Pterosaurs as a food source for small dromaeosaurs David Hone, Takanobu Tsuihiji, Mahito Watabe, Khishigjaw Tsogtbaatr PII:

DOI: Reference: To appear in: Received date: Revised date: Accepted date: S0031-0182(12)00094-6 doi: 10.1016/j.palaeo.2012.02.021 PALAEO 6050 Palaeogeography, Palaeoclimatology, Palaeoecology 11 November 2011 6 February 2012 14 February 2012

Please cite this article as: Hone, David, Tsuihiji, Takanobu, Watabe, Mahito, Tsogtbaatr, Khishigjaw, Pterosaurs as a food source for small dromaeosaurs, Palaeogeography, Palaeoclimatology, Palaeoecology (2012), doi: 10.1016/j.palaeo.2012.02.021

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Pterosaurs as a food source for small dromaeosaurs

1. School of Biology and Environmental Sciences, University College Dublin, Dublin 4, Ireland.

2. Department of Geology and paleontology, National Museum of Nature and Science, 3231 Hyakunincho, Shinjukuku, Tokyo 1690073, Japan. 3. Hayashibara Museum of Natural Sciences, 2-3, Shimoishii-1, Okayama 700-0907, Japan.

4. Paleontological Center, Mongolian Academy of Sciences, Ulaanbaatr, 210351, Mongolia.

Abstract

Stomach contents preserved in fossil specimens provide direct evidence for the diet of extinct animals. Such exceptional fossils remain rare for predatory non-avian dinosaurs and each can add significantly to our understanding of trophic interactions between various taxa. Here we present evidence for the dromaeosaurid theropod Velociraptor scavenging on the carcass of an azhdarchid pterosaur, with a long bone of the pterosaur being found as gut contents of the dinosaur. Despite previous inferences of dromaeosaurs as hyper-predators, scavenging appears to have been an important part of their ecology.

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David Hone*1, Takanobu Tsuihiji2, Mahito Watabe3, Khishigjaw Tsogtbaatr4.

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Keywords:

deinonychosaur, azhdarchid, scavenger, predator-prey, Cretaceous

1. Introduction

The preserved gut contents of fossil carnivorous dinosaurs provide direct evidence for their diet and help to establish trophic patterns, species interactions and the ecology of both taxa concerned (e.g. Charig and Milner, 1997; Varricchio, 2001). Such specimens are known for a variety of predatory non-avian theropod dinosaurs, though they are rare (see Hone and Rauhut, 2010). However, while these and similar ichnological records (e.g. bite marks, coprolites) do provide direct evidence of

and scavenging without exceptional evidence (e.g. Hone and Watabe, 2010). In general non-avian carnivorous theropods (and especially deinonychosaurs) have been considered active predators (Holtz, 2003), however evidence for scavenging in dromaeosaurs (Currie and Jacobsen, 1995; Hone et al., 2010) is now known. This should be no surprise few amniote carnivores are exclusively predatory or carnivorous and even the most dedicated carnivores will not turn down a free meal in the form of a carcass. Thus the debate of whether or not any given taxon was a predator or scavenger has moved on (e.g. Holtz, 2008) - the issue is where on the continuum between these extremes a given taxon may lie, and what evidence is

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carnivore-prey interactions it can be difficult to distinguish between active predation

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available to support this inferred trophic position. Here a specimen of the dromaeosaurid Velociraptor is described, preserved with

azhdarchid pterosaur and suggests that small non-avian deinonychosaurs were capable of consuming relatively large bones. While not the first evidence of theropod feeding on a pterosaur carcasss (Currie and Jacobsen, 1995; Buffetaut et al., 2004) this is the first known as gut contents.

2. Institutional Abbreviations

MPC-D: Registered number of dinosaur specimens stored at Paleontological Laboratory of Paleontological Center, Mongolian Academy of Science, Ulaanbaatar.

The skeleton was discovered in 1994 in the middle part of the geological section of the eolian sandstone complex at Tugrikin Shireh in the Gobi Desert, Mongolia. The eolian beds are 30 m thick, measured from the base of the cliff to the top and thus the fossil was recovered approximately 15 m from the base of the cliff. The cliff is south facing, forming the southern end of the mesa-like structure in Tugrikin Shireh. The locality is rich in dinosaur and other vertebrate fossils and the sedimentation is solely eolian in nature. We do not provide GPS co-ordinates owing to the prevelance of illegal fossil excavation at the site details are available on request. This locality is correlated with the Djadokhta Formation and thus of Campanian age (e.g.,

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3. Locality Information

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part of a bone in the chest cavity. This element can be identified as belonging to an

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Jerzykiewicz and Russell, 1991).

