taxonID	type	description	language	source
375A87FAFFEBFFFD6DA5FBF8FB54FFD8.taxon	description	urn: lsid: zoobank. org: act: B 56 C 993 F-A 5 C 1 - 4 ED 1 - A 3 E 2 - 05426 EA 112 EC	en	Gheerbrant, Emmanuel, Teodori, Dominique (2021): An enigmatic specialized new eutherian mammal from the Late Cretaceous of Western Europe (Northern Pyrenees). Comptes Rendus Palevol 20 (13): 207-223, DOI: 10.5852/cr-palevol2021v20a13, URL: https://doi.org/10.5852/cr-palevol2021v20a13
375A87FAFFEBFFFD6DA5FBF8FB54FFD8.taxon	type_taxon	TYPE SPECIES. — Azilestes ragei n. sp.	en	Gheerbrant, Emmanuel, Teodori, Dominique (2021): An enigmatic specialized new eutherian mammal from the Late Cretaceous of Western Europe (Northern Pyrenees). Comptes Rendus Palevol 20 (13): 207-223, DOI: 10.5852/cr-palevol2021v20a13, URL: https://doi.org/10.5852/cr-palevol2021v20a13
375A87FAFFEBFFF26D87FB3FFA64FC1B.taxon	description	(Figs 3 - 7) urn: lsid: zoobank. org: act: 52 DF 0 B 06 - 5 E 24 - 449 C-AD 95 - 24 C 10986 E 30 C	en	Gheerbrant, Emmanuel, Teodori, Dominique (2021): An enigmatic specialized new eutherian mammal from the Late Cretaceous of Western Europe (Northern Pyrenees). Comptes Rendus Palevol 20 (13): 207-223, DOI: 10.5852/cr-palevol2021v20a13, URL: https://doi.org/10.5852/cr-palevol2021v20a13
375A87FAFFEBFFF26D87FB3FFA64FC1B.taxon	etymology	DERIVATIO NOMINIS. — Azilestes: Azil from the name of the type locality, Mas-d’Azil, and lestes, Greek, robber, pirate by allusion to the small size of many insectivore-like mammals from the Cretaceous and Paleogene; ragei: dedicated to Jean-Claude Rage as a tribute to his memory and to his major contribution on the study of Cretaceous European microvertebrates.	en	Gheerbrant, Emmanuel, Teodori, Dominique (2021): An enigmatic specialized new eutherian mammal from the Late Cretaceous of Western Europe (Northern Pyrenees). Comptes Rendus Palevol 20 (13): 207-223, DOI: 10.5852/cr-palevol2021v20a13, URL: https://doi.org/10.5852/cr-palevol2021v20a13
375A87FAFFEBFFF26D87FB3FFA64FC1B.taxon	materials_examined	HOLOTYPE. — MNHN. F CTE 215, fragment of left dentary with damaged M 1 and M 2. LOCALITY AND AGE. — Mas-d’Azil (Capens), Ariège, Plantaurel massif, Northern Pyrenees, France; upper part of the Formation des Grès de Labarre, early Maastrichtian.	en	Gheerbrant, Emmanuel, Teodori, Dominique (2021): An enigmatic specialized new eutherian mammal from the Late Cretaceous of Western Europe (Northern Pyrenees). Comptes Rendus Palevol 20 (13): 207-223, DOI: 10.5852/cr-palevol2021v20a13, URL: https://doi.org/10.5852/cr-palevol2021v20a13
375A87FAFFEBFFF26D87FB3FFA64FC1B.taxon	diagnosis	DIAGNOSIS. — This species belongs to a tribosphenic therian mammal. Its molar morphology (e. g. hypoconulid and entoconid not twinned, paraconid much smaller that metaconid, trigonid compressed) differs from metatherians and marsupials, but indicates it belongs instead to a Cretaceous eutherian, and one the largest known. The partial dental formula preserved in the holotype is C 1, P 4, P 5, M 1 - 2 (incisors and last molar broken away). This species differs from all known Cretaceous eutherians by the following specialized traits: short and robust (deep) dentary; shortened anterior dentition lacking at least P 1 - 3 and any lower diastema, and with small single-rooted P 4; mandibular symphysis extended close to P 4 level; hypoconulid and postcristid forming an isolated transverse cingular-like structure at the posterior part of the talonid; presence an incipient but distinct hypolophid and a well-developed posterior fovea. The molars have a robust bunodont and grinding-crushing morphology that is associated with the presence an interradicular crest and radicular grooves.	en	Gheerbrant, Emmanuel, Teodori, Dominique (2021): An enigmatic specialized new eutherian mammal from the Late Cretaceous of Western Europe (Northern Pyrenees). Comptes Rendus Palevol 20 (13): 207-223, DOI: 10.5852/cr-palevol2021v20a13, URL: https://doi.org/10.5852/cr-palevol2021v20a13
