identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03A387A8FFE08F58FB8B580EFE48F923.text	03A387A8FFE08F58FB8B580EFE48F923.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Abelisauridae Bonaparte & Novas 1985	<div><p>Abelisauridae indet.</p><p>MATERIAL EXAMINED. — France • 44 (fossil teeth); Hérault, Cruzy, Massecaps; 43°21’37.56”N, 2°57’17.46”E; “ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=2.95485&amp;materialsCitation.latitude=43.36043" title="Search Plazi for locations around (long 2.95485/lat 43.36043)">Valdo-Fuvélien</a> ”, “Grès à reptiles” Formation; Campanian-Maastrichtian; 118 m a.s.l; ACAP leg; M6, M7, M9, M11, M12, M13, M14, M15, M16, M17, M213, M237, M257, M468, M519, M674, M703, M808, M1010-3, M147, M1103, M1129, M1174, M1203, M1439 (reference specimen, Fig. 4), M1482, M1509, M1573, M1778, M1817, M1854, M1921, M1965, M2154, M2402, M2600, M3570, M4070, M4334, M4803, M5065-2, M5246 (Figs 2, 3), M5262, M5289.</p><p>LOCALITY. — Massecaps (43°21’37.56”N, 2°57’17.46”E); 1.3 km east of Cruzy, Hérault, France.</p><p>STRATIGRAPHY. — “Valdo-Fuvélien”, “Grès à reptiles” Formation.</p><p>AGE. — Campanian-Maastrichtian (Buffetaut et al. 1999).</p><p>REFERENCE SPECIMEN</p><p>M1439 (Fig. 4) has been chosen as the reference specimen of all this material because it has a complete crown and preserves almost all the denticles of the mesial and distal carinae, particularly the most apical ones (Fig. 4G).</p><p>DESCRIPTION</p><p>Tooth slightly elongated baso-apically and ziphodont in shape (sensu Hendrickx et al. (2020): “labiolingually compressed crown with serrated carinae and apically oriented apex”). Crown height ranges from 17 mm to 55 mm (average height = 37 mm ± 10 mm). The mesial margin of the crown is more recurved than the distal margin, which can be almost straight. In lateral view the mesial carina is convex and the distal carina is concave. The mesial carina ends in the region of the cervix for lateral tooth and more apically for more anterior teeth. Marginal undulations, flutes, longitudinal grooves, or ridges are absent. The enamel texture is irregular and smooth, and slightly oriented apico-basally (Fig. 2G). The distal carina has significantly wider denticles than the mesial carina (Table 1; Fig. 6). The average length of 10 denticles at mid-crown varies from 2.8 mm to 4.2 mm for the mesial carina and from 3.2 mm to 4.6 mm for the distal carina (3.4 ± 0.4 mm vs 3.9 ± 0.4 mm). Denticle length is fairly constant along the mesial and distal carina except for the apical-most and basal-most denticles. Denticles are inclined towards the tip of the crown and the major axis of the denticle is not perpendicular to the mesial margin of the crown. In lateral view denticles are parallelogram in shape. In mesial view, the denticles have a round to crescent-shaped contour; in distal view, the denticles are crescent-shaped. The interdenticular space is narrow. There is no interdenticular sulcus.</p><p>Tooth M5246</p><p>Although the apex is missing, the tooth appears to be slightly elongated baso-apically (30 mm without the apex) and ziphodont in shape (sensu Hendrickx et al. 2020: “labiolingually compressed crown with serrated carinae and apically oriented apex”). Comparison with other large theropod teeth from Massecaps, indicates that M5246, with its flat morphology as well as its rather straight and aligned carinae, is a lateral tooth. However, compared with M1439, M5246 has a more ovoid cross-section in basal view (Fig. 2F vs Fig. 4F), is less compressed laterally, has a mesial carina that ends more apically and mesial and distal carinae that appear straighter in mesial and distal views. This suggests a more anterior position in the jaw and that it is probably an antero-lateral tooth.</p><p>The denticles are poorly preserved in the mesial carina and completely broken on the distal carina, with the exception of the most basal part, where a few are preserved. The basal part of the tooth is damaged in the cervix region, and it is difficult to ascertain whether part of the root is preserved distally. The tooth bears a mark at the base of the lingual surface, close to the mesial edge (Figs 2 D-G; 3).</p><p>The mesial carina bears 18 denticles per 5 mm and does not show density variations along the preserved section of the carina, except for its most distal part. The distal carina is not preserved, with the exception of the most basal part, making it impossible to count the density of denticles.