AMEGHINIANA (Rev. Asoc. Paleontol. Argent.) - 47 (3): 387-400. Buenos Aires, 30-09-2010 ISSN 0002-7014 Predation bite-marks on a peirosaurid crocodyliform from the Upper Cretaceous of Neuquén Province, Argentina Lucas Ernesto FIORELLI1 Abstract. Bite marks and injuries caused by intraspecific predation or aggressive social behavior are un- common in fossil vertebrates, and in the fossil record of suchian archosaurs they are extremely unusual. A peirosaurid crocodyliform collected from the Upper Cretaceous beds of the Neuquén Group (northern Patagonia, Argentina) shows abundant bite marks and injuries. These injuries are spread across all the preserved parts of the specimen, with the highest concentration of bite marks, perforations and breakage in the caudal region. Characteristics of these injuries are analyzed, and their possible origin and related taphonomic aspects are assessed. Results indicate that the injuries were not produced by intraspecific figh- ting, but probably by the predating action of a theropod dinosaur. Resumen. MARCAS DE PREDACIÓN EN UN CROCODILIFORME PEIROSAURIDO DEL CRETÁCICO SUPERIOR DEL NEU- QUÉN, ARGENTINA. El hallazgo de marcas de mordeduras y lesiones por predación o comportamiento so- cial agresivo intraespecífico en vertebrados fósiles es poco frecuente. En el registro fósil de arcosaurios su- quianos tales evidencias son extraordinarias. Aquí se reporta un crocodiliforme peirosáurido del Cretácico Superior del Grupo Neuquén (norte de Patagonia, Argentina) con abundantes marcas de mordeduras y lesiones. Estas heridas están localizadas en todas las partes preservadas del espécimen, con una alta con- centración de mordeduras, perforaciones y roturas en la región caudal. En el presente trabajo, se analizan las características de las lesiones, y también sus posibles causas y los aspectos tafonómicos relacionados. Los resultados muestran que las mordeduras fueron hechas por la acción de un predador y no por peleas intraespecíficas. Key words. Bite marks. Predation. Peirosaurid. Upper Cretaceous. Neuquén. Palabras clave. Mordeduras. Predación. Peirosáurido. Cretácico Superior. Neuquén. Introduction specimen of the crocodyliform Baurusuchus pachecoi Price, 1945, from the Adamantina Formation, Bauru Bite marks and injuries produced by predation Basin (Brazil), was recently reported (Avila et al., and scavenging in vertebrate fossils are uncommon 2004). These authors argued that such bite marks and only a few cases are well documented (Erick- were produced by cospecific combat. son and Olson, 1996; Chure et al., 1998; Frey et al., Ethology and social behavior of extant crocody- 2002; Everhart, 2005; Reisz and Tsuji, 2006; Bianucci lians is more complex than that of other non-archo- et al., 2010; Hone et al., 2010). Even less common are saurian reptiles (Avila et al., 2004). Many ethologi- cases in which the bite injuries are caused by intras- cal traits, a combination of sensory signals (olfac- pecific and cofamilial aggressive social behavior tory, visual, and acoustic), and several corporal (Molnar and Farlow, 1990; Sereno and Novas, 1993; postures and social displays are used as warnings Tanke and Currie, 1998; Bell and Currie, 2009). Ca- before escalation to physical intraspecific aggres- ses applicable to crocodylomorphs are also limited, sion occurs (Hunt, 1977; Garrick et al., 1978; Vliet, with only a few examples recorded in fossil mem- 1989; Thorbjarnarson and Hernández, 1993; Pough bers of this group (Gilmore, 1946; Buffetaut, 1983; et al., 1998; Senter, 2008). Injuries are caused fre- Sawyer and Erickson, 1998; Frey et al., 2002). A case quently when for some reason —e.g., disputes for of injuries and bite marks produced on the tail of a food or territory, dominance fights during repro- ductive period and courtship behavior, etc.— figh- ting takes place (Cott, 1961; Hunt, 1977; Vliet, 1989, 2001a, 2001b; Senter, 2008). Adult or subadult ex- 1Centro Regional de Investigaciones Científicas y Transferencia tant crocodilians have virtually no natural preda- Tecnológica de La Rioja, Entre Ríos y Mendoza s/n, 5301 Anillaco, tors. However, crocodilian nests, neonates and ju- La Rioja, Argentina. [email protected] veniles are predated by mammals, birds, fishes, ©Asociación Paleontológica Argentina AMGHB2-0002-7014/10$00.00+.50 388 L.E. Fiorelli snakes, turtles, and even other crocodilians (Scott, arid and dry continental climate was dominated by 2004). As adult members of this group represent the aeolian deposition, creating big dunes and inter-du- top predators in their trophic environments, cases ne lagoon basins skirted by fluvial deposits, criss- in which they have been attacked by other preda- crossed by streams and seasonal or ephemeral wa- tors have been very rarely recorded. ter