375A87FAFFEBFFF26D87FB3FFA64FC1B.taxon	description	DESCRIPTION Although damaged and incomplete, the specimen MNHN. F CTE 215 is one of the most complete jaw remains of eutherian mammals reported from the late Cretaceous of Europe. Dentary (Figs 3; 5) MNHN. F CTE 215 preserves only part of the mandibular corpus including the posterior part of the mandibular symphysis which extends far posteriorly, close to level of the unique P 4 alveolus. It is very short and robust. The mandibular corpus is dorso-ventrally high, and labially inflated. It is convex below the molars, but concave below the premolars (P 4 - 5) and the symphysis. Its dorso-ventral depth rapidly decreases at the level of the symphysis. The posterior mental foramen is rather large and located ventro-labially below and between the two roots of P 5, as in most Cretaceous eutherians. Based on the CT scans, the mandibular canal was reconstructed as large and located directly ventral to the tooth roots (Fig. 5). Laterally, it extends on the labial side of the roots, but not on the lingual side. Its diameter is more or less constant up to P 4 level along preserved corpus length. Its cross-section is typically dorso-ventrally flattened below the teeth, with a flat dorsal margin. Anteriorly, the mandibular canal divides in several small lateral branches (“ lat br ”, Fig. 5) labially to P 5 and P 4 (anterior) roots and lingually to the canine alveolus in the mandibular symphysis (Fig. 5 A, B; Appendices 1; 2). The preserved anteriormost part of the mandibular canal is smaller in cross-section and dorso-labial to the canine alveolus (Fig. 5 B, G). Tooth row The dentaryMNHN. F CTE 215 preserves two damaged molars, M 1 and M 2, and four well-developed empty anterior alveoli, interpreted as corresponding to a two-rooted P 5, and single-rooted P 4 and C 1 (Figs 5; 6 and see below). The anterior and posterior parts of the dentary are broken, so that M 3, the canine and the incisors and their alveoli (except part of C 1 alveoli) are not preserved. There is no diastema. Anteriorly to the P 4 alveolus, there is only one very large alveolus, most probably for the canine (instead for an enlarged incisor, see below). It suggests a strongly reduced anterior cheek dentition in this species (only two premolars, P 4 being reduced), which is also supported by the position of the mandibular symphysis close to P 4. Such a reduced tooth row is unique in stem eutherians. The dentary and its tooth-row curve lingually from the level of P 4. Teeth (Figs 3 - 6) Canine. The CT scans and our 3 D reconstruction (Figs 5 - 7) show the occurrence one large anterior alveolus extending within the mandibular symphysis in its posterior part. It is identified as the alveolus for the lower canine because it has a size proportion and relative position to the symphysis known for this tooth in many Cretaceous eutherians. Enlarged lower incisors are known in a few Cretaceous eutherians such as zalambdalestids, but they also have a canine (e. g. Wible et al. 2004), and they have distinctive molar morphology. The alternative interpretation of the large anterior alveoli of MNHN. F CTE 215 for the presence an enlarged incisor and the absence of a canine is unlikely for Cretaceous therians. Only the lower and posteriormost part of the large anterior alveolus is preserved (Figs 5 D, G, H; 6). The alveolus is open anteriorly at the anterior part of specimen MNHN. F CTE 215 where it is broken. Its orientation suggests a mesially inclined root within the dentary. A procumbent canine root is known in zhelestids such as Parazhelestes Nessov, 1993 (Archibald & Averianov 2012: figs 15 A; 22 B). The alveolus size (3.3 × 2.7 mm at anterior preserved section, measured from the 3 D digital model) indicates a large C 1. Large to very large canine are known in eutherians such as the zhelestid Eoungulatum Nessov, Archibald & Kielan-Jaworowska, 1998 (Archibald & Averianov 2012). Premolars. The premolars are broken and are only represented by their alveoli (Fig. 5). Only three distinct alveoli anterior