Maximum Bite Force and Prey Size of Tyrannosaurus rex and Their Relationships to the Inference of Feeding Behavior

The feeding behavior of the theropod dinosaur Tyrannosaurus rex is investigated through analysis of two variables that are critical to successful predation, bite force and prey body mass, as they scale with the size of the predator. These size-related variables have important deterministic effects on the predator’s feeding strategy, through their effects on lethal capacity and choice of prey. Bite force data compiled for extant predators (crocodylians, carnivorans, chelonians and squamates) are used to establish a relationship between bite force and body mass among extant predators. These data are used to estimate the maximum potential bite force of T. rex, which is between about 183,000 and 235,000 N for a bilateral bite. The relationship between maximum prey body mass and predator body mass among the same living vertebrates is used to infer the likely maximum size of prey taken by T. rex in the Late Cretaceous. This makes it possible to arrive at a more rigorous assessment of the role of T. rex as an active predator and/or scavenger than has hitherto been possible. The results of this analysis show that adult Triceratops horridus fall well within the size range of potential prey that are predicted to be available to a solitary, predaceous T. rex. This analysis establishes boundary conditions for possible predator/prey relationships among other dinosaurs, as well as between these two taxa.     

Los Cayos Dinosaur Tracksite: An Overview on the Lower Cretaceous Ichno-Diversity of the Cameros Basin (Cornago,La Rioja Province,Spain)

The Los Cayos dinosaur tracksite is located at the eastern sector of the Cameros Basin (Cornago Township, La Rioja province), NE Spain. The sediments consist of interbedded terrigenous siliciclastics and carbonates belonging to the Enciso Group (Early Cretaceous, Aptian in age). The sedimentological and faunal evidence suggests that these sediments were deposited in a low gradient lacustrine environment. Los Cayos constitutes a relatively wide area with at least 6 localities that have yielded dinosaur tracks. More than 2,000 dinosaur tracks have been discovered to date. Medium to large theropod dinosaurs constitute about 95% of the ichnofauna. Ornithopod dinosaur tracks have been reported from only one of the outcrops (Los Cayos D). Los Cayos S has yielded theropod and sauropod tracks. One sauropod trackway shows a narrow-gauge locomotion pattern and a manus print morphology suggesting that the trackmaker was a titanosaurid, or at least a titanosauriform sauropod. Some pterosaur manus impressions, avian-like footprints of small size and possible turtle tracks complete the assemblage of one of the most impressive and best-preserved dinosaur tracksites of the European Lower Cretaceous.     

Gar-Bitten Coprolite From South Carolina,USA

Coprolites can preserve a wide range of biogenic components. Of all the coprolites known from the fossil record, hitherto only two are known to preserve vertebrate tooth impressions (i.e., those of chondrichthyans). Here, a coprolite, from a thick lag deposit that includes a mixture of late Cretaceous, early Paleocene, and Plio-Pleistocene taxa at Clapp Creek in Kingstree, Williamsburg County, South Carolina, USA, preserves bite marks most consistent with having been made by a gar, Lepisosteus sp. (Lepisosteidae, Actinopterygii). This is the first-known coprolite to preserve actinopterygian tooth/bite marks. Aborted coprophagy seems unlikely; an accidental or serendipitous strike more likely describes the origin of the score marks over the surface of the coprolite. This coprolite also preserves small paired striations interpreted as evidence of coprophagy by an unknown organism.     