bodies. The horizon in which the fossil was Herein I report a peirosaurid crocodyliform from found is composed of coarse-grained, light violet to the Santonian Bajo de la Carpa Formation in Neu- pink sandstones of fluvial origin. quén Province, Argentina (Bonaparte, 1991; Fiorelli et al., 2007), with numerous injuries, perforations, breakage, and bite marks. These injuries are present Fauna of the Bajo de la Carpa Formation throughout most of the preserved parts of the speci- men being more frequent in the caudal section. Various outcrops of the Bajo de la Carpa For- mation have yielded a rich fossil vertebrate assem- blage (figure 2.2), including the remains of an inde- Material and methods terminate titanosaurid (Chiappe and Calvo, 1994; Parras and González Riga, 2001), cf. Laplatasaurus The specimen described here, MUCPv-27 (figure Huene, 1929 (Leanza et al., 2004), and the peculiar 1.1-1.8), represents a new species of a still unnamed beaked sauropod Bonitasaura salgadoi Apesteguía, peirosaurid crocodyliform (Fiorelli et al., 2007). 2004, found at the locality known as La Bonita, in Río Bonaparte (1991) briefly described this specimen and Negro Province. The theropod dinosaur fauna is al- suggested that it would represent a taxon closely re- so diverse, including the small abelisauroid thero- lated to Araripesuchus Price, 1959. Nevertheless, a re- pod Velocisaurus unicus Bonaparte, 1991, found in cent review of MUCPv-27 stated that it would not re- Neuquén (also in the Campus of the University of present a form closely related to the Notosuchia, but Comahue). Recently, the presence of carnotaurine instead exhibits affinities with neosuchian peirosau- abelisaurid theropods was reported from the top of rids (Fiorelli et al., 2007). Available fragmentary re- the unit in the La Invernada region , in northern mains include many elements in good state of pre- Neuquén (Porfiri and Calvo, 2006), and also at Paso servation but most of the fossil was lost (probably Córdova (Río Negro Province) (Ezcurra and owing to predation). Preserved elements of MUCPv- Méndez, 2009). Aditionaly, remains of indetermina- 27 (figure 1.1-1.8) include cranial remains, nine dor- te theropods coming from the Auca Mahuida region sal vertebrae, some articulated appendicular ele- (Neuquén Province) were considered by Coria and ments, numerous presacral osteoderms, and an al- Arcucci (2004) to be large basal tetanurans related to most complete caudal region that is fully enclosed by carcharodontosaurids. Porfiri et al. (2008) reported osteoderms. Materials were photographed, drawn, from Tratayén (Neuquén Province) an articulated and different digital programs were used for the postcranium belonging to an allosauroid related to analysis and design of the images and to highlight the Carcharodontosauria (sensu Benson et al., 2009). specific anatomical aspects. Coelurosaurian dinosaurs are also common and re- latively well-known, such as the alvarezsaurids Abbreviations. MUCP, Museo de Geología y Paleontología, Alvarezsaurus calvoi Bonaparte, 1991 (Campus of the Universidad Nacional del Comahue, Neuquén, Argentina. University of Comahue) and Achillesaurus manazzo- MUCPv-27 is currently housed at the Centro Paleontológico Lago Barreales -CePaLB-, State Route #51, Km 65, Neuquén Province, nei Martinelli and Vera, 2007 (Paso Córdova), the Argentina. enantiornithine Neuquenornis volans Chiappe and Calvo, 1994, and the basal ornithuromorph Patagopteryx deferrariisi Alvarenga and Bonaparte, Locality and horizon 1992 (both in the Campus of the University of Comahue). Also, bird egg-clutches with embryos Specimen MUCPv-27 comes from beds exposed (Schweitzer et al., 2002) (Campus of the University of within the Campus of the University of Comahue, Comahue), dinosaur eggs related to Megaloolithus pa- in the city of Neuquén (38º56’25”S - 68º3’10”W) (fi- tagonicus Calvo et al., 1997 (south of Neuquén city), gure 2.1). In this area, the outcrops are referred to and remains of the snake Dinilysia patagonica Wood- the Bajo de la Carpa Formation, Río Colorado ward, 1901 (Campus of the University of Comahue). Subgroup, Neuquén Group (middle-upper In addition, the chelid Lomalatachelys neuquina Lap- Santonian; Leanza et al., 2004) (figure 2.2). Unlike ot- parent de Broin and de la Fuente, 2001, comes from her sections in the basin, the rocks of this formation Loma de la Lata, in Neuquén Province (de la Fuente, in the city of Neuquén were deposited in fluvial and 2007) and constitutes a common faunal element. aeolian systems (Sánchez et al., 2006). Moreover, the Crocodyliforms are an important faunal compo- AMEGHINIANA 47 (3), 2010 Predation on a Cretaceous peirosaurid crocodyliform 389 Figure 1. Some elements of the MUCPv-27 peirosaurid. 1 and 2, caudal