</p><p>Mark</p><p>The mark is located on the lingual face of the tooth and has affected the enamel surface and extends into the underlying dentine. The mark has two faces that define a bevelled profile. The face located most basally in the tooth is flat (Fig. 3B [F1]). The face located more apically bears slightly curved internal striations, parallel to each other and oriented perpendicular to the axis of elongation of the bevel (Fig. 3B [F2]). This basal face forms an angle with the tooth surface less important than the apical face. These striations are composed of a succession of nine grooves separated by eight wrinkles (Figs 2; 3). The mark has a well-marked V-shaped profile in cross-section. The complete mark measures 4 mm in the direction of bevel elongation. The width of seven consecutive grooves measures 3.0 mm and the mean width of one groove 0.4 mm (Table 1). Two enamel tears are visible (Fig. 3B [S1, S2]), one extending from the bevel termination located distally, the other parallel to the first just below (Figs 2; 3).</p><p>Denticle measurements</p><p>The mid-crown denticles of the crocodile tooth (M5085) are irregular in size, averaging less than 0.1 mm (Fig. 5A). For the teeth of Dromaeosauridae indet. (M324, M561), the average length of mid-crown denticle is 0.15 mm (Fig. 5 E-F). For Abelisauridae indet. teeth (M1439, M1778), the mean length of mid-crown denticle is 0.4 mm (Fig. 5 I-J; Table 1).</p><p>Furthermore, Abelisauridae indet. teeth have wider denticles on the distal carina than on the mesial carina (Mean denticles length calculated from all Abelisauridae indet. teeth equals 0.39 mm vs 0.34 mm; t test p -value = 0.0001845; Table 1). For instance, 10 denticles located on the mesial carina of M1439 measure 4 mm whereas ten denticles located on the distal carina measure 4.4 mm (Table 1). Denticle length is fairly constant along the mesial and distal carina except for the last circa ten most apical (and basal) denticles, whose respective lengths decrease significantly to 0.09 mm and 0.11 mm (Fig. 4G). The length of the denticles of the mesial and distal carinae is positively and significantly correlated with the total height of the crown (respective p -values = 0.0047 and 6e-04; Fig. 6).</p><p>DISCUSSION</p><p>TAXONOMIC ASSIGNMENT OF TOOTH M5246</p><p>Abelisaurids are common in Campanian and Maastrichtian sediments of Europe. Their presence has been noticed in Spain from the Late Campanian of the Laño and Armuña localities (Astibia et al. 1990; Pérez-García et al. 2016; Isasmendi et al. 2021; 2022) from the Late Campanian - Early Maastrichtian locality of Poyos (Malafaia et al. 2023, 2025) and from the latest Maastrichtian Arén and Tremp formations in Ribagorza’s country (Isasmendi et al. 2024). Their presence has also been reported multiple time in France. They are represented in the Early Campanian of Lambeau du Beausset with the species Tarascosaurus salluvicus Le Loeuff &amp; Buffetaut, 1991 (Le Loeuff &amp; Buffetaut 1991); but also in the Late Campanian in Velaux-Bastide Neuve and Fox-Amphoux (Tortosa et al. 2014), in Trets-La Boucharde, with the specimens known as “La Boucharde specimen” (Allain &amp; Pereda-Superbiola 2003; Carrano &amp; Sampson 2008), in Pourcieux (Buffetaut et al. 1988), and in the locality of Jas Neuf Sud, which yielded the species Arcovenator escotae Tortosa, Buffetaut, Vialle, Dutour, Turini &amp; Cheylan, 2014 (Tortosa et al. 2014).</p><p>Finally, the presence of teeth, cranial and postcranial fragmentary elements referred to an Abelisauridae indet.have been documented multiple times in the area of Cruzy (Buffetaut 2005; Buffetaut et al. 1999; Ősi &amp; Buffetaut 2011; Tortosa et al. 2014). Recently, Isasmendi et al. (2024) hypothesized that some teeth previously assigned to dromaeosaurids by Chanthasit &amp; Buffetaut (2009) might belong to abelisaurids.</p><p>Abelisauridae dentition is well known from numerous taxa. However, Arcovenator Tortosa, Buffetaut, Vialle, Dutour, Turini &amp; Cheylan, 2014 presents numerous unique features in its dentition such as a distal carina strongly deflected labially and a mesial carina gently curved mesio-lingually toward the base that ends well-above the cervix as well as an apex placed distal to the basalmost point of the distal margin of the crown (Hendrickx et al. 2020). M5246 has a broken and missing apex and a damaged distal carina. However, it shows a mesial carina that does not reach the cervix and a distal carina slightly convex similar to Arcovenator escotae teeth (Tortosa et al. 2014: fig. 4) and teeth referred to Arcovenator sp. from the Laño locality (Isasmendi et al. 2021; 2022). The tooth M5246 