Gastroliths (stomach stones) are known from many extant and extinct vertebrates, including dinosaurs. Reported here is the first unambiguous record of gastroliths in an ornithopod dinosaur. Clusters of small stones found in the abdominal region of three articulated skeletons of Gasparinisaura cincosaltensis were identified as gastroliths on the basis of taphonomic and sedimentologic evidence. The large number of stones found in each individual, their size, and the fact that Gasparinisaura cincosaltensis was herbivorous, all suggest that they were ingested as a result of lithophagy rather than accidental swallowing.
Gastroliths or geo-gastroliths sensu Wings (2007) are known in many taxa of extant and fossil vertebrates (Whittle and Everhart 2000). Gastroliths have been occasionally reported in non-avian dinosaurs (Wings 2004) but only few cases can withstand rigorous testing. The clusters of stones associated with the prosauropod Massospondylus (Bond 1955) and the sauropods Seismosaurus hallorum (Gillette 1994), Dinheirosaurus lounnhanensis (Bonaparte and Mateus 1999), and Cedarosaurus weiskopfae (Sanders et al. 2001), indicate the occurrence of gastroliths in at least some sauropodomorphs. Also, the presence of stones in the abdominal cavities of the maniraptoriform theropods Sinornithomimus dongi (Kobayashi et al. 1999; Kobayashi and Lü 2003), Shenzhousaurus orientalis (Ji et al. 2003), Caudipteryx zoui, and Caudipteryx dongi (Ji et al. 1998; Zhou et al. 2000; Zhou and Wang 2000) suggests that these non-avian theropods were lithophagic.
Among ornithischians, unambiguous evidence supporting lithophagy is known from the basal ceratopsian Psittacosaurus (Osborn 1923; You and Dodson 2004). Supposed gastroliths reported in several thyreophorans and ornithopods are possibly secondarily transported clasts (Currie 1997; Lucas 2000). Stones associated with remains of the ornithopods Edmontosaurus (“Claosaurus”) (Brown 1907) and Iguanodon (Rivett 1956) appear to lack the taphonomic features that would allow their identification as true gastroliths (Wings 2004).
A partial skeleton of Gasparinisaura cincosaltensis Coria and Salgado, 1996 (MUCPv 213), a basal ornithopod from South America collected in 1992, contains a cluster of stones in the abdominal region which are here identified as gastroliths. In addition, two new specimens of G. cincosaltensis (MCSPv 111 and MCSPv 112) provide additional evidence of stomach stones in this species. These stones represent the first unambiguous evidence for lithophagy in ornithopod dinosaurs.
Material and geologic setting
Three specimens of Gasparinisaura cincosaltensis, MUCPv 213, MCSPv 111, and MCSPv 112, were collected near the city of Cinco Saltos (Río Negro Province, Patagonia, Argentina) (Fig. 1), in mudstones and sandstones of the early Campanian Anacleto Formation, in the uppermost portion of the Neuquén Group (Ramos 1981; Dingus et al. 2000). MUCPv 213 (Fig. 2A) consists of a partial skeleton that includes cranial and postcranial elements (see Salgado et al. 1997 for a detailed anatomical description). A portion of the preserved elements (both incomplete humeri articulated with both radii and ulnae, several posterior dorsal ribs from both left and right sides) are included in a compacted, reddish mudstone, whereas the posterior ribs are imbedded in a medium to coarse sandstone. Nine of the left ribs appear to be preserved in their original position. The bone surfaces do not show any significant preburial or diagenetic damage.
MCSPv 111 (Fig. 3A) is the most complete, articulated specimen of Gasparinisaura cincosaltensis collected to date, lacking only the skull and most of the tail. The left ventrolateral side of the skeleton is exposed, with the left hind limb in flexion over the abdominal area. Several characters allow the assignment of MCSPv 111 to G. cincosaltensis (Coria and Salgado 1996; Salgado et al. 1997): (1) long pubic peduncle of ilium, (2) greater and lesser trochanter of femur fully fused to each other, (3) absence of anterior intercondylar groove in the femur, (4) tibial malleoli approximately at the same level, and (5) metatarsal II transversely compressed and anteroposteriorly developed. The relatively small size of MCSPv 111 (around 43 % of the G. cincosaltensis largest individual, MUCPv 213) suggests that it is a juvenile specimen. MCSPv 111 is preserved in friable, medium- to coarse-grained, quartzose sandstone.
Specimen MCSPv 112 (Fig. 3B) was found approximately 0.3 mm from MCSPv 111 (Leonardo Salgado, personal communication 2008). It consists of the skull, several dorsal and caudal vertebrae, and the pelvic girdle in articulation with both hind limbs. Several characters including the ascending process of jugal anteroposteriorly wide, chevrons in midcaudal region roughly triangular, anteroposteriorly expanded, and characters (1), (2), and (3) as in MCSPv 111 allow its asignment to Gasparinisaura cincosaltensis. MCSPv 112 is also a juvenile, even slightly smaller than MCSPv 111.
