, 9, 12, or to the hypocone (CSBX 13, 29) or to the posteroloph (CSBX 28). Te strong bulbous metaconule bears a short mesiobuccal ridge, ending free or connected to the metalophule I. Tere are more or less numerous low extra-ridges or granules in the mesoflexus. M 1 (Fig. 30) and M 2 (Fig. 31). If M 1 and M 2 display a strong morphological variability, a common pattern is recognizable. All these teeth are pentalophodont, with a short mesoloph. Te paracone and metacone are salient; the protocone and hypocone are sharp on weakly worn teeth and blunt when worn. Te protocone is stretched mesiodistally with its pre- and post-protocristae. Te hypocone is smaller than the protocone, with lower and weaker arms. Mesiodistally, the endoloph is short, thick, and high but a little lower than the protocone and hypocone summits (as seen on lingual views of unworn teeth; Fig. 30 a 2, b 2, c 3, f 2). Te sinus is barely marked or absent. Te paraconule is protruding mesially from the protoloph; it is rounded or a little stretched distomesially, and generally nearly as strong as the metaconule. Te metaconule is generally more bulbous than the paraconule, except on the holotype where it is stretched obliquely. Te connection between the protoloph and the center of the protocone, as well as the connection between the metaconule and the prehypocrista, are generally weak and very low, except on the holotype. Te buccal metaloph is generally absent, the metaconule being separated from the hypocone. Te main variation involves the development of the parastyle and anterostyle, the shape and thickness of the mesostyle, the length and development of the mesoloph, the relationships of the latter with the mediobuccal crest of the metaconule, the development of metalophule I, the connection between the metaconule and the posteroconule, the extra-ridges and granules, and the enamel ornamentation. Te parastyle is often more swollen on M 1 than on M 2, but not always. It generally fuses with the anteroloph; except on some unworn specimens like CSBX 67: Fig. 30 b, CH 41: Fig. 30 c, BUX 67 – 28, and BUX 67 – 54: Fig. 30 h. Te anteroloph is higher than the posteroloph and located between the parastyle and the anterostyle. Te latter is generally swollen and connected to the end of the preprotocrista, which is more or less oblique linguo-mesially. On BUX 67 – 61: Fig. 30 f, the preprotocrista is slightly mesiodistally oriented. Te anteroloph is more or less crenulated, with very short mesiodistal wrinkles. One of these wrinkles sometimes joins an extra-ridge from the protoloph. Tis extra-ridge can be present, absent, or duplicated and parallel to the paraconule. Te distal flank of the paracone rarely shows a short postparacrista (M 1: BUX 67 – 61: Fig; 30 f, BUX 67 – 54: Fig. 30 h CH 49, Fig. 30 l; M 2: BUX 67 – 44: Fig. 31 e, BUX 67 – 2, BUX 67 - 56: Fig; 31 I, CSBX 36). Te premetacrista is thick and always present. Te metalophule II is short and rarely connected to the metaconule. Te metalophule I is absent or reduced to weak extraridges. Te surface of the mesostyle can be underlined by a hawkshape ridge (BUX 67 – 59, 64), which disappears with wear in a thick mesostyle. Short mesiodistal ectocinguli rarely frame the mesostyle (Fig. 30 d). When present, the granules and short low extra-ridges are located in the mesoflexus, from the base of the protocone to the mesoloph and metaconule. Te mediobuccal crest of the metaconule is generally weak, and sometimes directed towards the lingual extremity of the mesoloph, without merging with it. BUX 67 – 70 departs from other molars by its more triangular shape. It probably corresponds to an M 1, as the posteroloph and the hypocone are not really reduced. Te preprotocrista is relatively short and the anterostyle small. Te parastyle is not salient. Te protruding paraconule is small. All the other features are close to those described above. On the M 2 CSBX 30, the post-hypocrista is absent and the posteroconule is linked to a short lingual part of the metaloph, which descends from the hypocone; the posteroloph is reduced to a short crest. M 3. (Fig. 32). Most of the M 3 are unworn or weakly worn (17 / 21); they display a wide range of size variation (Fig. 27 A), but their lengths and widths show a coefficient of variation v compatible with a single population (Table 4 A) (Legendre & Vianey-Liaud, 1986). Among the three biggest teeth, BUX 67 – 27 could belong to? Protadelomys cf. alsaticus morphotype 3. We have not found singularities for the two other large M 3 (BUX 67 – 85, 86), which could justify to exclude an attribution to? Protadelomys alsaticus. Among the smallest specimens, BUX 67 – 88 and BUX 67 – 87 are referred to? Protadelomys cf. alsaticus morphotype 2. Te remaining 18 specimens display a long anteroloph and a distinct parastyle. Te paracone represents the higher cusp. Te metacone is salient from the buccal ridge it forms with the mesostylar area and the metalophule II. Te paraconule is small and protruding. Te development of the metaconule is variable; it is salient on unworn teeth and otherwise usually weaker than the paraconule, but not always (BUX 67 - 84). Te central basin is wide and filled with extra-ridges as strong as in the M 1 – 2, in the area close to the mesostyle and metaconule. Tese extra-ridges are more or less numerous. Te distal part of the teeth is highly variable. Te metalophule II is short, connected to the metacone and oriented distomesially. It can be separated from the metacone or absent, with the metacone connecting with the posteroloph. Te hypocone, more or less strongly reduced, is more buccal than the protocone. Te posteroloph always joins the post-hypocrista. Te sinus is often present but weakly marked; it rarely communicates with the opposite synclines. Wrinkles can be present on the lingual slope of the crown. Lower teeth. (Figs. 33, 34, 35, 36, 37) six specimens are considered as dp 4 of? Protadelomys alsaticus. Te size distribution (Fig. 27 B; Table 4 A) is large, dp 4 being the smallest, then p 4 smaller than molars, the m 2 being slightly larger than the m 1. Te crowns of all the teeth are as high buccally as lingually, but their lingual flank is relatively flat whereas the buccal flanks of the protoconid and hypoconid are bulged. Wrinkles, weak ridges or granules are variably present in the basins. dp 4. (Fig. 33 a to f) Te metaconid is the highest and mesial-most cusp. It separates from the protoconid, which is smaller. Te anteroconid is absent; a weak and short mesiobuccal ridge (preprotocristid) is only visible on two teeth (CSBX 50: Fig. 33 f, and BW 56 – 55; Fig. 33 e). A short buccal metalophulid I is only visible on CH 55. Te metaconid prolongs distally in a high postmetacristid. A small mesostylid is isolated at the distal extremity of this crest on two specimens (CSBX 50 and BW 56 – 54). All specimens display a low mesiodistal extra-ridge, descending from the metaconid to the mesoflexid. Tis extra-ridge is high on CSBX 43 (Fig. 33 a) less on the others, discontinuous on CSBX 50, CH 55 and BW 56 – 55, and almost erased on BW 56 – 54. Tere are a few granules and / or low ridges in the basins (meso- and posteroflexids). Te postprotocristid is nearly mesiodistal and connected to the mesoconid; the latter is a little swollen and the ectolophid can be more lingually located, after making an angle distal to the mesoconid (CSBX 43). Tis ectolophid lowers between the mesoconid and the short prehypocristid. Te entoconid is salient and isolated from the postmetacristid – mesostylid. Te entolophid is complete and very low (6 / 9), joining the extremity of the short prehypocristid; it is incomplete on BUX 67 – 100 and 104. On one specimen only (CSBX 43), the lingual half of the entolophid is distally directed towards the posterolophid. Te hypoconulid is swollen and connected to a long post-hypocristid and to a short posterolophid. Te latter ends at the level of the distal flank of the entoconid, and closes the posteroflexid lingually. p 4. (Fig. 33 g to m) Te posterior root of p 4 is vertical and strong. Te length between the entolophid and the mesial border is only a little longer than to the distal bor- der; it is shorter than in the other Protadelomys species. Te metaconid is the highest and mesial-most cusp, and extends in a long lingual postmetacristid. Te metaconid separates from the protoconid, which is smaller and weaker. Te protoconid is usually low and less distinct than that of dp 4; it is rarely well distinct from the metaconid (CH 26, BW 56 – 57, and BUX 67 – 97). Te anteroconid is absent; there is only a weak short mesiobuccal ridge on BW 56 – 55, and a shallow anterosinus on BUX 67 – 103 (Fig. 33 k). Te wrinkling of the enamel is stronger on the latter than on the other eight p 4. Te mesiodistal ridge is usually low, continuous or more or less interrupted. Te granules and extra-ridges are rare. BUX 67 – 103 and 107 are the most ornamented p 4. Te mesoconid is weak, the mesial ectolophid is indistinct from the postprotocristid – mesoconid, and the distal ectolophid is short and broken at the level of the prehypocristid, as seen on unworn or weakly worn teeth (CH 26: Fig. 33 g; BW 59 - 57: Fig. 33 h). An ectostylid is present buccally at the base of the sinusid on BUX 67 – 101 and 103. Te sinusid is asymmetrical and pinched. Te entolophid is low and rarely continuous (BUX 67 – 103 and 105), directed to or connected to the short prehypocristid. Te hypoconulid is not well individualized; the posterolophid ends at the base of the entoconid. m 1 (Fig. 34) - m 2. (Fig. 35). Te two loci can be distinguished by a set of morphological characters. Te hypoconid is as strong as the protoconid, and slightly placed more buccally on m 1 than on m 2. Te distal root is generally more vertical on m 1 than on m 2, where it is oblique posteriorly and a little twisted buccally. On m 1, the metaconid – protoconid width is lower than the entoconid – hypoconid width, the anteroconid is usually less stretched and shorter, and the anterolophid is most often absent. For both m 1 and m 2, the morphological variation is important. We did not identify clear-cut groupings among the molars, but by combining size variation and a few discrete features, a characterization of three different tooth morphotypes was made possible (see below). On all m 1 and m 2, the metaconid is the higher cusp, followed lingually by a high and strong postmetacristid descending more or less steeply towards the lingual opening of the mesoflexid. Te mesostylid is absent on the m 1, except on CSBX 48, and present on a few m 2 as a swelling at the distal end of the postmetacristid (BSK-A 28, BUX 67 – 110: Fig. 35 h, 127: Fig. 35 d, 132,? 