lacks some characteristics of Arcovenator such as a mesial carina gently curved mesio-lingually toward the base. The tooth M5246 and the 43 other teeth belonging to the same morphotype have mean denticle densities at mid-crown height of 12.8/ 5 mm and 14.7/ 5 mm for the mesial and distal carina, respectively, which are slightly lower than those reported for the three teeth attributed to A. escotae (MHNAix-PV. 2011.12.15; MHNAix-PV. 2011.12.20 and MHNAix-PV.2011.12.187). Contrarily to A. escotae, the denticles of the distal carina are longer than those of the mesial carina (Table 1 and Fig. 6). Apart from these morphological variations, we tentatively attribute M5246 tooth (as well as M1439 and M1778 teeth, that we selected and figured for comparison, and 41 other teeth [see Systematic palaeontology]) to Abelisauridae indet. due to their overall resemblance with the teeth of A. escotae and Arcovenator sp. from contemporaneous sediments of France and Spain, as well as the fact that Abelisauridae are, so far, the only medium to large size theropods known in Europe for this period.</p><p>MARK IDENTIFICATION</p><p>The parallel striations of this mark correspond to the definition of the ichnospecies Knethichnus parallelum Jacobsen &amp; Bromley, 2009 (Jacobsen &amp; Bromley 2009). This mark is the product of serration traces, which originate when a theropod tooth, or other denticulate tooth, was dragged across a bone (or other hard surface such as tooth) at an angle at which only the denticles on the tooth meet the bone. We therefore interpret this mark as a bite mark produced by a denticulate tooth. The internal striations of the mark correspond to the marks left by the denticles of the tooth (“denticles drag marks”; Rogers et al. 2003, 2007; Jacobsen &amp; Bromley 2009; Drumheller et al. 2020). The ridged shape of the wrinkles and the more rounded shape of the grooves (Figs 2; 3) correspond respectively to the interdenticular space between two consecutive denticles and to the apex of a denticle (Fig. 3B). This bite mark resembles certain bite marks on bones (cf. Rogers et al. [2003: fig. 3A, B]; Jacobsen &amp; Bromley [2009: fig. 3A]; Drumheller et al. [2020: fig. 2A]). It also resembles the bite mark left by Tyrannosauridae on Tyrannosauridae teeth (cf. Abler [1992: fig. 10E] and Jacobsen [1995: fig. pD52]). However, illustrations in these studies are of poor quality and do not allow for a clear view of this type of mark, particularly for detailed comparison with the one shown in this study.</p><p>The relative movement of the tooth is complicated to determine and could either have a first order direction either apico-basally or basoapically (Fig. 3D). The bevel, sub-perpendicular to the shaft, is due to the mesial or distal carina of another tooth meeting the labial surface of tooth M5246 and probably marks a change in the relative second order labiolingual component (or conversely; Fig. 3D).</p><p>PRODUCER ASSIGNMENT</p><p>Following the definition of the ichnospecies Knethichnus parallelum we compare this mark caused by the dragging of denticles to the different denticulate tooth morphotypes recovered in the locality of Massecaps, which belong to three taxa: small ziphosuchian teeth (cf. Company et al. 2005; Rabi &amp; Sebők 2015; Fig. 5A, B), theropod teeth attributed to Dromaeosauridae indet. (Fig. 5 C-F) and larger theropod teeth attributed to Abelisauridae indet. (Fig. 5 G-J). In the case of the ziphosuchian tooth, the denticles are irregular in size, averaging less than 0.10 mm (Fig. 5A). For the teeth of Dromaeosauridae indet., the average width of a denticle is 0.15 mm (Fig. 5E, F). For Abelisauridae indet. teeth, the average width of a denticle is 0.40 mm (Fig. 5I, J; Table 1). The width of the grooves (0.4 mm) observed in the bite mark of the tooth M5246 therefore matches perfectly that of the denticles of Abelisauridae indet. teeth recovered from the same locality. We therefore assign its producer to an Abelisauridae indet. Furthermore, the regression models carried out for the mesial (n = 20) and distal (n = 27) carinae also clearly suggest that the average width of the grooves is compatible with the width of denticles located on the distal carina (Fig. 6). Taken together, this implies that the bite mark observed on the tooth M5246 was applied by the distal carina of another tooth of Abelisauridae indet.</p><p>BEHAVIOURAL IMPLICATIONS</p><p>This type of bite mark has been reported on several teeth of Tyrannosauridae (Abler 1992; Jacobsen 1995; and Caleb M. Brown pers. comm.) but remains poorly documented in the literature. To our knowledge, this is the first reported occurrence of this type of mark on an Abelisauridae tooth.