to M 1 are preserved in addition to the anteriormost large alveolus for the canine. They correspond to a two-rooted P 5 and a likely single-rooted P 4 (Figs 5; 6). Judging from the alveoli size, P 5 was a little smaller than M 1, and P 4 was much smaller than P 5. The P 5 alveoli are widely spaced and have a similar diameter. The alveolus of the single-rooted P 4 is half the size (diameter) of the anterior alveolus of P 5. It is also significantly lingually shifted in the dentary with respect to the anterior root of P 5, and it is located close to the mandibular symphysis. The anterior part of the dentary shows a small bony gap (hole) located lingually and in front of the P 4 alveolus (labelled “? ” in Fig. 7) which could be interpreted as a small tooth alveolus, either for an anterior root of P 4 or a small P 3 (as seen in early eutherians such as zhelestids). In the tomographies this structure is seen as a darker (empty) space corresponding to small vertical cavity in the bone (Fig. 7). However, similar and probably homologous bone cavities of the same size are also found between the roots of p 5, m 1 and m 2, positioned lingually to the line of the alveoli (Fig. 7 D, E). They correspond to bone vascularisation or pneumatization (air cells) rather than tooth alveoli. The CT scans do not show any trace of other alveoli anterior to P 4, other than the very large and procumbent one for the canine. Actually, there is no space available for the anterior premolars (P 1 - 3) between the alveoli of P 4 and C 1 (Figs 5; 7). This evidence, together with the mandibular symphysis extending close to P 4 in MNHN. F CTE 215, supports this as a natural condition representative of the species. As a result, the anterior cheek dentition of the Mas-d’Azil eutherian species is significantly and unexpectedly reduced, with only two premolars, one being reduced in size (single-rooted). Molars. Only M 1 and M 2 are preserved (the posterior molar, M 3, is broken). The molars are typically bunodont with inflated crown and cusps, and with large talonid and postfossid. They have two long separate roots that are wide transversely. There is typically a well-developed interradicular crest and a radicular groove on internal side of the root (Fig. 6), both for a firm bony anchorage of the tooth in the dentary. Interradicular crests are known in a few taxa of very different groups of mammals such as multituberculates, meridiolestidans, lipotyphlans (soricids and erinaceids) and rodents; however, the distribution of these roots features is poorly known in eutherians. The anterior root is smaller in diameter than the posterior one. The roots are robust and slightly dilated at their apex (Fig. 6). The ratio of the roots height to the crown height is 2.65 to 1.8. M 1 (Figs 3; 4). The trigonid is broken but the talonid is well preserved. The trigonid occlusal outline is quadrate and it is wider than it is long (trigonid mesio-distally narrow). The trigonid width is about equal to that of the talonid. The talonid is high, especially labially below the hypoconid. It is long: longer than wide, and much longer than the trigonid. The postfossid is extended. The hypoflexid is wide and shallow. There is no trace of ectocingulid and mesoconid. The hypoconid is by far the largest talonid cusp. It might have been at least as large as the protoconid. It has a well-marked lingual wear facet which corresponds to the lingual phase grinding wear with the protocone (facet 9 of Crompton 1971; facet 10 of Butler 1952; hd-l in figure of Schultz et al. 2018). This lingual phase wear facet is derived among stem eutherians that have mostly shearing teeth with restricted labial phase movements of mastication (Crompton 1971; Crompton & Kielan-Jaworowska 1978). The cristid obliqua is long and longitudinally oriented; it joins the trigonid in its labial mid-width. Its labial side shows a well-developed wear facet that extends low onto the hypoconid and displays subvertical striae for shearing with the postparacrista (facet 