134, BUX 68 – 44: Fig. 35 k) or as a separated small cuspid (BUX 67 – 116, CSBX 46). Te valley between the postmetacristid and the entoconid is generally obtuse but it appears more constricted, acute, on BUX 67 – 136, 137, CSBX 47, and BUX 68 – 33, or even reduced when the mesostylid is present. An anterolophid is present, as a short lingual prolongation of the anteroconid, on m 1 BUX 136 only. On m 2, the anteroconid is generally slightly bulbous, more or less stretched buccolingually. It disappears in the slender anterolophid on BUX 68 – 44 and 46, BUX 67 – 112: Fig. 35 f, 117, 118, and 141. On BUX 67 – 141, the anterolophid is longer than on the other m 2, whereas the thick lingual metalophulid I is not mesial and runs obliquely (mesiobuccally) from the top of the metaconid to meet the transverse buccal metalophulid I. Te complete metalophulid I usually closes the anteroflexid distally (m 1: BUX 67 – 25, CSBX 67 – 52; m 2: BUX 67 – 110: Fig. 35 h, 116, 119: Fig. 35 i, 120) at least at its base; but sometimes it is open distally when the connection does not exist (m 1: BUX 68 – 33, 39; CSBX 42, BUX 67 – 11; m 2: BSK-A 21, A 28; BUX 67 – 15, 112: Fig. 35 f, 118, 121, 139; CH 13; CSBX 49). Te anteroflexid is reduced when the lingual metalophulid I is mesial, as a premetacristid (m 2: BUX 67 – 19 and 133). Te anteroflexid is slightly opened buccally on weakly worn teeth, but a low buccal elevation of enamel is present on its floor. If the buccal half of the metalophulid I always retains the same cuspid arrangement and buccolingual orientation, a strong variation is observed on the lingual metalophulid I. Te buccal metalophulid I always starts from the apex of the protoconid to run linguobuccally towards the midwidth of the crown. Te main variation occurs at the connection with the lingual metalophulid I. Tey generally separates with a mesiodistal narrow groove, more or less wide or deep. In these cases, the lingual metalophulid I can be short and regularly thin (e. g., BUX 67 - 33: Fig. 35 l) or thick (e. g., 130: Fig. 35 c, 132: Fig. 35 b, 133, 141; CH 22), or bulbous at its buccal end (e. g., BUX 68 – 39; BUX 67 – 129). Te lingual and buccal parts of the metalophulid I can, in some cases (CSBX 40, 44, 46, 52; BUX 67 - 113, 132, 140), be strongly connected. Te lingual metalophulid I is usually mesial, and occupies the position of the premetacristid (e. g., m 1: BUX 68 – 39; BSK-A 24, 25, 27; BUX 67 – 25, 114, 128, 135, 136, CSBX 48; m 2: BUX 67 – 110, 112, 113, 116, 117, 120, 133, 140; BUX 98 – 44; BSK-A 21, A 28, A 29; CSBX 40, 41, 49, 53). It can be slightly displaced or angled distally (m 2: BUX 67 - 11, 15, 115, 121,132, 134; BSK-A 26; CSBX 42, 46, 47, 54), or splitted in two branches more or less separated (e. g., m 1: BSK-A 22; BUX 67 – 5, 19; BW 56 – 59; CH 13, 17, 24; CSBX 44, 52, 56; m 2: BUX 67 – 127, 130). On BUX 67 – 141, the strong lingual metalophulid I is transversely in line with the buccal metalophulid I. When distinct from the buccal metalophulid I, the lingual metalophulid I can be long (half-width of the metalophulid I length) or short (e. g., m 1: BUX 67 – 115, 123; CH 22; m 2: CSBX 54; BUX 67 – 115). Te overall shape of the lingual and buccal metalophulids I makes an angle open mesially (e. g., BUX 67 – 124, 126, 132, 140; CSBX 46), but it can be (See figure on next page.) Fig. 34 Lower m 1 of? Protadelomys alsaticus Hartenberger, 1969 from Bouxwiller (Bas-Rhin). a BUX 67 - 136, right m 1; a 1, occlusal view; a 2, buccal aspect; a 3, lingual aspect. b CSBX 52, left m 1; b 1, occlusal view; b 2, buccal aspect. c: BUX 67 - 135, left m 1; c 1, occlusal view; c 2, buccal aspect; c 3, lingual aspect. d BUX 68 - 39, right m 1; d 1, buccal aspect; d 2, lingual aspect; d 3, occlusal view. e BUX 67 - 5, left m 1; e 1, occlusal view; e 2, buccal aspect; e 3, lingual aspect. f BUX 68 - 33, right m 1; f 1, occlusal view; f 2, buccal aspect. g BUX 67 - 128, left m 1; g 1, buccal aspect,; g 2, occlusal view; g 3, lingual aspect. h CH 22, left m 1; h 1, occlusal view; h 2, buccal aspect. i BSK-A 24, left p 4; i 1, occlusal view; i 2, buccal aspect. j: BUX 67 - 111, right m 1; j 1, occlusal view; j 2, buccal aspect; j 3, lingual aspect. k: BUX 67 - 114, right m 1; k 1, occlusal view; k 2, buccal aspect; k 3, lingual aspect. l: BUX 67 - 115, right m 1; l 1, occlusal view; l 2, buccal aspect. Scale bar = 1 mm strictly transverse (e. g., BUX 67 – 113, 131, CSBX 44, 52). hypoconulid), and even ridges (posterolophid and extra- A lower and generally weaker ridge descends from the ridges), which are all thick and low. Te mesial part of apex of the metaconid, distally to the main lingual metal- the ectolophid is mesiodistally oriented and very short ophulid I (CSBX 41, 47, 48; BSK-A 21, A 29; BUX 67 - 127, and weak; its distal part is directed buccolingually to con- 128, 130, 135, 136, 140; CH 17, 22). It is generally slen- nect the postmesoconid, and is linked to the extremity of der, weak, and short, but can also be absent. In few cases, the prehypocristid. In only a few cases (BUX 67 – 19 and the connection of this ridge with the buccal metalophulid CSBX 52), the mesolophid is facing the lingual flank of I occurs through this distal lingual metalophulid I. It is the mesoconid, and is reduced to a spur. Te other molars the case for instance on BUX 67 – 25,123, where the met- do not show a true mesolophid. A postmesolophid-like alophulid I is complete and angles medially; or on BUX structure is sometimes visible (BUX 67 – 5, 114, 115, 67 – 116: Fig. 35 a, 136, 137, and BUX 68 – 44: Fig. 35 k, 123, 124, 125, 126, 135: Fig. 34 c, 136, 137, BSK-A 27, BW where it is straight and transverse. 56 – 59, CH 17, 22, 24; CSBX 47), the postmesoconid spur, Te postprotocristid is strong, thick distally and which is more or less bulged or elongated obliquely. Te oblique mesiobuccal to distolingual; it is always longer mesoconid and postmesoconid are, in some cases, undisthan the prehypocristid. tinguishable (CSBX 47, 54; CH 24; BUX 67 – 116). Te ectolophid area, defined from the distal end of the Te entolophid can be almost transverse or slightly postprotocristid to the mesial end of the prehypocris- curved distally. It usually joins the mesial end of the pretid, appears variable. Te Figure 36 illustrates this vari- hypocristid, together with the low distal ectolophid. It ation with schematized drawings of the buccal area of lowers at the level of this junction, sometimes interrupts. the m 1 – m 2 from Bouxwiller. Te mesoconid can occupy On BUX 67 – 133, the entolophid is not continuous and most of the ectolophid (BUX 67 – 19, 111; BUX 68 – 33). formed only by small isolated oblique ridges. Te buccal On both sides of this mesoconid, the ectolophid is short. junction of the entolophid often occurs either more lin- Its mesial part is generally in the continuity of the post- gually, to the postmesoconid or the postmesoconid spur; protocristid, and hardly distinct. Its distal part is lower, or more distally, to the anterolingual wall of the hypocwhich interrupts on weakly worn or unworn teeth. How- one (BUX 67 – 130, 132; BUX 68 – 39; BW 56 – 59; CSBX ever, the path of this ectolophid + mesoconid varies. 54). Te entolophid usually bears an entoconulid, which Te postprotocristid is thick and frequently swollen at is however frequently worn and difficult to identify; it its end in a “ premesoconid ” - like structure (BUX 67 – 11, is only visible on BUX 67 – 5, 19, 25, BW 56 – 59, CH 24, 25, 141; BSK-A 26, A 28). Tis structure prolongs in a CSBX 40, 47, 56. One, rarely two, distomesial spur (s) or lingual spur of variable size (BUX 67 – 25, 110, 111, 112, ridge (s) frequently descend (s) from the entoconulid in 123, 130, 132, 136; CH 22, 24; CSBX 40, 41, 44, 46, 47, the central basin. 48), two lingual spurs (BUX 67 – 132, 140), or one buccal A few extra-ridges, wrinkles or granules are present in spur (CH 13). On the m 1 BW 56 - 59, this spur is long at the central basin, but often erased by wear. Some of these the end of the postprotocristid but does not present any are organized in oblique ridges from the distal extremlingual swelling. It is indistinct or absent on the m 2 BUX ity of the lingual metalophulid I, or / and from the lingual 67 – 116, 117, and 129. On worn teeth, the swelling or extremity of the buccal metalophulid I to the basin floor. the spur of the premesoconid are not visible (BSK-A 27: Tere are two well-distinct oblique extra-ridges (BSK Fig. 36, BSK-A 29 BUX 67 – 23, 120, 139). Te short mesial A 22, 24), one on BUX 67 – 19, 128, 137, CSBX 40, 47, 52, part of the ectolophid is higher than the distal one, and 56, and CH 22, three on BUX 67 – 114, 136, BUX 68 – 36, rarely distinct from the junction premesoconid – meso- 39, and BW 56 – 59. Te extra-ridges are numerous on conid. Te shape of the mesoconid varies with wear: BUX 67 – 114, 137, and BW 56 – 59; CSBX 52. it can be bulbous, reduced or stretched. BUX 67 – 115 m 3. (Fig. 37). Among the measured m 3, three are much display a peculiar morphology, with bulbous cuspids, larger than the others are, and are described below as? P. conids (mesoconid, premesoconid at the extremity of the cf. alsaticus morphotype 1 and? P. cf. alsaticus morphopostprotocristid, postmesoconid spur, anteroconid, and type 3 (Table 4 B). If another teeth is much shorter (BUX (See figure on next page.) Fig. 35 Lower m 2 of? Protadelomys alsaticus Hartenberger, 1969 from Bouxwiller (Bas-Rhin). a BUX 67 - 116, right m 2; a 1, occlusal view; a 2, buccal aspect; a 3, lingual aspect. b BUX 67 - 132, left m 2; b 1, occlusal view; b 2, buccal aspect; b 3, lingual aspect. c: BUX 67 - 130, right m 2; c 1, occlusal view; c 2, buccal aspect; c 3, lingual aspect. d BUX 67 - 127, left m 2; d 1, occlusal view; d 2, buccal aspect; d 3, lingual aspect. e CSBX 49, right m 2; e 1, occlusal view; e 2, buccal aspect; e 3, lingual aspect. f BUX 67 - 112, right m 2; f 1, buccal aspect; f 2, lingual aspect; f 3, occlusal view. g CSBX 56, left m 2; g 1, occlusal view; g 2, buccal aspect; g 3, lingual aspect. h BUX 67 - 110, left m 2; h 1, occlusal view; h 2, buccal aspect; h 3, lingual aspect. i BUX 67 - 119, right m 2; i 1, buccal aspect; i 2, lingual aspect; i 3, occlusal view. j: BUX 67 - 141, right m 2; j 1, buccal aspect; j 2, occlusal view; j 3, lingual aspect. k: BUX 68 - 44, left m 2; k 1, occlusal view; k 2, buccal aspect; k 3, lingual aspect. l: BUX 67 - 33, left m 2; l 1, occlusal view; l 2, buccal aspect; i 3, lingual aspect. m: BUX 67 - 133, right m 2; m 1, occlusal view; m 2, buccal aspect; m 3, lingual aspect. n: BUX 67 - 140, left m 2, occlusal view. Scale bar = 1 mm 67 – 155), its features seem to enter the whole variability. BUX 68 – 37, is here referred to? P. cf. alsaticus morphotype 2. It has a relatively short oblique postprotocristid, a low mesial ectolophid; the mesoconid is strong and prolonged in a thick postmesoconid. Te distal ectolophid is higher than the mesial one; the prehypocristid is thin and short. It differs from BUX 67 – 155 and from all the other specimens in the absence of a thick postprotocristid and of a premesoconid spur or ridge. In all m 3, the lingual metalophulid is mesial, except on BUX 67 – 160, where it is distally oriented. Te occurrence and strength of the ectomesolophid and of a more or less mesiodistal ridge from the lingual metalophulid are variable among all the m 3. Te ectomesolophid is long on BSK-A 25, BUX 67 – 151, 153, 159, 165, and 168. A mesostylid, more or less distinct from the postmetacristid, occurs on CSBX 45, BUX 67 – 14, 151, 152, 157, 159, 160, and 165. On BSK-A 29 and BUX 67 – 168, the mesostylid prolongs buccally in a short lingual mesolophid, the metaconid appears more massive, thicker at its base, than on the other specimens. We observed the same main variation along the postprotocristid – ectolophid area, but less pronounced than on m 1 – 2. Te post-hypocristid is relatively short up to the hypoconulid area. Te hypoconulid is weakly bulged, single or often duplicated (e. g., BUX 68 – 37; 67 – 53, 151, 154, 155, 158, 164, 168). Te posterolophid is short, generally ending at the distal base of the entoconid.