</p><p>The fact that the bite mark affects both enamel and underlying dentine suggests that relatively high pressure has been applied to the tooth surface. This is possible provided that the tooth was immobile at the time the pressure was applied. A first hypothesis would be that the tooth was firmly anchored in the jaw when the mark was produced. The orientation of the bevel (sub-perpendicular to the shaft of the tooth) and the parallel striations indicate that the tooth that produced the mark was sub-perpendicular to the M5246 tooth that bears it. This implies that the mark could not have been left by an upper jaw tooth on a lower jaw tooth (or vice versa) of the same individual. The bite mark then could have been produced by one individual on another. This could have been produced on the occasion of agonistic behaviours involving head-biting, which is widely documented in present-day and fossil vertebrates (Tanke &amp; Currie 1998 and reference therein; Chimento et al. 2019; Benoit et al. 2021; Iyoda &amp; Yanagihara 2024) and in particular within non-avian theropods (Tanke &amp; Currie 1998; Peterson et al. 2009; Brown et al. 2022). The definition of agonistic behaviour was initially proposed by Scott &amp; Fredericson (1951) and clearly defined by Scott (1966) as “a behavioural system composed of behaviour patterns having the common function of adaptation to situations involving physical conflict between members of the same species”. Even if access to food represents one of the most favourable theatres for the expression of these agonistic behaviours, we cannot exclude the hypothesis that this mark could have been produced during other types of social interaction involving agonistic behaviour (e.g., territorial occupation, sexual reproduction). Secondly, given that the taxonomic identification of these teeth in this study is limited to the family rank ( Abelisauridae indet.), we cannot be absolutely certain that the bite mark involves two individuals of the same species. Thus, if we follow the definition stricto sensu, which circumscribes agonistic behaviour only to individuals of the same species, we can only hypothesize that this bite mark is evidence of confamilial social interaction.</p><p>However, the location of the striations on the lingual face of the tooth, their orientation, implying that the jaws of two individuals meet at a 90° angle and the fact that the striations probably correspond to the denticles of a distal carina make this first hypothesis mechanically unlikely. Alternatively, this bite mark may be the result of cannibalism, already documented in another Abelisauridae, Majungasaurus crenatissimus Depéret, 1896, from the Late Cretaceous of Madagascar (Rogers et al. 2003, 2007). Cannibalistic behaviour has also been documented in other theropods, such as Utahraptor Kirkland, Gaston &amp; Burge, 1993 from the Early Cretaceous (Cedar Mountain Formation) of the United States (Britt et al. 2009), the Tyrannosauridae, including Tyrannosaurus rex Osborn, 1905, Daspletosaurus Russell, 1970 and Albertosaurus sarcophagus Osborn, 1905 (Longrich et al. 2010; Hone &amp; Tanke 2015; McLain et al. 2018; Dalman &amp; Lucas 2021; Coppock &amp; Currie 2023). Possible cannibalistic behaviour in Allosaurus Marsh, 1877 is also hypothesized from an Upper Jurassic theropod bite mark assemblage coming from the Mygatt-Moore Quarry (Drumheller et al. 2020). A Tyrannosauridae tooth embedded in the dentary of another Tyrannosauridae has also been reported from the Maastrichtian, Alberta, Canada, and is either interpreted as confamilial or cannibalistic scavenging or fatal agonism (Bell &amp; Currie 2010). Cannibalistic behaviour has also been reported in the theropod Coelophysis bauri Cope, 1889 (Nesbitt et al. 2006, and references therein), butNesbitt et al. (2006) refute previous demonstration of cannibalistic behaviour for this species and question the commonality of such behaviour among non-avian dinosaurs.However,cannibalism has been documented in many current predatory species such as the Komodo dragon ( Varanus komodoensis Ouwens, 1912) and in particular in taxa that bracket phylogenetically extinct non-avian theropods such as Crocodilia Owen, 1842 and Aves Linnaeus, 1758 (Roach &amp; Brinkman 2007 and references therein). Furthermore, several studies published after Nesbitt et al. (2006) have documented cannibalism among theropods (Rogers et al. 2007; Britt et al. 2009; Longrich et al. 2010; Hone &amp; Tanke 2015; McLain et al. 2018; Dalman &amp; Lucas 2021; Coppock &amp; Currie 2023). Both present and past data thus support the view of Roach &amp; Brinkman (2007) that intraspecific predation was also a significant factor in the biology of non-avian theropods.