3 of Crompton 1971; facet 6 of Butler 1952; hd-mb in figure 2 of Schultz et al. 2018). The wear facet posterior to the hypoconid, corresponding to shearing with premetacrista (facet 4 of Crompton 1971; facet 7 of Butler 1952; hd-d in Schultz et al. 2018), is comparatively much smaller. The postcristid is reduced (weak and short) on the distal flank of the hypoconid. The hypoconulid has a distinctive ridge-like morphology instead of being cuspate: together with the postcristid it forms a blunt cingular-like structure at the distal rim of the talonid (Fig. 4). This cingular-like postcristid-hypoconulid structure is isolated by notches from both the hypoconid and the entoconid. It is more inflated on its lingual side. As a result, the hypoconulid is closer to the entoconid than to the hypoconid. However, it is not twinned with entoconid. The entoconid is inflated, acute at apex (mesio-distally compressed) and slightly higher than the hypoconulid. It is far distal from the trigonid and aligned transversely with the hypoconid. The postfossid is opened disto-labially between the hypoconid and hypoconulid by a notch bearing a large wear facet, possibly corresponding to a lingual extension of facet 7 of Butler (1952). The transverse extension of the wear facet 7 is usually seen in lophodont mammals such as perissodactyls; it corresponds to a specialized transverse shearing with the metaloph. There is a faint but distinct transverse crest at the bottom of the postfossid that is extended between the hypoconid and the entoconid (Fig. 4); it is in the topographic position of the hypolophid and it delimits a posterior fovea. It has a tiny distal wear facet extending on the entoconid, which corresponds to facet 8 of Butler (1952) (ed-d in figure 2 of Schultz et al. 2018), and to shearing with the hypocone. The mesio-lingual side of the entoconid is truncated by a wear facet (wear facet 9 of Butler 1952; ed-m in Schultz et al. 2018), resulting from shearing with the protocone (distal side). The postfossid is closed lingually (no talonid notch). There is a distinct and inflated entoconulid. The crown is only moderately exodaenodont above the roots. Observations of the reconstructed 3 D digital models of the teeth from CT scans shows the absence of postcingulid in M 1 (see Appendix 1). M 2 (Figs 3; 4). The crown of M 2 is nearly entirely broken. It preserves only part of the metaconid and the entoconulid. The metaconid was as high as the hypoconulid of M 1. The occlusal outline of the trigonid is very narrow mesio-distally, and more so than in M 1 (seeTable 1). The 3 D digital reconstruction from the CT scans supports the presence of a small paraconid in lingual position and closely appressed to the metaconid (Fig. 6 A; Appendix 1). The tooth occlusal outline (Figs 3; 4) and the 3 D models from CT scan indicate that M 2 is slightly longer than M 1. Dental occlusion and function Several features of lower molars of specimenMNHN. F CTE 215 suggest occlusion with upper molars bearing a developed hypocone: the cingular-like postcristid and hypoconulid, the hypolophid present and delimiting a posterior fovea, the postfossid opened between hypoconulid and hypoconid with a wear facet in between, and the likely presence of a wear facet 8 ofButler (1952). In particular, the hypolophid usually occludes in between the protocone and the hypocone, i. e., in the interloph. The bunodont molar construction, the large postfossid and hypoconid, the wear pattern including grinding wear facet 10 of Butler (1952), and the development of an interradicular crest and radicular grooves indicate strong power stroke and a developed crushing-grinding function, presumably corresponding to a (relatively hard) plant diet.	en	Gheerbrant, Emmanuel, Teodori, Dominique (2021): An enigmatic specialized new eutherian mammal from the Late Cretaceous of Western Europe (Northern Pyrenees). Comptes Rendus Palevol 20 (13): 207-223, DOI: 10.5852/cr-palevol2021v20a13, URL: https://doi.org/10.5852/cr-palevol2021v20a13