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF98DF06C73943E1FEB1FA79.taxon	discussion	Remarks and comparisons Even if? P. alsaticus has its postprotocristid thick, long and oblique with its distal end often swollen with a premesoconid spur, like in P. lugdunensis, morphological differences involving the lower teeth are observed, together with the differential development of extra-ridges and granules: in P. lugdunensis, the cusps are more bulged, the entolophid is more continuous, the postprotocristid is fused to the mesoconid area (no distinct pre, meso and postmesoconid elements).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF8ADF05C73943E6FA36FC79.taxon	materials_examined	Material and measurements (Additional file 4: S 4, Table 4 B). We refer two P 4, one M 1, two M 2, two dp 4, one p 4, two m 1, two m 2, one m 3, to this morphotype.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF8ADF05C73943E6FA36FC79.taxon	description	Description Upper teeth P 4. (Fig. 38 a) One P 4 (CSBX 8) displays a simpler occlusal morphology than other specimens, and cusps more bulbous. Te paracone and metacone are very close to one another, and the mesoflexus is narrow. Te mesostyle is small, adjacent to the metacone, and the mesoloph is absent. Another P 4 (CSBX 29) could be close to CSBX 8, with its bulbous cusps and small hypocone, but it is too damaged, especially in the area between the paracone and metacone. M 1. (Fig. 38 b) One tooth (CSBX 35) has been questionably refered to an M 2 of? P. alsaticus (Hartenberger, 1969: pl. 2, Fig. 7). Tis weakly worn M 1 displays an anterosyncline wider than that of the other M 1. Te parastyle is strong, slightly separated from the anteroloph, and weakly connected to a sloping preparacrista. Te anteroloph is weakly worn, whereas the parastyle is unworn. Te anterostyle is strong, connected to both the preprotocrista and the anteroloph. Te endoloph is short and thick, nearly as high as the lingual main cusps. Te hypocone is slightly smaller than the protocone, its anterior and posterior arms being short. Te parastyle presents one additional ridge connected to the anteroloph in the anterosyncline. Te paracone is slightly swollen. Te buccal part of the rectilinear protoloph is as high as the paracone, and its lingual part is lower. Te protoloph connects to the protruding paraconule and to the middle of the protocone (protocrista). Te paraconule is mesial to the protoloph and slightly less swollen than the metaconule. Te paracone is framed by the pre- and post-paracristae. Te latter is strong, and interrupted anterior to the mesostyle. Te mesostyle prolongs in a strong mesoloph, joining a low cusp (mediobuccal) against the metaconule. A short ectocingulum is present on the distal flank of the mesostyle. Te premetacrista is strong and the metacone a little thicker than the paracone. Te metalophule II is high and the metalophule I is low, descending mesially from the latter to the base of the metaconule. Te connection between the metalophule II and the metaconule is low as is the one between the latter and the mid-hypocone. Tere is a strong posteroconule, lowly attached to the metaconule and to the mesiodistal post-hypocrista. Te sinus forms only a very shallow depression below the endoloph. Some rare very low ridges, present on the floor of the flexi, connect the main structures. (See figure on next page.) Fig. 36 Schematic representations of the variations of the outer wall components of the lower molars of? Protadelomys alsaticus Hartenberger, 1969 from Bouxwiller (Bas-Rhin). Color caption: the colors represent different cuspids, cristids or lophids: 1: Brown = protoconid; 2: buccal part of the metalophulid I = pale violet; 3: postprotocristid = brown like the protoconid; 4: premesoconid swelling (spur or ridge) = brown, at the distal end of the postprotocristid; 5: mesial ectolophid = green; 6: mesoconid = blue; 7: ectomesolophid = blue; 8: postmesoconid ridge and swelling = ochre; 9 distal ectolophid = Green; 10: prehypocristid = black; 11: hypoconid = black; 12: entolophid = black; 13: poshypocristid = black M 2. (Fig. 38 c) Among the largest M 2, BUX 67 - 37 and 67 – 77 are referred to this morphotype. Tey show distinct and swollen conules. Te metaconule is simple. If the thick buccal metalophule II is transverse on BUX 67 – 37, it thickens lingually and curves distally on BUX 67 – 77 to join the posteroconule. Te mesostyle and the buccal mesoloph are present but their width seems to vary: on BUX 67 – 37, it is thicker and longer and reaches the metaconule. LoWer teeth dp 4. (Fig. 38 d, e) Two dp 4 differs from the others by having a transverse metalophulid I, as well as a short anterolophid closing a small anteroflexid. On BUX 67 – 96, the mesiodistal ridge is barely marked, as is the buccal half of the entolophid. Te lingual part of the latter is short and distally oriented towards the posterolophid. CH 25 is much worn with a damaged mesial flank, but the “ trigonid ” area is preserved and similar to that of BUX 67 – 96. p 4. (Fig. 38 f) On CH 18, the metaconid is strong and prolonged buccally in a high and short anterolophid up to the preprotocristid. A short transverse metalophulid I runs parallel to the anterolophid, both being separated by a minute anteroflexid. Te short postmetacristid descends abruptly towards the lingual opening of the mesoflexid, mesially to the entoconid, which is massive. Tis lingual opening is V shaped and the vertical surface of enamel is concave at the junction of the lingual flanks of metaconid and entoconid. Tis is due to the bulging of the buccal flanks of the metaconid + postmetacristid, and the bulging of the buccal flanks of the entoconid. One thick mesiodistal ridge runs from the metaconid to the premesoconid spur; lingual to it, there is an additional parallel crest, more discontinuous. Tree short extraridges fill the space between the main ones. Te ectolophid makes an angle behind the mesoconid; the latter is slightly stretched mesiolingual to bucco-distal. A short and strong ectomesolophid curves and its buccal end connects a hypoconid spur, which is mesiobuccal. Te lophids, ridges, and conids appear thick when worn. m 1 – 2. Te peculiar shape of the buccal wall of the p 4 and m 3 is not so easy to distinguish on the m 1 – 2: the recess between the bulging flanks of the metaconid and the entoconid is not strongly marked, and varies with wear. Among the largest m 2, BUX 67 – 138, 139: Fig. 38 i, 142, and CSBX 53 show these features; BUX 67 – 121 does not. We found this character on the m 1 BUX 67 – 125 and 137. BUX 67 – 125 has a short and swollen anteroconid; the lingual metalophulid I is mesial then turns distally in a short ridge, which meets the distomesial hook located at the end of the transverse buccal metalophulid I. Te weak and low additional lingual metalophulid I makes a long mesiodistal ridge up to the center of the basin, where it meets a few low and slender extra-ridges. Another extra-ridge runs on the buccal flank of the metaconid lingually to this additional metalophulid I. A short premesoconid spur is present. Te mesoconid is a little stretched obliquely (mesiolingual to bucco-distal); it bears a barely marked ectomesolophid and a weak mesiobuccal spur of the hypoconid. Its distal end is linked to a thick postmesoconid. Te distal ectolophid is low and very short, and this is especially clear on unworn and weakly worn teeth. Te entolophid is almost straight and connected to the prehypocristid. Te post-hypocristid is short, the hypoconulid strong, and the posterolophid displays two successive bulges. Te occlusal face of BUX- 137 is damaged and all occlusal structures are worn. BUX 67 – 138 is a pristine m 2, reduced to its enamel cap (Fig. 38 h). At its lingual end, the stretched anteroconid connects with a thin discontinuous mesiodistal ridge running towards the center of the basin. Te lingual metalophulid I is mesial and short as a premetacristid. Te buccal metalophulid I is longer, being transversely oriented at its most buccal part then turning slightly backwards. An additional lingual metalophulid I is thin and low, like the extra-ridges located in the basin. Te posprotocristid ends in a premesoconid spur. It strongly connects to the mesoconid, which is stretched buccomesial to linguodistal. A strong ectomesolophid descends mesiodistally along the buccal flank of the mesoconid, while a thin mesiobuccal spur climbs on the hypoconid flank. Two postmesoconid spurs emerge from a unique point at the distal end of the mesoconid. Te distal ectolophid is low, and connects to the short prehypocristid. Te entolophid is almost straight and complete; it is linked to the extremity of the prehypocristid. Te hypoconulid swells at the extremity of the post-hypocristid, and the posterolophid is long, relative to the post-hypocristid length. m 3. (Fig. 38 j) BUX 68 – 23 is the larger m 3 of the theridomorph assemblage from Bouxwiller. Its morphology is relatively simple when compared to other m 3, and the extra-ridges are very low and rare. Its metaconid and entoconid are more massive. Te lingual metalophulid I is mesial as it is the case on nearly all the m 3 from Bouxwiller; it is short and its buccal end joins the anterolophid lingual end. Te additional more distal lingual metalophulid I is slender and low; it descends in the direction of the transverse buccal metalophulid, without connecting it. Tere is no mesostylid. Te postprotocristid is long and thick, and connects to the mesoconid by a low mesial part of the ectolophid. Te mesoconid is a little stretched mesiodistally, with two tiny mesial and distal bulges with short lingual spurs. A thick linguodistal short ridge starts in front of the distal bulge. We interpret this ridge as the postmesoconid ridge. Indeed, there is no trace of a lingual part of the entolophid. Te entoconid is free from the posterolophid. Te latter is very short, making a short and thick curve with the hycoconulid and the post-hypocristid. Te hypoconulid is unique and not swollen.