Opportunistic exploitation of dinosaur dung: fossil snails in coprolites from the Upper Cretaceous Two Medicine Formation of Montana

Multiple associations of fossil snails with dinosaur coprolites demonstrate that snails and dinosaurs not only shared ancient habitats but were trophically linked via dinosaur dung. Over 130 fossil snails representing at least seven different taxa have been found on or within herbivorous dinosaur coprolites from the Upper Cretaceous Two Medicine Formation of Montana. The terrestrial snail Megomphix is the most common taxon, but three other terrestrial taxa (Prograngerella, Hendersonia and Polygyrella) and three aquatic snails (Lioplacodes, ?Viviparus and a physid) also occur in coprolites. At least 46% of the shells in the faeces are whole or nearly so, indicating that most (if not all) of the snails were not ingested by dinosaurs, but were post‐depositional visitors to the dung pats. The sizeable, moist and microbially enriched dinosaur faeces would have provided both food and roosting sites for the ancient snails, and the large number of snail–coprolite associations reflect recurring, opportunistic exploitation of dung. The terrestrial taxa in the coprolites suggest that this Late Cretaceous locality included sufficiently moist detrital or vegetative cover for snails when dinosaur dung was not present. The aquatic snails probably entered the faeces during flood events. Dinosaur dung would have provided an abundant but patchy influx of resources that was probably seasonally available in the ancient environment.     

Nesting Habitat Creation Enhances Recruitment in a Predator‐Free Environment: Malaclemys Nesting at the Paul S. Sarbanes Ecosystem Restoration Project

Aquatic turtles worldwide are plagued with habitat loss due to development and shoreline alteration that destroys the terrestrial–aquatic linkage which they must cross to reproduce successfully. Furthermore, nesting habitat loss can concentrate nesting, increasing nest predator efficiency. We describe how the Paul S. Sarbanes Ecosystem Restoration Project at Poplar Island created nesting habitat for Malaclemys terrapin (Diamondback Terrapin), and document nesting success in response to construction progress and the absence of raccoons and foxes, the primary nest predators. We monitored terrapin nests throughout the nesting seasons from 2002 to 2011 to determine overall and within‐nest survivorship. Female terrapins began nesting on the restoration project within 1 year but planned construction during the study eliminated some nesting areas and opened previously inaccessible areas. Overall, nest survivorship was considerably higher than mainland nesting areas due to the absence of raccoons and foxes on the island and within‐nest survivorship was similar. Egg size, hatchling size, and the frequency of shell scute anomalies were similar to other terrapin populations, suggesting normal developmental conditions on the island. We documented annual variation in hatchling size that correlated negatively with mean air temperature during the incubation season. Our results indicate that restored or created isolated island habitat can be located rapidly by terrapins and can become an important source of recruitment in regions where nesting habitat is limited and predation is high. Poplar Island illustrates how habitat loss and restoration can affect turtle populations by revealing the changes in nesting patterns and success in newly created, predator‐free habitat.     

The first dinosaur egg from the Lower Cretaceous of Western Siberia,Russia

The Lower Cretaceous Ilek Formation in Western Siberia (Russia) has yielded various vertebrate fossils, including skeletal remains of dinosaurs. Here we report on a fragmentary theropod egg from the vertebrate locality Shestakovo 3 of the Ilek Formation in Kemerovo Province. We assign the specimen to the oogenus Prismatoolithus (oofamily Prismatoolithidae) as Prismatoolithus ilekensis oosp. nov., on the basis of the following unique combination of characters: ovoid-shaped egg; thin eggshell 300–330 μm thick; angustiprismatic morphotype; eggshell with three different layers; gradual transition between mammillary layer and prismatic layer; abrupt contact between prismatic layer and external layer; mammillary layer to prismatic layer to external layer thickness ratio is 1:3:0.6; prismatic layer with ill-defined squamatic texture; angusticanaliculate pore system; and smooth outer surface. Like other Early Creataceous Prismatoolithus, the egg of Prismatoolithus ilekensis oosp. nov. was laid by a small bodied theropod dinosaur (troodontid or primitive bird) and this taxonomic attribution is supported by results of our phylogenetic analysis. Prismatoolithus ilekensis oosp. nov. is the first Early Cretaceous ootaxon from Russia.