Near the town of Cinco Saltos, the Anacleto Formation is composed of two lithofacies: (1) cross-stratified sandstones and (2) reddish mudstones (Salgado et al. 1997; Heredia and Salgado 1999). MUCPv 213 was found in the mudstones, and MCSPv 111 and MCSPv 112 in the sandstones. Field observations of both lithofacies showed no accumulations of stones similar to those contained inside the specimens of Gasparinisaura cincosaltensis.
Most of the following description is based on specimen MUCPv 213, from which the stones were removed and examined in a scanning electron microscope (Philips SEM 515). The stones found in specimens MCSPv 111 and MCSPv 112 were studied in situ.
All three Gasparinisaura cincosaltensis individuals were found with stone clusters. In MUCPv 213, a cluster of stones occupies about 70 cm2 in the rib cage on the left side of its abdominal area. Several stones are flanked by both left and right dorsal ribs, preserving several contacts between bones and the other stones (Fig. 2B, C). There are no stones outside the abdominal region on the lateral side of the rib cage. In total, 180 stones were counted (only stones larger than 4 mm in diameter were considered). The average length of the stones was 7.9 (±2.5) mm and the length of the largest stone was 17 mm (Fig. 2B). The total mass of the stones was 51.1 g.
Most of the stones were of igneous rock (47.22%). Other materials include quartz (27.22%), sandstone (23.89%), and three were composed of metamorphic rock (1.67%). Stones ranged from light brown and light gray to dark brown or nearly black in color. Following the roundness classification proposed by Powers (1953), the stones are mainly sub-rounded with dull surfaces. Sphericity (Blatt et al. 1972) varied from 0.51 to 0.91 with an average of 0.73 (±0.08). The 10 stones analyzed with SEM show smooth surfaces and a lack of pits or grooves as observed in other vertebrate gastroliths (Fig. 2D) (Whittle and Oronato 2000).
A conspicuous, three-dimensional accumulation of stones was exposed during preparation in the abdominal region of MCSPv 111. The cluster is located caudal to the rib cage and anteroventral to the postpubic processes (Fig. 3A). More than 40 rounded and sub-rounded, igneous stones can be seen in the cluster (few stones in contact with bones), all accumulated within an area of approximately 11 cm2.
In MCSPv 112, two clusters of stones were found. The largest cluster is located anteroventral to the pelvic girdle, underneath the last dorsal vertebra and the first sacral vertebra (Fig. 3B). Many stones of this cluster are in contact with ribs and vertebrae. The smallest accumulation is located in front of the main cluster, near the skull. Both accumulations seem to be part of a major cluster, partially lost by weathering. As in MCSPv 111, the stones were only observed inside the abdominal region.
The inorganic nature of gastroliths makes their identification difficult. Stones associated with bones are identifiable as gastroliths if they are clustered in the abdominal area of relatively complete skeletons (Wings 2004: 113). In addition, Wings (2004: 84) proposed a series of detailed taphonomic and sedimentologic criteria for identification of gastroliths. Many of these criteria are met by the rocks associated with the three specimens described here: (1) all the stones were found in situ in the abdominal area of the skeletons, like in MUCPv 213 and MCSPv 112, the stones were inside the rib cage; (2) the specimens are almost complete (MCSPv 111 and MCSPv 112) and articulated (MUCPv 213, MCSPv 111 and MCSPv 112); (3) there is a direct association between bones and stones; (4) the stones were found in clusters; (5) the matrix that supports each specimen lacks similar stones, isolated or forming clusters; (6) there were no isolated stones forming accumulations in the outcrops where the skeletal remains were collected; (7) there are three individuals of the same species with rocky accumulations in the gut area. Hence, the most parsimonious explanation for the stones is to consider them as true gastroliths (category 1 in the taphonomic classification of gastroliths finds proposed by Wings [2004: 113]).The presence of gastroliths in Gasparinisaura cincosaltensis is the first unequivocal evidence for lithophagy in Ornithopoda, thus increasing the number of documented cases of lithophagy among the Dinosauria.