</p><p>Interestingly, present-day scavenging on large mammal carcasses focuses mainly on the most nutritious and easily accessible body areas in the anus and rib cage (White &amp; Diedrich 2012); although the taxa involved are not phylogenetically close to those discussed here. Skulls appear less targeted but may nevertheless bear bite marks (White &amp; Diedrich 2012; Diedrich 2014). Hone &amp; Watabe (2010) report a specimen of Saurolophus Brown, 1912 humerus bearing numerous bite marks left by a specimen of Tarbosaurus Maleev, 1955 . The authors show that the arrangement and type of bite marks are indicative of scavenging behaviour. As mentioned above a Tyrannosauridae tooth embedded in the dentary of another Tyrannosauridae may be interpreted as confamilial or cannibalistic scavenging (Bell &amp; Currie 2010).</p><p>However, the fact that tooth M5246 has no root suggests that it is probably a shed tooth, ruling out the possibility that the mark was caused by cannibalistic behaviour on a tooth still anchored in the jaw of a dead individual. Furthermore, at the locality of Massecaps, bones attributed to Abelisauridae indet. are relatively rare. They consist in one tibia and one maxilla out of more than 5000 vertebrate fossils inventoried (Authors pers. obs.). The 44 isolated teeth attributed to Abelisauridae indet. that have been recovered at the locality of Massecaps, including the tooth M5246, do not have the root preserved and therefore probably correspond for most of them to shed teeth (although it cannot be excluded that some teeth may have lost their roots during biostratinomic processes, prior to burial). The presence of relatively abundant rootless theropod teeth combined with the scarcity of other skeletal elements belonging to these same theropod taxa is a pattern frequently observed in Mesozoic bonebeds (e.g., Buffetaut &amp; Suteethorn 1989; Lauters et al. 2008; Britt et al. 2009; Bell &amp; Campione 2014; Eberth 2015; Evans et al. 2015; Fanti et al. 2015; Ullmann et al. 2017; Drumheller et al. 2020; Snyder et al. 2020; Allain et al. 2022). This is interpreted as the fact that, before they were buried, the carcasses attracted theropod dinosaurs that came to feed on them and lost teeth in the process. This is consistent with the presence of numerous fossilized bone remains of herbivorous dinosaurs ( Rhabdodontidae, Titanosauria and Nodosauridae) in the vertebrate fossil assemblage of the locality of Massecaps, some of which preserve bite marks that are currently being studied.</p><p>In view of these elements, it seems unlikely that the bite mark on this tooth is the product of cannibalistic (or at least confamilial) scavenging. Alternatively, in view of the paleontological and taphonomic context described in this study, the most parsimonious hypothesis is that this bite mark was inflicted by one individual of Abelisauridae indet. on one of its own shed teeth, which becomes stuck between two occluding teeth or between a tooth and a bone while the animal is feeding on a carcass. This is consistent with both published studies and numerous unpublished observations of tyrannosaurid teeth bearing tyrannosaurid bite marks (Abler 1992; Jacobsen 1995; Caleb M. Brown pers. comm.). These findings may be attributed to the high tooth replacement rate in theropods, combined with frequent tooth loss during feeding and an exceptionally strong bite force. As a result, shed teeth were occasionally bitten or crushed along with the prey, and could be pressed against hard part such as bones during the consumption of a carcass (Fig. 7).</p></div>	https://treatment.plazi.org/id/03A387A8FFE08F58FB8B580EFE48F923	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Boschetto, Damien;Maggia, Bruno;Brito, Léa De;Borgese, Julie;Clavel, Didier;Chenet, Jean-Pierre;Roques, Thomas;Sèbe, Stéphane;Veyssières, Jean-Marc;Goedert, Jean	Boschetto, Damien, Maggia, Bruno, Brito, Léa De, Borgese, Julie, Clavel, Didier, Chenet, Jean-Pierre, Roques, Thomas, Sèbe, Stéphane, Veyssières, Jean-Marc, Goedert, Jean (2026): Unusual Abelisauridae tooth mark on an Abelisauridae tooth from the Upper Cretaceous of Cruzy (Hérault, France): implications for feeding behaviours. Geodiversitas 48 (5): 69-85, DOI: 10.5252/geodiversitas2026v48a5