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF89DF00C73940F2FE93F99F.taxon	materials_examined	Material and measurements. (cf. Additional file 4: S 4; Table 4 B) We refer six M 1 – M 2, two M 3, one dp 4, three p 4, one m 1, one m 3 to this morphotype.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF89DF00C73940F2FE93F99F.taxon	description	Description Upper teeth M 1. (Fig. 39 a, b, c). Te smaller M 1 of the assemblage (BUX 67 – 78; Fig. 39 a) has a relatively simple occlusal pattern, without wrinkles of the enamel on the outskirts of the crown, only very small and tiny roughnesses. Te parastyle is weakly swollen. Te main cusps and conules, as well as the anterostyle and posteroconule, are bulbous. Te metacone bears a premetacrista. Te mesostyle is strong and placed a little more buccally than the paracone and metacone. Te paraconule prolongs mesially in a short thick ridge; it is a little smaller than the metaconule, which is bulbous and strong. Te mesoloph reaches the center of the mesoflexus but its lingual part is discontinuous and low. Te protoloph is strong only in its buccal part; its lingual part is very low and slender, and connects with the center of the protocone. Te buccal part of the metalophule II is strong; it ends lingually in a distomesial spur, which could represent a vestigial metalophule I. Te lingual metalophule is as weak as the very low distal connection between the metalophule II and the metaconule, and the low distal connection between the metaconule and the center of the hypocone. Te other M 1 (BUX 67 – 48: Fig. 39 c, BUX 67 - 49: Fig. 39 b, BUX 67 - 63: Fig. 39 d = M 1 or M 2; BUX 67 - 78: Fig. 39 a) are small but display the same occlusal pattern, with some variations in the mesoloph length (shorter in BUX 67 – 48) and in its lingual connection: to the metaconule mesial spur (BUX 67 - 48, 49), or free (BUX 67 – 63). Te parastyle is more swollen on BUX 67 – 48 and 49. M 2 (Fig. 39 d, e). Te M 2 show the same features as the M 1, but the connection between the mesoloph and the buccal ridge of the metaconule is stronger on BUX 67 – 26, which also shows a stronger distal spur on the paraconule. Te mesoloph is short and ends free in the mesoflexus on BUX 67 – 41. BUX 67 – 26 shows a short ectocingulum on the distal flank of the mesostyle. BUX 67 – 41 has both postparacrista and premetacrista while BUX 67 – 26 has only the premetacrista, like on the M 1. M 3. BUX 67 – 87: Fig. 39 f, and 88: Fig. 39 g, are small, their occlusal outline is almost circular in shape with rare extra-ridges and small conules. Te metaconid is weakly protrudring from the bucco-distal cingulum. LoWer teeth dp 4. (Fig. 39 h). BUX 67 – 95 is much smaller than the other dp 4. Despite an important mesial wear, it is possible to distinguish a mesial junction between the metaconid and the protoconid (anterolophid), closely followed distally by the metalophulid I. Te mesiodistal ridge is incomplete and low. Te postprotocristid makes a short low lingual spur at its distal extremity. Te mesoconid is barely marked on the short mesiodistal ectolophid. Te latter lowers at the level of its junction with both the prehypocristid and the buccal end of the entolophid. Te hypoconulid is slightly protruding above the prehypocristid and posterolophid. Tere is no distinct extra-ridge or granule, and the enamel is smooth. p 4. (Fig. 39 i, j, k). Te three p 4 are shorter than but almost as wide as those of? Protadelomys alsaticus. Te extra-ridges are absent, apart from a short mesiodistal ridge, which connects to the metaconid (BUX 67 – 98 and BW 56 - 54 bis) or to the protoconid (BUX 67 – 106). Te latter and BUX 67 – 98 show a small mesial notch (not visible for BW 56 - 54 bis); it is limited linguo-mesially by a short cingulid on BUX 67 – 106. molars. On the smallest lower molars previously referred to as P. alsaticus, no peculiar morphological characters can justify an attribution to P. cf. alsaticus morphotype 2. However, we tentatively refer two teeth to this morphotype: one m 1 (BSK-A 27: Fig. 39 l) and one m 3 (BUX 68 – 37: Fig. 39 m).), which are clearly narrower than the other m 3.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF8CDF7EC4834546FB64FD19.taxon	materials_examined	Material and measurements. (Additional file 1: S 4; Table 4 B). We refer two DP 4, one P 4, one M 2, one M 3; one m 1, two m 2, and three m 3 to this morphotype.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF8CDF7EC4834546FB64FD19.taxon	description	Description Upper teeth DP 4. Te larger DP 4 (CSBX 2: (Fig. 40 a).) of the Bouxwiller assemblage displays a more intricate pattern than the other DP 4 (Fig. 40 b). Te paraconule is relatively reduced and aligned with the straight protoloph, while the metaconule is duplicated, as are the mesostyle and the mesoloph. Te metalophule I joins the mediobuccal ridge of the metaconule, together with the main mesoloph. Te outskirts of the crown display irregular wrinkles, thicker than the thin ornamentation of? P. alsaticus. P 4. (Fig. 40 c, d). CSBX 12 and 13 show common features with the P 4 of? P .. alsaticus, but still differs from them in some aspects. Tey show a longer anteroloph with a preparacrista, which is stronger; the connection between the postparacrista, the small mesostyle, and the long premetacrista limits the shallow mesoflexus buccally. Te metalophule I is reduced or almost absent. Te hypocone is almost as large as the protocone. M 2. (Fig. 40 e). BUX 67 – 3 is weakly worn, especially in its medial part. Te extra-ridges are more numerous than on the M 1 BUX 67 – 35 (morphotype 1). Te paracone is bulged and the postparacrista is present. Te protoloph turns a little distally, buccal to the paraconule. Te latter separates from the protoloph by an oblique notch; it is stretched forward, inside the anteroflexus. Te paraconule attaches to the short transverse lingual part of the protoloph, linked at mid-protocone, just mesially to a low protocrista. Te premetacrista, the metacone, and the metalophule II merge into a loph, which is arcuate up to the metaconule level. Te metaconule is complex, which makes it difficult to identify a swollen conule present in this area alongside at least three very close ridges. Te mesostyle is complex, prolonged by a bifurcated mesoloph. A ridge extends from the end of the buccomesial border of the metacone and runs parallel to the mesoloph. Another ridge descends from the metalophule II in the basin; it could constitute a remnant of the metalophule I. Te endoloph is high and thick. Te hypocone is more buccal than the protocone. It separates from a distinct posteroconule by a shallow and narrow slot. Te posteroloph is very short. M 3. (Fig. 40 f). Te unworn M 3 BUX 67 – 27 appears more concave than the other molars, as the cusps and the surrounding bucco-distal ridge or the lingual one are not high or thick. Te paracone is not much prominent, and only slightly higher than the protoloph. Te mesially protruding slender paraconule is oriented towards the anterostyle. Te latter, not well individualized, only represents a swelling at the extremity of the preprotocrista. Te extra-ridges and the granules are numerous on the mesoflexus floor. Te metacone is low, not salient, and located above the bucco-distal crest joining the mesostyle area. Tis metacone is similar in height as the posteroloph, which is short. Te protocone is a little elongated mesiodistally from the preprotocrista to the postprotocrista. A thickening, which actually corresponds to the high and strong endoloph observed on the M 2, prolongs the postparacrista (BUX 67 - 3). A linguobuccal lophid, as high as the paraconule, faces the protocone and reaches the central extra-ridges. Te hypocone is not clearly identifiable, as it is not swollen; it is strongly linked to the postprotocrista. Tere is no trace of a sinus. Te metaconule is lower than the paraconule, and reduces to a thin ridge postwardly oriented. Smooth vertical wrinkles, diverging from its occlusal surface, adorn the lingual surface of the protocone. LoWer teeth m 1 – m 2. (Fig. 40 g, h, i). Te m 1 BW 56 – 59 has numerous extra-ridges and wrinkles. Te elements of the postprotocristid + ectolophid area are in line obliquely, mesiobuccal to distolingual, like on the two m 2 (BUX 67 – 129 and 142). BUX 67 – 129 is little worn, as are their short anteroconid whereas the premetacristid (mesial lingual metalophulid I) is longer than on the m 1 and it bears two long mesiodistal ridges, parallel to the one starting at the lingual end of the buccal metalophulid I. Te postmetacristid is steep and long. On BUX 67 – 142 (Fig. 40 i), which is not too much worn but a little damaged, the premesoconid – mesoconid – postmesoconid elements are obliquely in line with the postprotocristid, like on the m 1. Te ectomesolophid is faint. Te lingual metalophulid I, transverse, is almost in line with the buccal one, without any break. Even worn, it is possible to distinguish numerous low extra-ridges in the central basin and a few wrinkles descending from the postprotocristid and mesoconid area in the sinusid. m 3. (Fig. 40 j, k). BUX 67 – 169 differs from the other large m 3 BUX 68 – 23 (morphotype 1) in its mesial part that is relatively wider than the distal one. It also differs from it in the presence of a mesostylid at the extremity of the postmetacristid closing lingually the mesoflexid, and in the presence of an entolophid. A few wrinkles descend buccally from the postmetacristid and from the mesostylid; some granules are visible in the basin, even if the tooth is worn. Te lingual metalophulid I is mesial (premetacristid) and bears two long low mesiodistal ridges running in the basin, parallel to the short ridge emitted by the lingual end of the transverse buccal metalophulid I. A premesoconid spur is well distinct, directed lingually from the end of the postprotocristid. Te lophid developed from the postprotocristid to the prehypocristid is mesially oblique and then oriented mesiodistally, with the distal part of the ectolophid. Te ectomesolophid is absent (contra on BSK-A 25 where it is well present). Te buccal part of the entolophid connects to the strong postmesoconid ridge and to the prehypocristid. Te lingual entolophid is duplicated and the distal one is slightly directed distally. While worn, the hypoconulid seems to be also duplicated, with a distomesial ridge running towards the buccal entolophid. Te extra-ridges and granules are numerous in the low basins of BSK-A 25.