The specimen is of a largely complete and articulated skeleton of the velociraptorine dromaeosaurid Velociraptor mongoliensis (MPC-D100/54), missing the right forelimb and most of the tail (this material will be described fully in a future publication). Five dromaeosaurid species are currently recognized from the Djadokhta Formation or coeval sediments in the Gobi Desert of Mongolia and China: Velociraptor mongoliensis, V. osmolskae, Tsaagan mangas, Linheraptor exquisitus, and Mahakala omnogovae. With a skull length of approximately 230 mm, MPC-D100/54 is considerably larger than Mahakala (Turner et al., 2007) and this can be eliminated from further consideration. MPC-D100/54 lacks autapomorphies used

lacks the elongated maxillary fenestra and elongated promaxillary fenestra that are characteristic of V. osmolskae (Godefroit et al., 2008), the elongated basipterygoid process and pendulous paroccipital process without distal twisting observed in T. mangas (Norell et al., 2006), or an enlarged maxillary fenestra diagnostic of L. exquisitus (Xu et al., 2010). The skull of MPC-D100/54 does however bear a longitudinal ridge dorsal to a row of neurovascular foramina in the maxilla (although only in the posterior part of the bone), which is considered diagnostic for V. mongoliensis, (Barsbold and Osmlska, 1999). Accordingly, MPC-D100/54 is referred to V. mongoliensis.

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to diagnose V. osmolskae, T. mangas, or L. exquisitus. For example, MPC-D100/54

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4. Description

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The specimen has undergone extensive preparation, and numerous elements have been separated from the matrix, although the chest cavity (dorsal vertebrae, ribs,

articulated piece (Fig. 1). The animal was injured or recovering from an injury at the time of death with one broken dorsal rib showing signs of regrowth.

The specimen represents a young individual, perhaps a sub-adult, based on the incomplete fusion of the right scapula and coracoid, the separate sternal plates, and that the sacral ribs are not fully fused to the ilia. The femora are 192 and 194 mm long (left and right respectively) compared to a length of nearly 240 mm measured from an adult specimen of V. mongoliensis (Norell and Makovicky, 1999).

4.1 Gut contents

14). The anteriorly positioned part is fragmentary, with a more complete posterior part. The two parts lie in a direct line with one another and as far as can be determined are identical in bone wall thickness and have similar shapes, strongly suggesting that this was originally all part of a single piece. The anteriormost part of the bone lies below the fourth dorsal of the Velociraptor and the posterior piece below the fifth through seventh dorsals (Fig. 2). While the chest has collapsed somewhat, the relative positions of the bones have not been affected as this collapse is purely vertical (i.e. mediolateral directions of the body) and not lateral. The current position of this element is therefore considered genuine or close to the original position and thus

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Part of a single bone lies preserved in the chest cavity of the Velociraptor (Figs.

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sternal plates, gastralia, right scapula and both coracoids) is retained as a single

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would in life have been present in the upper part of the gastrointestinal tract, most likely the stomach. It is improbable that this entered the area post-mortem given the

to this being an eolian deposit.

The preserved bone would have been approximately 75 mm in length including the broken proximal part and 12 mm in diameter (only approximate values were obtained owing to the position of the overlying ribs and sterna). The bone is slightly oval in cross section and has a very thin cortex of approximately 0.2 mm in thickness (as measured from photographs owing to the inaccessibility of the material within the chest cavity).

The surface of the bone is smooth and in good condition, showing no unusual traces of marks or deformation that could be attributed to digestive acids. The edges

breakage occurred before, or as part of, ingestion. This also indicates, that the bone had not lain long in the gut as it would have been broken up, or at least the rough edges would be smoothed out. This also therefore suggests that the individual died shortly after ingesting the bone.

5. Discussion 5.1 Identity of the pterosaurian element The bone preserved in the Velociraptor chest cavity is here identified as that of a pterosaur. The extremely thin walled nature of the bone is unique to pterosaurs

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are broken however, and not smooth but jagged and rough. This suggests that either

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lack of disturbance to the material and the narrow spaces between the ribs, in addition

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(Fastacht, 2005), even in elements with large diameters such as that preserved here. Indeed the bone walls seen here are especially thin having a ratio of radius / wall

than a number of previously measured pterosaurs (Fastnacht, 2005) though comparable with others (Witton, 2008). The nature of the bone (see description above) suggests that it has not undergone damage from erosion or digestion and that the thin walls are original and unmodified. This discovery also marks the first record of a pterosaur from Tugrikin Shireh.