The large number of gastroliths observed in each individual, their size, and the fact that Gasparinisaura cincosaltensis was a herbivorous dinosaur (Norman et al. 2004), indicate that the stones were ingested as a result of lithophagic behaviour rather than accidental swallowing. Keeping in mind all possible functions of gastroliths in vertebrates (Wings 2007), the most probable utility of these stones in Gasparinisaura seems to have been the trituration of plant materials, perhaps because the food was especially tough and thus needed trituration to make it digestible. This function was certainly linked to the presence of a gizzard, where food is mechanically broken down as in many extant birds (Gionfriddo and Best 1999). However, the existence of a gastric mill in sauropod dinosaurs has recently been denied, since the amount of gastroliths reported in these dinosaurs was too small (0.03% of their body mass) to support a gastric mill as present in extant birds (Wings and Sander 2007). In MUCPv 213, the total mass of gastroliths (51.1 g) amounts to about 0.3% of the body mass which is estimated (based on the femur circumference of 71 mm) as 18 kg (Anderson et al. 1985). Thus the relative mass of gastroliths in MUCPv 213 is much higher than that estimated for sauropod dinosaurs, but similar to that calculated for several extant birds (Wings and Sander 2007). If Gasparinisaura utilized gastroliths in the same way as in living birds, the amount of gastroliths would have been sufficient to support a gastric mill.
This work summarizes a part of my unpublished thesis at the Universidad Nacional del Comahue, Centro Regional Universitario Bariloche, on the gastroliths from South American ornithopods. I thank my graduate advisors Rodolfo Coria (Museo Carmen Funes, Neuquén, Argentina) and Gabriela Cusminsky (Centro Regional Universitario de Bariloche, Río Negro, Argentina) for their constant support, Olga Hevia (Museo Cinco Saltos, Río Negro, Argentina) for access to specimens under her care, Alberto Garrido (Museo Carmen Funes) for geological help in the field, and Raul Bollatti (Ente para el Desarrollo de la Margen Sur, Río Negro, Argentina) and Marcos Poblete for help with excavation work. Oliver Wings (Universität Tübingen, Tübingen, Germany), Mike Everhart (Sternberg Museum of Natural History, Hays, Kansas, USA), and Zhonge Zhou (Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China) provided information on gastroliths in vertebrates. Rodolfo Coria, Leonardo Salgado (Museo Universidad Nacional de Comahue, Neuquén, Argentina), Spencer Lucas (New Mexico Museum of Natural History, New Mexico, USA), and Mike Everhart provided useful comments on an early draft of the manuscript. Peter Galton and Oliver Wings provided constructive reviews. Specimens MCSPv 111 and MCSPv 112 were found by Walter Lagos and Sara Poblete (Cinco Saltos, Río Negro, Argentina). Carlos Cottaro (Centro Atómico de Bariloche, Río Negro, Argentina) provided assistance with the SEM.
- J.F. Anderson , A. Hall-Martin , and D.A. Russell 1985. Long-bone circumference and weight in mammals, birds and dinosaurs. Journal of Zoology 207: 53–61. Google Scholar
- R.R. Andreis , A.M. Ińiguez Rodriguez , J.J. Lluch , and D.A. Sabio 1974. Estudio sedimentológico de las Formaciones del Cretácico Superior del área del Lago Pellegrini (provincia de Río Negro, República Argentina). Revista de la Asociación Paleontólogica Argentina 29: 85–104. Google Scholar
- H. Blatt , G. Middleton , and R. Murray 1972. Origin of Sedimentary Rocks. 634 pp. Prentice-Hall, New Jersey. Google Scholar
- J.F. Bonaparte and O. Mateus 1999. A new diplodocid, Dinheirosaurus lourinhanensis gen. et sp. nov., from the Late Jurassic beds of Portugal. Revista del Museo Argentino de Ciencias Naturales 5: 13–29. Google Scholar
- G. Bond 1955. A note on dinosaur remains from the Forest Sandstone (Upper Karoo). Occasional Papers of the National Museum of Rhodesia 2: 795–800. Google Scholar
- B. Brown 1907. Gastroliths. Science 25: 392. Google Scholar
- R.A. Coria and L. Salgado 1996. A basal iguanodontian (Ornitischia: Ornithopoda) from the Late Cretaceous of South America. Journal of Vertebrate Paleontology 16: 445–457. Google Scholar
- P.J. Currie 1997. Gastroliths. In: P.J. Currie and K. Padian (eds.), Encyclopedia of Dinosaurs, 270. Academic Press, San Diego. Google Scholar
- L. Dingus , J. Clarke , G.R. Scott , C.C. Swisher III , L.M. Chiappe , and R.A. Coria 2000. Stratigraphy and magnetostratigraphic/faunal constraints for the age of sauropod embryo-bearing rocks in the Neuquén Group (Late Cretaceous, Neuquén Province, Argentina). American Museum Novitates 3290: 1–11. Google Scholar