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFF8CDF7EC4834546FB64FD19.taxon	discussion	Remarks and comparisons Te larger lower molars of? Protadelomys cf. alsaticus morphotype 3 are similar in size as the ones of Masillamys parvus fom Messel. Tey share the numerous extra-ridges and the organization of the lingual metalophulid I area. However, the oblique postprotocristid + ectolophid is longer, the sinusid is deeper linguobuccally. Moreover the trigonid of p 4 in this species is better developed than in p 4 of Masillamys parvus, with a distinct protoconid, and the entolophid and extraridges are thicker in the species from Bouxwiller than in M. parvus.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFF2DF7CC73941C1FB85FDD9.taxon	discussion	Remarks Tese rodents have been referred to as Protadelomys (cf.) alsaticus (Marandat et al. 1993) based on the presence of frequent enamel wrinkles and size similarities, which are smaller than that of the teeth of P. lugdunensis.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFF2DF7CC73941C1FB85FDD9.taxon	materials_examined	Material and measurements (Additional file 5: S 5, Fig. 41 Table 5). Most of the teeth, except the two lower dp 4, are slightly larger than the teeth of P. alsaticus from Bouxwiller (Table 4). Te difference is only statistically significant for the m 1 (Table 8). Morphological comparison Upper teeth. (Fig. 42 a to m). Te two DP 4 from Cuzal display some differences with P. alsaticus from Bouxwiller. On the DP 4 from both localities, the protoloph is oriented forward and connects to the junction anteroloph – preprotocrista. Te paraconule is absent in the species from Cuzal whereas it is is always present in P. alsaticus from Bouxwiller. Two mesostyles and mesolophs are present on the two DP 4 like in Bouxwiller. Contrary to P. alsaticus from Bouxwiller, the hypocone is as strong as the protocone, which is more lingual. Te metalophule I is not distinct, while it is on a few DP 4 from Bouxwiller. Te metaconule is not bulged and reduced to extraridges, which join the hypocone and the posteroloph (CUZ 176). In P. alsaticus, the metaconule is strong with no lingual connection or junction with the postprotocrista. Te P 4 from Cuzal are triangular with a reduced hypocone, whereas the hypocone is less reduced in Bouxwiller. Te floor of the mesoflexus of the P 4 is flatter and more wrinkled than in P. alsaticus. On CUZ 188, the anteroloph is very short and weakly swollen at the level of the anterostyle, which joins the preprotocrista, not on CUZ 177 and CUZ 189 Te paracone is slightly smaller than the metacone. A thin postparacrista plunges to the mesostyle area. On CUZ 188 (Fig. 42 d), the buccal protoloph is forked, one part oriented forwards to the anterostyle, the other backwards, directed to the lingual metaloph. Tey are less distinct on the other P 4 and correspond to low extraridges. Te P 4 lack a paraconule. Te mesostyle prolongs in a mesoloph on CUZ 188 and 189, it is duplicated mesially on the former, and interrupted on CUZ 177. A short metalophule I joins the distal mesoloph on CUZ 188. Te metaloph II is thicker and directed towards (CUZ 177) or connected (CUZ 189) to the robust metaconule. Te latter is more (CUZ 188) or less (CUZ 177, 189) fused with the postprotocrista. A small ridge connects the metaconule to the hypocone, which is weak and stretched on CUZ 188. Te upper molars of cf.? P. alsaticus from Cuzal display some variable features. On all of them, as well on the M 1 as on the M 2, the hypocone is only slightly less robust than the protocone. Te smallest M 1 (CUZ 154: Fig. 42 k, and 155: Fig. 42 j) have shorter protocone + pre- and post-protocristae, but the other features vary like in the other M 1 from Cuzal (CUZ 178, 181, 182, 186) and like in? P. alsaticus from the type locality, Bouxwiller, with numerous extraridges. Elsewhere, we noted a few differences with? P. alsaticus from Bouxwiller. Te size difference between the two main lingual cusps, the protocone and hypocone, is more important in typical alsaticus. Te paraconule and metaconule are present but less bulged than in the type population. In the upper molars from Cuzal, the protoloph connects the preprotocrista or the protocone; the buccal metaloph II is nearly always present and connects the hypocone and often both the posteroloph through mesiodistal extraridges (CUZ 156, 178, 182); the extraridges are more numerous in the flexi; a metalophule I is present or sketched (6 / 8). A mediobuccal crest of the metaconule is generally present, merging with various extraridges. A narrow pinched sinus is more frequently present, below the endoloph. Te morphology of upper M 3 (at least CUZ 180) is close to that of the large M 3 from Bouxwiller. Lower teeth. (Fig. 42 n to x). Te lower dp 4 have a narrower pre-lobe, due to the absence of the protoconid and metalophulid I. Te protocristid is low, and an ectostylid limits the base of the sinusid on CUZ 187 (Fig. 42 n). Te entolophid is low and weak. Te posterolophid is absent on CUZ 161 and replaced by a linguodistal notch. Like the dp 4, the p 4 differs from typical? P. alsaticus in the absence of protoconid and in the ratio of the length of the anterior lobe, markedly longer than the posterior lobe. Te anterior is delimited lingually by the metaconid + the postmetacristid and buccally by the mesiodistal protocristid + the postprotocristid + the ectolophid and mesoconid, the posterior lobe extends from the hypoconid, through the post-hypocristid + the hypoconulid + the posterolophid to the entoconid. Unlike typical? without? P. alsaticus, the anterolophid is more often present as a lingual prolongation of the relatively bulbous anteroconid on m 1 and m 2. It is generally distinct from the premetacristid, which is generally absent. It connects to the metaconid flank only on a few worn teeth (m 1, m 2: CUZ 164). Like in typical alsaticus, the anteroflexid is closed distally by a complete (buccolingual) metalophulid I on lower molars. A low anterolophulid can divide the anteroflexid (m 1 – 2: CUZ 168, 169, 170, 175; m 3: CUZ 164). Te path of the metalophulid I is more constant than in typical? without? P. alsaticus. It is complete in all cases, rectilinear from the anterior part of the metaconid to the protoconid, and slightly oblique mesiolingual to distobuccal. Its lingual part is never completely mesial, as it is typically in? without? P. alsaticus. Additional thin ridges (one to three) descend from the lingual metaconid flank to the talonid basin. Te postprotocristid is strong, thick distally and oblique buccomesio-distolingually; it is always longer than the prehypocristid, like in typical? P. alsaticus. Tere is less variation in the ectolophid area, from the distal end of the postprotocristid to the mesial end of the prehypocristid. Te mesoconid occupies most of the ectolophid, which is short. Te direction of the ectolophid mesial part generally makes an angle with that of the postprotocristid. Similarly, its short distal part makes an angle with the prehypocristid; it is slightly lingual than the mesial one. Te mesial and distal parts are similar in height but low, seeming interrupted on unworn teeth. Te premesoconid, mesolophid, and postmesoconid ridges are always distinct, more or less long. At least one ectomesolophid is present (13 / 15 m 1 – 2), but it can be duplicated (CUZ 170). Te entolophid is more or less continuous, connecting the entoconid to the prehypocristid or to the postmesoconid ridge. Two m 3 only are present; their morphology is similar to that of m 1 – 2, but with a reduced posterior lobe. Comparison. Overall, we found more similarities between the species of Cuzal and? P. alsaticus than with the other species of Protadelomys (P. cartieri, P. lugdunensis) and? Protadelomys (? P. maximini and? P. nievesae), notably in its size, the organization of the postprotocristid – ectolophid, the connections between the lophs and the lingual cusps, the alignment of the pre- and post-cristae, and the numerous extra-ridges. Among the differences, some could indicate a more progressive grade than typical? P. alsaticus, like the more robust hypocone, the less bulged conules, and the transverse lophs and metalophulid I that are better defined than in typical alsaticus. However, the evolutionary significance of the absence of the protoconid on p 4 and dp 4 remains unclear, as it is present on p 4 and dp 4 of? P. nievesae,? P. alsaticus, and P. cartieri. In contrast, it is poorly developed or absent on p 4 and dp 4 of? P. maximini and P. lugdunensis. Te weak morphological variation of the lower molars is also noticeable.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFF0DF6EC7394181FA37FDF9.taxon	materials_examined	Holotype. M 2 (or M 1?); SMX 1 - 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 dp 4, wider DP 4 and significantly larger M 1; larger than? P. nievesae. Enamel of the teeth generally ornamented with weak and low extra-ridges and granules. Upper teeth. DP 4 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. P 4 as length as and slightly wider than the DP 4. 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; m 3 with metacone slightly prominent; paraconule protruding and metaconule crestiform. Lower teeth. All the lophids and ridges are thin. On dp 4, anteroconid absent, and anterocingulid variably present; protoconid small, and lower than the metaconid; no strong mesiodistal ridge descending in the mesoflexid. On p 4, 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 dp 4; protoconid absent or indistinct from the metaconid on p 4; 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 DP 4 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 dp 4 with a flat basin; the p 4 with an underdeveloped protoconid; the lower molars with a more discontinuous and low entolophid. Material and measurements. (Additional file 6: S 6; Tables 6, 8; Figs. 43, 44). Upper teeth: Probably due to different taphonomic conditions, the DP 4 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 M 2 from 1.63 × 1. 65 mm to 1.92, to 2.03 mm (i. e., the type) (Table 6 A, Additional file 6: S 6). Only two M 3 (SMX 1 - 291 and SMX 2 - 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, 12). Lower teeth: Fourteen dp 4 are present in the sample from Saint-Maximin. Tey appear slightly larger than the dp 4 of? P. alsaticus from Bouxwiller (Tables 4 A, 6 A). Te size of the p 4 of? P. maximini is only slightly smaller than that of? P. alsaticus from Bouxwiller (Tables 4 A and 6 A). It is larger than that of? P. nievesae from Casa Ramon (Additional file 7: S 7, 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), but they are significantly larger than those of? P. nievesae from Casa Ramon (Table 8).