The diversity of Late Cretaceous pterosaurs is limited, with only the derived azhdarchids, nyctosaurs and the unusual istiodactylids being currently known from this time. The latter is represented by a single isolated jaw from Canada (Arbour and Currie, 2010, though see Witton, 2012) and as istiodactylids bear numerous diagnostic

is not considered a likely candidate. A single nyctosaur humerus has been reported from the Late Cretaceous of Brazil (Price, 1953) but these taxa are both rare and exclusively confined to marine sediments (Unwin, 2005). Thus an azhdarchid identity is favoured and at least some azhdarchoids have especially thin walled bones, even compared to some other pterosaurs (Elgin et al., 2009), which tentatively supports this identification. Finally, an azhdarchid pterosaur has been recorded at the nearby Bayshin Tsav locality (Watabe et al., 2009) that is of similar age to (though older than) this specimen, and the azhdarchids are one of the few pterosaurian groups that likely favoured terrestrial environments (Witton and Naish, 2008). While clearly the bone is

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teeth which have never been reported from Tugrikin Shireh or related sediments, this

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thickness of 30 (6 mm / 0.2 mm). This ratio is higher (i.e. the bone wall is thinner)

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not definitively diagnostic, there is strong circumstantial evidence to support the referral of the material to the Azhdarchidae.

and distal ends of pterosaurian long bones (e.g. a condyle, trochanter, tapering or expansion of the shaft etc.), and so the bone would originally have been longer than the measured 75 mm. It would most likely have exceeded 100 mm, and could potentially have been much longer. In the azhdarchids, long, straight, and thin-walled elements with a sub-circular cross-section that are significantly longer than their diameter correspond to a series of major elements including the humerus, the ulna, radius, wing metacarpal, femur, tibia and the first wing phalanx. Many pterosaur long bones, especially wing elements, are considerably longer than their diameter and taper only a little along their length. Based on the near-complete azhdarchid

may more than 20 times longer than the midshaft width (and thus the bone could potentially have been closer to 250 mm in length originally). While exact determination of the bone here is uncertain, using Zhejiangopterus (Cai and Wei, 1994) as a model, a bone of 100 mm in length or 12 mm in diameter would scale to a variety of wingspans. For example, a humerus of 12 mm diameter would scale to around 2 m, whereas using the radius as a model would produce a value closer to 3.5 m. Witton (2008) calculated the mass of a 2.9 m wingspan Zhejiangopterus as over 9 kg, with a very large azhdarchid (>10 m in wingspan) potentially weighing around 250 kg. Thus the minimum likely size of the pterosaur

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Zhejiangopterus (Cai and Wei, 1994) wing phalanges and long bones such as the tibia

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The bone as preserved lacks of any obvious features associated with the proximal

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was 2 m in wingspan, was probably closer to 3 m, and was potentially from an animal considerably larger still.

5.2 Scavenging or active predation?

Turner et al. (2007) calculated a mass of 24 kg for an adult Velociraptor (of femur length 238 mm), and the specimen here is not mature and rather smaller. Based on the supplementary data of Turner et al. (2007) the animal here was around 13 kg. While pterosaurs are light for their size, an active predator such as an azhdarchid (Witton and Naish, 2008) with a wingspan probably greater than the total length of the dromaeosaur and weighing 9 kg or more would be a difficult, and probably even dangerous, target from a young dromaeosaur. Thus, unless the pterosaur was already ill, infirm or injured, it seems unlikely that this would be a case of predation.

dromaeosaur given that it would be a very substantial part of the carnivores mass. Consuming parts of large bones when there would be substantial amounts of meat available on a newly dead animal suggests that this was late-stage carcass consumption (see Hone and Watabe, 2010 and reference therein). The dromaeosaur was consuming bone (perhaps with some trace flesh attached) presumably because there was little else to eat on the carcass, although it may simply have been seeking a source of minerals. Velociraptorines are known to bite on bones leaving scrape marks (Currie and Jacobsen, 1995; Hone et al., 2010) during late-stage carcass consumption of sizeable

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Furthermore, such a large animal would provide a substantial bounty for a

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carcasses, while smaller food items could be consumed whole (OConnor et al., 2011). While the articular ends of the bone are missing and may have been bitten through,

teeth indeed all the tooth rows appear complete and undamaged, despite the propensity of velociraptorines to shed teeth during feeding (Currie and Jacobsen, 1995; Hone et al., 2010). Thus it appears that the bone was swallowed with little or no oral (or other form of) processing by the dromaeosaur. Here the element in question likely had little in the way of muscle tissue attached to it (since the major muscle groups would attach to the missing articular ends), and the normal delicate feeding of dromaeosaurs suggest that ingestion of large bone elements was not part of their normal feeding routine (e.g. see Hone et al., 2010) and nor is this a common pattern of carcass consumption large bones are consumed whole when there is no alternative.

consume.