- D.D. Gillette 1994. Seismosaurus, the Earth Shaker. 205 pp. Columbia University Press, New York. Google Scholar
- J.P. Gionfriddo and L.B. Best 1999. Grit use by birds—a review. Current Ornithology 15: 89–148. Google Scholar
- S. Heredia and L. Salgado 1999. Posición estratigráfica de los sustratos supracretácicos portadores de dinosaurios en Lago Pellegrini, Patagonia septentrional, Argentina. Ameghiniana 36: 229–234. Google Scholar
- Q. Ji , P.J. Currie , M.A. Norell , and S.-A. Ji 1998. Two feathered dinosaurs from northeastern China. Nature 393: 753–761. Google Scholar
- Q. Ji , M.A. Norell , P.J. Makovicky , K.-Q. Gao , S. Ji , and C. Yuan 2003. An early ostrich dinosaur and implications for ornithomimosaur phylogeny. American Museum Novitates 3420: 1–19. Google Scholar
- Y. Kobayashi and J.-C. Lu 2003. A new ornithomimid dinosaur with gregarious habits from the Late Cretaceous of China. Acta Palaeontologica Polonica 48: 235–259. Google Scholar
- Y. Kobayashi , J.-C Lu, Z.-M. Dong , R. Barsbold , Y. Azuma , and Y. Tomida 1999. Herbivorous diet in an ornithomimid dinosaur. Nature 402: 480–481. Google Scholar
- S.G. Lucas 2000. The gastromyths of “Seismosaurus”, a Late Jurassic dinosaur from New Mexico. In: S.G. Lucas and A.B. Heckert (eds.), Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science Bulletin 17: 61–68. Google Scholar
- D.B. Norman , H.-D. Sues , L.M. Witmer , and R,A. Coria 2004. Basal Ornithopoda. In: D.B. Weishampel , P. Dodson , and H. Osmólska (eds.), The Dinosauria, Second Edition, 393–437. University of California Press, Berkeley. Google Scholar
- H.F. Osborn 1923. Two Lower Cretaceous dinosaurs of Mongolia. American Museum Novitates 95: 1–10. Google Scholar
- M.C. Powers 1953. A new roundness scale for sedimentary particles. Journal of Sedimentary Petrology 23: 117–119. Google Scholar
- V.A. Ramos 1981. Description geológica de la Hoja 33 c. Los Chihuidos Norte, Provincia del Neuquén. Servicio Geológico Nacional, Boletín 182: 1–103. Google Scholar
- W.H.E. Rivett 1956. Reptilian bones from the Weald Clay. Proceedings of the Geological Society of London 1540: 110–111. Google Scholar
- L. Salgado , R.A. Coria , and S. Heredia 1997. New materials of Gasparinisaura cincosaltensis (Ornithischia: Ornithopoda) from the Upper Cretaceous of Argentina. Journal of Paleontology 71: 993–940. Google Scholar
- F. Sanders , K. Manley , and K. Carpenter 2001. Gastroliths from the Lower Cretaceous sauropod Cedarosaurus weiskopfae. In: D.H. Tanke and K. Carpenter (eds.), Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie, 166–180. Indiana University Press, Bloomington. Google Scholar
- C.H. Whittle and M.J. Everhart 2000. Apparent and implied evolutionary trends in lithophagic vertebrates from New Mexico and elsewhere. In: S.G. Lucas and A.B. Heckert (eds.), Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science Bulletin 17: 75–82. Google Scholar
- C.H. Whittle and L. Oronato 2000. On the origin of gastroliths determining the weathering environment of rounded and polished stones by scanning-electron-microscope examination. In: S.G. Lucas and A.B. Heckert (eds.), Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science Bulletin 17: 69–74. Google Scholar
- O. Wings 2004. Identification, Distribution, and Function of Gastroliths in Dinosaurs and Extant Birds with Emphasis on Ostriches (Struthio camelus). 187 pp. Published Ph.D. thesis, University of Bonn, Bonn. (Accessible online at http://nbn-resolving.de/urn:nbn:de:hbz:5N-04626). Google Scholar
- O. Wings 2007. A review of gastrolith function with implications for fossil vertebrates and a revised classification. Acta Palaeontologica Polonica 52: 1–16. Google Scholar
- O. Wings and P.M. Sander 2007. No gastric mill in sauropod dinosaurs: new evidence from analysis of gastroliths mass and function in ostriches. Proceedings of The Royal Society 274: 635–640. Google Scholar
- H.-L. You and P. Dodson 2004. Basal Ceratopsia. In: D.B. Weishampel , P. Dodson , and H. Osmólska (eds.), The Dinosauria, Second Edition, 478–493. University of California Press, Berkeley. Google Scholar
- Z. Zhou and X.-L. Wang 2000. A new species of Caudipteryx from the Yixian Formation of Liaoning, northeast China. Vertebrata PalAsiatica 38: 111–127. Google Scholar
- Z. Zhou , X. Wang , F. Zhang , and X. Xu 2000. Important features of Caudipteryx: evidence from two nearly complete new specimens. Vertebrata PalAsiatica 38: 241–254. Google Scholar