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFF0DF6EC7394181FA37FDF9.taxon	description	Description Upper teeth DP 4. (Fig. 45 a to i). Probably due to different taphonomic conditions, the DP 4 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 DP 4 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 DP 4 (SMX 1 - 253: Fig. 45 a) 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 DP 4. Te postparacrista of SMX 1 - 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 (SMX 1 - 254: Fig. 45 b, 259, 260: Fig. 45 g, 264; SMX 2 - 216: Fig. 46 h, 217: Fig. 45 j). On the unworn SMX 1 - 255: Fig. 45 c and weakly worn SMX 1 - 256: Fig. 45 d, the paraconule is strong and indistinct from the anterostyle. Te protoloph is discontinuous on SMX 1 - 255. On the latter specimen, like on SMX 1 - 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 SMX 1 - 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 (SMX 1 - 256, 258; SMX 2 - 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 (SMX 1 - 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 (SMX 1 - 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 (SMX 1 - 256, 259, 260) or vestigial. Te metaconule generally separates from the lingual cusps, except on SMX 1 - 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 DP 4 (SMX 1 - 264: Fig. 45 k) 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 SMX 1 - 260: Fig. 45 g, which is smaller. Te SMX 1 - 252 is a pristine DP 4 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. P 4. (Fig. 45 j 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 (SMX 2 - 218: Fig. 45 j). 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 P 4 differ from SMX 2 - 218 in their smaller size and slender roots. Teir shape is more trapezoidal (SMX 1 - 262: Fig. 45 l, and 263: Fig. 45 m) or rounded (SMX 1 - 265: Fig. 45 n), with a very small to absent hypocone, longer (although short) anteroloph and posteroloph (except SMX 1 - 263 where the anteroloph is as reduced as on SMX 2 - 218). A strong conule widens the mesial end of the preprotocrista on SMX 1 - 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 SMX 1 - 263 and present on SMX 1 - 262 and 265. On the latter, a narrow and shallow break separates the mesostyle from the postmetacrista. M 1 – 2. Description of the type SMX 1 - 281. (Fig. 46 a). Te type is the largest upper molar of the sample (1. 91 × 2. 02 mm; Figure 5 – 8 in Escarguel, 1998). It could represent an M 2, 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 Teeth of? Protadelomys alsaticus from Cuzal (Quercy). a CUZ 183, left DP 4; occlusal view. b CUZ 176, left DP 4; occlusal view. c: CUZ 177, left P 4; occlusal view. d CUZ 188, right P 4 – M 1; occlusal view. e CUZ 189, left P 4 – M 1; occlusal view. f CUZ 155, left M 1; occlusal view. g CUZ 156, right M 1; occlusal view. h CUZ 182, left M 1; occlusal view. i CUZ 178, right M 1; occlusal view.: CUZ 155, left M 1; occlusal view. g, right M 1; occlusal view. h CUZ 182, left M 1; occlusal view. i CUZ 178, right M 1; occlusal view. j: CUZ 186, left M 1; occlusal view. k: CUZ 154, right M 2; occlusal view. l: CUZ 179, right M 2; occlusal view. m: CUZ 180, right M 3; occlusal view. n: CUZ 187, left dp 4; n 1, occlusal view; n 2, buccal aspect. o: CUZ 161, right dp 4; occlusal view. p: CUZ 152, left p 4; p 1, occlusal view; p 2, buccal aspect. q: CUZ 164, right m 1 – m 2 – m 3; occlusal view. r: CUZ 168, right m 1; occlusal view. s: CUZ 163, left m 1; occlusal view. t: CUZ 153, left m 1; occlusal view. u: CUZ 167, right m 1; occlusal view. v: CUZ 172, right m 2; occlusal view. w: CUZ 169, left m 2; occlusal view. X: CUZ 173, left m 3; occlusal view. Scale bar = 1 mm Other M 1 – M 2. (Fig. 46 b to o). Te molars (SMX 1 - 269, 270, 271, 272, 275, 278, 279, 282, 284, 285, 286, 288; SMX 2 - 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 M 2 is shorter than the anterior part, the hypocone and metacone being closer than on M 1. 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 (SMX 2 - 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 SMX 1 - 275: Fig. 46 f 3). One or two (e. g., SMX 1 - 275) lingual metalophules attach to the hypocone. Te endoloph is thick and high. On SMX 1 - 278: Fig. 46 h, a low entostyle is located below the endoloph. Te posteroloph is short and rarely reaches the distolingual corner of the metacone (SMX 1 - 271: Fig. 46 e). Some M 1 (SMX 1 - 273) and M 2 (SMX 1 - 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 M 1 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 M 2 SMX 1 - 276: Fig. 46 o, which however displays numerous extra-ridges. Te M 2 SMX 1 - 280, (Fig. 5 – 5, in Escarguel, 1998: Fig. 46 m) 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. M 3. (Fig. 46 p). 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 dp 4. (Fig. 47 a to g). Teir morphology is relatively homogeneous, but a few features are only present in the smallest dp 4, which are here attributed to? Protadelomys cf. maximini morphotype 2. Te two roots diverge, as usual on dp 4. Te protoconid is present, small, and lower than the metaconid. A mesial notch separates the two cusps mesially (except on SMX 1 - 203), but a short metalophulid I can be present. Te latter is absent on SMX 1 - 201 and SMX 2 - 205, and interrupted on SMX 1 - 206. A short anterocingulid descends from the protoconid on some teeth (SMX 2 - 204, 205: Fig. 47 e, f). On all the dp 4 s, 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 (SMX 1 - 203, 206: Fig. 47 b, d). 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. 47 b 1, d 2), which is linked to a short ectocingulid that closes the base of the sinusid (SMX 1 - 203, 206). Tis cingulid can be simply a flat and not a ridge (SMX 1 - 201; SMX 2 - 205: Fig. 47 a, f). Te short prehypocristid descends towards the ectolophid. Te entoconid is small and conical. When present, the entolophid is low, complete (SMX 1 - 203; SMX 2 - 204) or incomplete (SMX 1 - 202, 204, 206; SMX 2 - 205; SMX 3 - 26). It is connected, or directed, to the junction prehypocristid – ectolophid, or to the hypoconid (SMX 1 - 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 (SMX 1 - 202, 203; SMX 2 - 202, 204, 205) to numerous (SMX 1 - 206: Fig. 47 d). p 4. (Fig. 47 h to k). Tere are seven p 4 from Saint-Maximin, only four of them can be described in details, the others being too much worn or damaged. On p 4, the main cusps are more bulbous and robust than on dp 4. 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 p 4 only (SMX 1 - 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 SMX 1 - 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 SMX 1 - 208. A spur on the protoconid distal flank prolongs in an ectostylid on SMX 1 - 210. A short ectomesolophid is noticeable on SMX 1 - 208 and 210. m 1 – m 2. (Fig. 48 a to g). Like in Bouxwiller, we observed a large morphological variation among the 34 m 1 – m 2 from Saint-Maximin, which it is worth comparing to the size variation. For both m 1 and m 2, 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 m 2 SMX 1 - 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 (m 1 – m 2) are considered as typical? P. maximini. (See figure on next page.) Fig. 46 Upper molars of? Protadelomys maximini Escarguel, 1998 from Saint-Maximin (Gard). a SMX 1 - 281, left M 1, Holotype; a 1, occlusal view; a 2, lingual aspect; a 3, buccal aspect. b SMX 1 - 269, left M 1 or 2; b 1, occlusal view; b 2, buccal aspect. c: SMX 1 - 287, left M 1, enamel cover only; occlusal view. d SMX 1 - 273, right M 1; d 1, occlusal view; d 2, buccal aspect; d 3, lingual aspect. e SMX 1 - 271, left M 1; e 1, occlusal view; e 2, lingual aspect; e 3, buccal aspect. f SMX 1 - 275, right M 1; f 1, lingual aspect; f 2 ,, buccal aspect; f 3; occlusal view. g SMX 2 - 224, right M 1; g 1, occlusal view; g 2, lingual aspect; g 3: buccal aspect. h SMX 1 - 278, right M 1; h 1, occlusal view; h 2, lingual aspect; h 3: buccal aspect. i SMX 2 - 221, left M 1; i 1, occlusal view; i 2, lingual aspect; i 3, buccal aspect. j: SMX 2 - 222, right M 1 or M 2; j 1, occlusal view; j 2, lingual aspect; j 3, buccal aspect. k: SMX 2 - 220, left M 1; k 1, lingual aspect; k 2, occlusal view; k 3, buccal aspect. l: SMX 1 - 223, left M 2; l 1, occlusal view; l 2, lingual aspect; l 3, buccal aspect. m: SMX 1 - 280, left M 2; m 1, occlusal view; m 2, lingual aspect; m 3: buccal aspect. n: SMX 1 - 268, right M 2; occlusal view. o: SMX 1 - 276, left M 2; o 1, occlusal view; o 2, lingual aspect. p: SMX 1 - 291, left M 3; p 1, occlusal view; p 2, 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 SMX 1 - 230. All the lophids and ridges are thin. Te anteroconid (mainly distinct on m 1) and the anterolophid are present but are thinner than in? P alsaticus. Te anteroflexid is generally closed distally by a complete (buccolingual) metalophulid I (SMX 1 - 213, 214, 216, 218, 220, 221, 227, 228, 229, 230, 246, 250; SMX 2 - 212); it is rarely opened distally when there is no connection between the buccal and the lingual halves of the metalophulid I (SMX 2 - 208, 209, 213, 215; SMX 1 - 232, 238 (= 298 in Escarguel, 1998, Fig. 4 – 9), 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 (SMX 1 - 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., SMX 1 - 227, 238, SMX 2 - 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., SMX 1 - 227, 230; SMX 2 - 209). A postmesoconid spur is present on some specimens (e. g., SMX 2 - 212, SMX 1 - 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. m 3. (Fig. 48 h, i). All eight m 3 are quite larger than the m 1 – 2. Tey show the same features as the other molars.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFE0DF6BC483456EFBF7FE19.taxon	description	DP 4. (Fig. 50 a). Te smallest DP 4 from Saint-Maximin (SMX 3 - 44), trapezoidal, differs from the small DP 4 attributed to? P. cf. maximini morphotype 1, in its narrower size, and relatively longer anteroloph. Its hypocone occupied a very lingual position when compared to the protocone, whereas the two cusps are roughly located at the same lingual level on typical? P. maximini, and slightly less lingual in? P. cf. maximin i morphotype 1. M 1 – M 2. (Fig. 50 b, c). Tree M 1 – 2 (SMX 3 - 46, 47, and 48) are smaller than the molars of typical? P. maximini. Te enamel surface is slightly ornamented, with only a few granules. Te hypocone is conical. Te arms of the protocone are mesiodistal, the posterior one being short and ending at the level of the short endoloph. Te postparacrista is weak while the premetacrista is slightly stronger and frames a small mesostyle, which prolonged in a short and slender buccal mesoloph. Paraconule and metaconule are protruding and nearly equal in size. Te metaconule is linked to the apex of the hypocone. Tere is a mid-protocone ridge (protocrista) directed towards the paraconule on SMX 3 - 47 (Fig. 50 c). On the latter, unworn, the metalophule II is straight, and does not connect to the metaconule. Short ridges represent vestigial metalophule I, which is directed towards, on the unworn M 1 or joining the mesoloph on SMX 3 - 48. LoWer teeth dp 4 .. (Fig. 50 d to f). Te specimen SMX 1 - 207 previously considered as a p 4 (Escarguel, 1998: 372), is here considered as a dp 4 due to the distal direction of its posterior root and to a distinct low protoconid close to the metaconid. Tis specimen is one of the shortest dp 4 (1. 