5.3 Paravian carnivory Evidence for both predation and scavenging in seen for dromaeosaurs (Norell and Makovicky, 2004; Hone et al., 2010; OConnor et al., 2011) and troodontids (Makovicky and Norell, 2004), and at least some early birds such as Archaeopteryx (Elanowski, 2002) are considered carnivorous / insectivorous (although see also Zanno and Makovicky, 2011). However, it is notable that a similar fossil to that described here is known from the Late Cretaceous of Canada. Currie and Jacobsen

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A bone of such diameter and length would presumably have been a challenge to

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the preserved part shows no feeding traces. Nor does the skull show damage to any

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(1995) described a large azhdarchid tibia that shows both bite marks and an embedded tooth from the dromaeosaur Sauronitholestes. As with the specimen here, it seems

Therefore it is reasonable to infer that at least some carnivorous paravians scavenged large carcasses regularly and it may have been a common behaviour. At the very least, based on the increasing amount of evidence for scavenging in members of this group, they should not be characterised as hyper-specialised predators as they have been on occasion (e.g. Ostrom, 1990). In addition, the rarity of pterosaur fossils in general, and specifically those showing evidence of having been killed or scavenged means that evidence of two separate incidents involving dromaeosaurs and pterosaurs may be more than a coincidence and may speak of potentially close ecological ties in come communities.

bones while feeding, clearly at least some also ingested large bones (or significant parts of them) whole. While this practice is more commonly associated with large theropods (see Hone and Rauhut, 2010 and references therein), small paravians at least had the capacity to swallow relatively large food items whole. In this regard, the gut contents of small theropods such as Compsoganthus (Ostrom, 1978) and Microraptor (Larsson et al., 2010; OConnor et al., 2011) may represent the normal condition for ingestion of food by all non-avian theropods: large parts of, or the whole prey item, would be consumed without extensive oral processing before swallowing. This pattern is also seen in both modern crocodilians and raptorial birds such as owls

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Notably, while some paravians are known to leave extensive bite marks on large

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unlikely that such a small carnivore brought down such a relatively large prey item.

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suggesting a common pattern throughout Archosauria. Nevertheless, presuming that animals did not normally ingest relatively large

not easily digest, this suggests that even small carnivores scavenged regularly. As such, they may have represented an important part of ecosystem recycling in clearing up large carcasses. If the recent model of Carbone et al. (2011) is correct, then numerous small paravians in a given Late Cretaceous terrestrial community might have reached such carcasses rapidly, and were clearly potentially capable of consuming not just soft tissues, but significant parts of the skeleton as well.

Acknowledgements

We thank Fabio Dalla-Vecchia for a photograph of the skull of the Velociraptor

Witton for detailed and helpful comments on an earlier version of the manuscript. Mr. Ken Hayashibara financially supported the fieldwork in Mongolia. The specimen was prepared by a Mongolian preparator, Lkhagvasuren, with his usual consummate skill. CT scanning images of the studied skeleton (supplementary data) were taken in Miyamoto Orthopedic Hospical, Okayama, Japan. The authors thank the division of radiology of the hospital, especially Mr. Fumihiko Kakuta, for their dedication and cooperation.

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Figure Captions:

The broken rib is marked with a white arrow. The two parts of the preserved bone are marked with black arrows (the main part to the left and a smaller broken piece to the right). Scale bar is 50 mm.

Figure. 2. Line drawing of chest cavity of Velociraptor, MPC-D100/54 in right ventrolateral view. The preserved pterosaur bone is coloured grey. Anatomcail abbreviations are as follows: co, coracoid; g, gastralia; r, rib; sc, scapula; st, sternal plate; v, dorsal vertebra. Scale bar is 50 mm.

cavity in posterioventral view showing the cross section shape and the extreme thinness of the cortex. Scale bar is 10 mm.

Figure. 4. Close-up of the bone in the chest cavity in ventral view showing the posteriorly positioned part (to the left) and the associated fragments in line with this (to the right). Scale bar is 10 mm.

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Figure. 3. Close-up of the posteriorly positioned part of the bone in the chest

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Figure. 1. Chest cavity of Velociraptor, MPC-D100/54 in right ventrolateral view.

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22 Highlights - The first theropod dinosaur with a pterosaur bone preserved as gut contents. - This supports previous interpretations of Velociraptor as scavenging. - Pterosaurs were perhaps regularly part of the diet of carnivorous dinosaurs.

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