13 × 1. 37 mm), with SMX 3 - 40: Fig. 50 d, SMX 3 - 41: Fig. 50 f, SMX 3 - 42: Fig. 50 e, and SMX 2 - 202 (the latter being figured as P. maximini: Fig. 4 – 3, in Escarguel, 1998). Te metaconid and protoconid are almost unworn on SMX 1 - 207, the second being much lower and slender than the metaconid, and placed a little more distally, which indicates that this tooth likely corresponds to a dp 4 rather than a p 4; a mesial notch separates the protoconid from the metaconid. Te long posprotocristid ends with a weak premesoconid swelling, before joining the mesiodistal ectolophid, which is small. Te mesoconid is tiny. Te center of the basin is not well distinct on SMX 1 - 207, but a long mesiodistal ridge, a short mesolophid, and an entolophid nearly complete are perceivable. Te posterolophid does not reach the entoconid and the hypoconulid. SMX 3 - 42 display similar features as SMX 1 - 207. SMX 2 - 202 and SMX 3 - 41 have neither a long mesiodistal ridge, nor a long entolophid, but show small low granules and ridges in the basin. On SMX 3 - 40, the entolophid is better individualized. m 1 – m 2. (Fig. 50 g, h). Te m 1 SMX 1 - 222 and 226 have the anterolophid reduced to the anteroconid, whereas it is longer on SMX- 225. Tis anteroconid – anterolophid does not connects the short premetacristid (= mesial metalophulid I) on SMX 1 - 222 and 226; but it does on SMX 1 - 225. A more distal lingual metalophulid I descends from the metaconid and joins the transverse buccal metalophulid I on the three m 1. Te thick oblique postprotocristid develops from the protoconid apex; it bears a low premesoconid swelling or a spur, in front of the mesoconid, which is slightly stretched and in line with the postprotocristid. A postmesoconid spur is present at the junction with the distal ectolophid, which is lower than the postprotocristid. Te prehypocristid is low and short. Some elements of the entolophid join this spur. Tere are low and thin extra-ridges in the basins. Te posterolophid is long, connecting the distal flank of the entoconid, through a postentocristid. Te hypoconulid is small. Te entolophid is low, variably long and incomplete and does not reach the hypoconid or ectolophid. SMX 1 - 225 also presents a short ectomesolophid. Te unique m 2 SMX 1 - 400 displays a similar morphology to that of the m 1, but the anterolophid is longer, slender, and connected to the premetacristid. Its lingual metalophulid I does not fuses the buccal metalophulid I. Te weak entolophid is reduced to its lingual part.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFE0DF6BC483456EFBF7FE19.taxon	materials_examined	Type locality. Casa Ramòn (Huesca, Spain); middle Eocene (MP 12?).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFE0DF6BC483456EFBF7FE19.taxon	discussion	Remarks. P. Peláez-Campomanes described the new species Protadelomys nievesae (1995) from Casa Ramòn (Huesca, Spain). Here we provide a specimen list with their measurements as Additional file 7: Data S 7 and compare them with? P. maximini (Escarguel, 1998),? P. alsaticus, Protadelomys cartieri, and P. lugdunensis (Hartenberger, 1969). Original diagnosis. Peláez-Campomanes, 1995: 301. “ Te cheek teeth are small. Te permanent and the deciduous premolars are smaller than the molars. Te metalophule is directed towards the protocone instead of towards the hypocone in DP 4 and P 4. Te anterior lobe of P 4 is slightly larger than the posterior one. Te lower molars do not have a mesoconid ”. “ P. nievesae differs from P. cartieri by its smaller premolars and its P 4 which has a poorly developed anterior part. P. nievesae differs from P. alsaticus by the following features: DP 4 with well-developed mesostyle and the metaconule and the paraconule connected to the adjacent cusps; P 4 and DP 4 with the metalophule directed towards the protocone; P 4 with the anterior lobe poorly developed and lower molars without mesoconid. P. nievesae differs from P. lugdunensis by its smaller size, its lower molars without ectomesolophid and its P 4 with the anterior lobe poorly developed ”.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFE0DF6BC483456EFBF7FE19.taxon	materials_examined	Material and measurements. (Additional file 7: S 7; Fig. 51; Table 7). Te holotype RP 022 is probably an m 2, because the mesial (trigonid) and distal (talonid) sides show similar widths. Te teeth are significantly smaller than those of the other species of? Protadelomys as well as from Protadelomys cartieri and P. lugdunensis. (Bivariate graphs in Figs. 3, 14, 28, 44, 51; Tables 6, 7, 8.)	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFE0DF6BC483456EFBF7FE19.taxon	discussion	Remarks about differential diagnosis, and comparisons? Protadelomys nievesae differs from? P. maximini and? P. alsaticus in its DP 4 showing a less stretched protocone and hypocone, shorter pre- and post-protocristae and hypocristae. DP 4 of? P. nievesae and? P. maximini share well-developed buccolingually transverse protoloph and metaloph. Te DP 4 and molars of? P. nievesae and of? P. alsaticus differ from? P. maximini in showing a less reduced hypocone. Te reduced anterior part of the P 4 (anteroloph short to absent and reduced anteroflexus) is present in 7 / 9 teeth of? P nievesae; the anteroloph is otherwise long and presents a narrow anteroflexus on 2 / 9. Te reduced anterior part is not a unique feature of? P nievesae, but mainly characterizes? P. maximini (5 / 5 P 4). Te number of P 4 with reduced anterior part varies also in P. cartieri (5 / 9) and P. lugdunensis (3 / 5). Owing to the weak number of P 4, we consider that these differences are not significant. On the upper molars of? P. nievesae and? P. alsaticus, the cusps and conules are more bulged and distinct from the lophs than in? P. maximini, and the endoloph is lower. The sinus is variably present in? P. nievesae and less individualized in? P. alsaticus, whereas it is nearly absent in? P. maximini. The hypocone is generally distinct on unworn P 4 of? P. nievesae. On upper molars, a lingual metaloph is not always distinct. When present, it joins the hypocone or the endoloph in? P. maximini, although more rarely. In the latter, it is very low as on molars of? P. nievesae. In? P. nievesae, the metaloph is sometimes oriented towards the protocone on DP 4 (2 / 5) and on P 4 (1 / 8), towards the hypocone on DP 4 (2 / 5) and more frequently on P 4 (5 / 8) and towards the posteroloph on P 4 (2 / 8). This lingual metaloph is more often distinct on molars of P. cartieri and P. lugdunensis even if these teeth are only moderately worn; on these molars it connects the hypocone, or the endoloph. In Protadelomys and? Protadelomys species, the size of the mesostyle is variable, and the mesoloph is variably long, although it never bypasses the metaconule level. Te dp 4 basin floor of? P. maximini is flatter than that of the dp 4 of? P. alsaticus or? P. nievesae. Te p 4 protoconid is often absent, exceptionally present and well distinct in? P nievesae (2 / 7) whereas it is present only on one p 4 of? P. maximini, on which it is low. On lower molars, the entolophid is generally relatively high and complete (11 / 15) or shortly interrupted (4 / 15) in? P. nievesae, whereas it is more often discontinuous and low in? P. maximini. It is generally complete in P. cartieri, P. lugdunensis, and? P. alsaticus.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFE7DF66C73941FDFDE0FC99.taxon	materials_examined	Holotype. Ek H 004, lower left tooth row, with p 4 – m 3. Type locality. Egerkingen γ (Swiss Jura); middle Eocene (MP 14).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFE7DF66C73941FDFDE0FC99.taxon	diagnosis	Diagnosis. Te lower molars display some features considered derived for theridomyids (e. g., Vianey-Liaud & Marivaux 2016; 2021): p 4 longer than m 1, with metaconid and protoconid mesially positioned; long thick postprotocristid; high ridges on the lingual (postmetacristid) and distal (post-hypocristid + posterolophid) edges of the teeth; numerous granules and low extra-ridges and granules in the basin; metalophulid I complete, developed from the mesial edge of the metaconid (premetacristid) to the preprotocristid. Tey also show: dp 4 shorter than p 4. Lower molars lengths increasing from m 1 to m 3. Anteroconid absent on p 4 and mesiobuccally situated on molars; short on molars but extending buccolingually more and more from m 1 to m 3, reaching the middle of the mesial border on m 3. Entolophid low and discontinuous. Ectolophid mesiodistal from p 4 to m 2 (nearly aligned with the postprotocristid on p 4, a little oblique on m 3), with mesoconid weakly marked. Ectomesolophid present on p 4. Te molar morphological features differs from that of Treposciurus in showing lower crowned and more concave occlusal surface of teeth, less individualized lophids (metalophulid I and entolophid), the metaconid fused in a curved and high crest with the postmetacristid, a smaller mesostylid, and smaller granules on enamel surface.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFE7DF66C73941FDFDE0FC99.taxon	materials_examined	Material. Te holotype Ek 1004 (p 4 to m 3); Ek 79, a right lower dp 4, and possibly a left M 1 (Ek 85). Even if the material is poor, it is possible to erect a new taxon, due to the peculiar features of the lower teeth. Upper tooth M 1 (Fig. 52 c). Compared to Protadelomys of the same locality, the weakly worn M 1 Ek 85 displays some peculiar features. It is larger, its paraconule and metaconule showing similar size and being both isolated. Te parastyle is mesiobuccal; the anterostyle is present. Te protoloph reduces to a short thin buccal ridge posteriorly oriented; it is followed lingually by the distinct paraconule A well-marked protocrista is present, its mesial end being free. Te strong mesostyle stretched mesiodistally and surrounded by two small cusps, closes the mesoflexus buccally. Te metacone shows a short premetacrista; the buccal metalophule II is present, but short. Two small granules could constitute the remnants of the buccal metalophule I. Te metaconule, not larger than the paraconule, is isolated both from the buccal metalophuleI and from the lingual cusps, protocone and hypocone. Te mesoflexus displays a few extra-ridges. Te hypocone appears smaller than the protocone. Te endoloph is relatively long, thin and high. It is slightly marked lingually by a shallow and short sinus. LoWer teeth dp 4 (Fig. 52 b). Te base of the crown EK 79 is poorly preserved and partly covered with glue. Despite these facts, at the metaconid level, the crown is clearly higher than that of dp 4 of the other species of the same locality. Te postmetacristid is high and buccodistally compressed, overhanging the lower central basin; a tiny mesostylid is distinct. Te protoconid is merged in the oblique protocristid, which ends in a weak mesoconid. Te distal ectolophid is interrupted in front of the hypoconid. Te prehypocristid is long and oblique bucco-mesially. Te hypoconid and entoconid are equally robust, bulbous with their upper part tapered. Wrinklings are numerous on the enamel surface, some making a kind of ectocingulid at the buccal opening of the sinusid. p 4. Te specimen Ek H 004 (Fig. 52 a) was first described and figured by Hartenberger (1969, p. 54; pl. 1, Fig. 4) as gen. and sp. indet, close to Suevosciurus; then as Treposciurus in Hartenberger (1990) and Escarguel (1999: 246). It is a little longer (2. 20 mm) than m 1 (2.09 mm). Te protoconid is close to the metaconid, but it is somewhat lower and separated by a shallow and narrow groove. Te metaconid curves mesio-lingually and prolongs in the high and narrow mesiodistal postmetacristid. Te postmetacristid ends at the base of a low and small entoconid, leaving the wide mesosynclinid opened lingually. Several ridges run obliquely in both the mesosynclinid and the posterosynclinid. Te lophids are crenulated by short wrinkles. Te postprotocristid is slightly oblique, long, and marked at the base of its distal end by a short edge. Te sinusid is shallow, and the crown high below. Te ectolophid is located in the continuity of the postprotocristid; it is long and bears a long ectomesolophid reaching the floor of the sinusid. Te ectolophid joins a short prehypocristid. Te entolophid is low and interrupted between its buccal and lingual halves. One thin ridge connects the lingual half with the posterolophid, which does not join the apex of the entoconid. Lower molars. Te lower molars have a concave talonid, with low and discontinuous transverse lophids and high edging ridges. Te basin enamel surfaces are strongly wrinkled, with short ridges and granules, while the lophids are crenulated. Te antesinusid – anteroflexid always opens buccally and lingually. m 1 – m 2. Te m 1 (2.09 × 1.75 mm) differs from the m 2 (2.34 × 2.13 mm) in displaying a smaller size, a metaconid closer to the protoconid than the entoconid to the hypoconid, a smaller anteroconid located closer to the protoconid than to the metaconid. On the m 2 and m 3, the anteroconid is equidistant from the protoconid and metaconid, and stretched in a short and low anterolophid. In both cases, the anteroconid does not connect with the metalophulid I; the anterolophulid is lacking but two to four low mesiodistal wrinkles are present in the anteroflexid. Te metalophulid I is rectilinear due to the alignment of its lingual and buccal halves; it connects to the preprotocristid. Te postprotocristid is thick, long, oblique, and its end bears a short premesoconid spur on the m 1, but not on the m 2 – 3. Te ectolophid is mesiodistally oriented, long, and swollen at the level of the mesoconid; it makes an obtuse angle with the postprotocristid. Its distal extremity is lower than the level of the connection between the prehypocristid and the entolophid. Te latter angles at an entoconulid level and it is low and incomplete lingually. Te post-hypocristid and the posterolophid are continuous. Te hypoconulid is not distinct or salient. m 3 (2.66 × 2.03 mm) is longer than m 2. Tis elongation affects the anterior part of the tooth; while the posterior lobe width is slightly reduced with respect to m 2. Consequently, its anterolophid is longer than that of the m 2, ditto for the postmetacristid and the postprotocristid. Te short ectolophid is lower than the postprotocristid. Comparison With Treposciurus mutabilis [from Ehrenstein 3 (MP 18, upper Eocene)] Because these specimens have been previously referred to Treposciurus, we compared them to a (younger) species of this genus, T. mutabilis, whose teeth are highly ornamented. Te major similarities between Homoetreposciurus and Treposciurus lie in the ornamentation of the enamel surface, with numerous wrinkles and granules. However, the wrinkles and ridges are more continuous and reticulated in Treposciurus. We also found a number of differences between the two genera. Compared to Homoetreposciurus, Treposciurus mutabilis is characterized by higher crown and higher main cuspids; a p 4 smaller than or equal to m 1, whereas m 3 is equal or smaller than m 2; better defined and higher transverse lophids (metalophulid I and entolophid), even if the anterolophid remains low; a longer and lingually developped anterolophid, while the anteroconid remains buccal and close to the protoconid; a higher metaconid that is tapered at its apex and not fused in a curved and high crest with the postmetacristid; a less oblique postprotocristid; a more salient mesoconid, with the ectomesolophid present on all the molars; a mesostylid often distinct and sometimes extending in a short lingual mesolophid; a less reduced posterior lobe of m 3. Similar differences were observed with older species of Treposciurus, such as T. preecei from Creechbarrow (Bartonian), even if their teeth are smaller, with less ornamented enamel.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFEADF67C7394573FD6AFDB9.taxon	description	(Fig. 53 c to g).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFEADF67C7394573FD6AFDB9.taxon	type_taxon	Type species. Eoelfomys lapradensis nov. gen. nov sp.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFEADF67C7394573FD6AFDB9.taxon	materials_examined	Type locality. Laprade (Lot, Quercy), middle Eocene (MP 14).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFEADF67C7394573FD6AFDB9.taxon	diagnosis	Diagnosis. Tat of the type species Eoelfomys lapradensis nov. gen. nov. sp.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFEADF67C7394573FD6AFDB9.taxon	discussion	Remarks Te smallest teeth from Laprade were referred to Elfomys nov. sp. by Hartenberger (in Sudre et al., 1990) and this identification was taken over by Comte et al. (2012). However, even if these teeth display some morphological and length similarities with Elfomys engesseri Hooker & Weidmann, they are larger and show unique feature combination, which justifies the creation of a new taxon, to which the mandible Ek H 003 from Egerkingen could belong.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFEBDF67C4834172FB95FA79.taxon	description	Elfomys sp. in Comte et al., 2012: Fig. 17; Pl. 8, Fig. x to z	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFEBDF67C4834172FB95FA79.taxon	materials_examined	Holotype. LAP 243, unworn right upper M 1. (Fig. 53 c).	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFEBDF67C4834172FB95FA79.taxon	diagnosis	Diagnosis. Size close to that of the large Elfomys engesseri Hooker & Weidmann, 2007; smaller than E. catalaunicus Bonilla-Salomon et al., 2016. Differs from the Eocene Elfomys species in: - Te occurrence of a strong and high endoloph, which prevents the communication between the sinus and the mesoflexus; - Te absence of connection between the metalophule II and the posteroloph on the upper teeth, the absence of a long prehypocrista connecting a mesiobuccal metaconule crest, the presence of a strong metaconule; - Te occurrence of a long low mesolophid on m 2 – m 3.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFEBDF67C4834172FB95FA79.taxon	materials_examined	Material and measurements. Appendix, S 3: two M 1, one M 2, one m 2, one m 3. (cf. Figure 17, in Comte et al., 2012). Teir size is close to that of Elfomys engesseri Hooker & Weidmann, 2007, and larger than E. catalaunicus Bonilla-Salomon et al., 2016.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFEBDF67C4834172FB95FA79.taxon	description	Description. Te crowns are unilaterally hypsodont but low. M 1 – 2. Te parastyle of the two M 1 is slightly curved towards the paracone. LAP 243, the holotype (Fig. 53 c), shows a premetacrista, not easily distinguishable on the other two upper molars (LAP 244 and 245), which are worn. Te posterior area is narrower on the M 2 (LAP 244) than on the M 1 (LAP 243, 245). Te three teeth have a high endoloph separating the shallow sinus from the mesoflexus. Te paraconule stretches forward and protrudes mesially. Te attachment of the protoloph to the mid-protocone is weak. Te mesostyle is present. Te mesoloph either reaches the buccal flank of the metaconule (LAP 245: Fig. 53 d), or is shorter (LAP 244: Fig. 53 e) or replaced by a low ridge reaching the buccal base of the metaconule (LAP 243). On the latter, a very low short ridge is reminiscent of a vestigial metalophule I. Te metalophule II is short and weakly connected to the base of the metaconule. Te latter also weakly attaches to the mid-hypocone. Tere are scarce granules on the enamel surface. Te main differences with the late Eocene species of Elfomys are the strong endoloph together with the absence of communication between the sinus and the mesoflexus, the absence of relationship of the metalophule II with the posteroloph, the absence of a long prehypocrista connecting to the mesiobuccal metaconule crest. m 2 - 3. Te teeth are flatter than that of the late Eocene Elfomys, the metaconid less salient, the anteroconid smaller. On m 2 (LAP 242; Fig. 53 f), the anterolophid is very short, the anteroflexid reduced, and the anterocingulid absent. Te two parts of the metalophulid I are continuous, even if their extremities are not completely fused. Te long and oblique postprotocristid makes an angle with the short mesiodistal ectolophid. From the junction between the postprotocristid and the ectolophid, a few low granules punctuate the path of a mesolophid. Te mesoconid is weak. Te entolophid is straight and connects the junction of the prehypocristid to the ectolophid. Te sinusid and mesoflexid do not communicate. On m 3 (Fig. 53 g), the anterolophid is a little longer and it is weakly attached to the short lingual part of the metalophulid I. Te buccal part of the metalophulid I is free and distally oriented. Small and very low granules trace the mesolophid path. Te posterior lobe is reduced.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
497F1B1DFFEBDF67C4834172FB95FA79.taxon	discussion	Remarks and comparisons As for basal Teridomorpha and the middle Eocene species of Masillamys or Protadelomys, this genus displays a high endoloph. Eoelfomys could represent an early stage preceeding Elfomys, genus in which the endoloph disappears, the prehypocrista lengthens to the mesoloph, and the distal ectolophid reduces. Te size and morphology of its lower molars is similar to those of the mandible Ek H 003 from Egerkingen.	en	Vianey-Liaud, Monique, Hautier, Lionel (2022): Revision of the genus PRotadelomYS, a middle Eocene theridomyoid rodent: evolutionary and biochronological implications. Swiss Journal of Palaeontology (8) 141 (1): 1-98, DOI: 10.1186/s13358-022-00245-3, URL: https://doi.org/10.1186/s13358-022-00245-3