urn:lsid:zoobank.org:act:734EAD40-86C3-488B-A61E-B5FF7378BC0E     

A diversified vertebrate ichnite fauna from the Dasheng Group (Lower Cretaceous) of southeast Shandong Province,China

The Qingquan dinosaur tracksite, from the Lower Cretaceous Dasheng Group, Shandong Province, China adds to the growing record of saurischian-dominated ichnofaunas of the region. The site reveals the presence of avian theropods (Koreanaornis) and non-avian theropods tentatively referred to Jialingpus. Sauropod tracks are referred to Brontopodus. One site shows evidence of extensive trampling attributable to sauropods and theropods that moved in the same westerly direction, though not necessarily at exactly the same time. This site is reminiscent of the famous Davenport Ranch site in Texas which has provoked much debate about the herding behavior of sauropods as a defense strategy against theropod predators.     

The First Dinosaur from Washington State and a Review of Pacific Coast Dinosaurs from North America

We describe the first diagnostic dinosaur fossil from Washington State. The specimen, which consists of a proximal left femur, was recovered from the shallow marine rocks of the Upper Cretaceous (Campanian) Cedar District Formation (Nanaimo Group) and is interpreted as pertaining to a large theropod on the basis of its hollow medullary cavity and proximally placed fourth trochanter. The Washington theropod represents one of the northernmost occurrences of a Mesozoic dinosaur on the west coast of the United States and one of only a handful from the Pacific coast of Laramidia during the Cretaceous. Its isolated nature and preservation in marine rocks suggest that the element was washed in from a nearby fluvial system. If the femur pertains to a tyrannosauroid, which seems likely given its size and the widespread occurrence of the group across Laramidia during Late Cretaceous times, then it would represent an earlier occurrence of large body size than previously recognized (complete femur length estimated at 1.2 meters). Uncertainty surrounding the latitude of deposition of the Nanaimo Group (i.e., the Baja-British Columbia hypothesis) precludes assigning the Washington theropod to either of the putative northern or southern biogeographic provinces of Laramidia.     

Dinosaur footprints from the Sao Khua Formation (Early Cretaceous) of Phu Kao National Park,Northeast Thailand

The first documented dinosaur footprints from Southeast Asia were theropod footprints found in Thailand at Phu Luang Wildlife Sanctuary. Since then, Mesozoic vertebrate footprints have been discovered in twelve different localities in Thailand. In 2000, dinosaur footprints were found in Phu Kao National Park in Northeastern Thailand, however, they have never been studied in detail. The following study presents the first map, the complete measurements, the geology, and stratigraphy of the site and surrounding area. The study discusses the nature of the trackmaker and the digitigrade character of the footprints. This discovery represents the first documented occurrence of dinosaur footprints inside the Early Cretaceous Sao Khua Formation from Thailand.     

Saurischian (theropod–sauropod) track assemblages from the Jiaguan Formation in the Sichuan Basin,Southwest China: ichnology and indications to differential track preservation

Saurischian (theropod and sauropod) tracks and trackways from the Jiaguan Formation (Lower Cretaceous) of the Sichuan Basin are exposed as natural casts with associated undertrack or transmitted print casts. The theropod tracks (cf. Eubrontes and Grallator) were left by differently sized trackmakers. This is a further example for the occurrence of characteristic Lower Jurassic ichnotaxa in the Cretaceous that obviously had a more extended stratigraphic range in East Asia. The sauropod trackway is tentatively assigned to cf. Brontopodus based on imprint morphology and (nearly wide) gauge. The tracks, however, allow a detailed study of their formation and the taphonomic processes under different substrate conditions. Differential preservation and erosion of primary sedimentary structures, and post-burial deformation structures, give insight into a complex preservational history during a low energy phase interrupting the deposition of a sequence of thick high energy sandstones. This is the sixth report of dinosaur tracks from the Jiaguan Formation and the fifteenth report from the Lower Cretaceous of Sichuan Province. Thus, the tetrapod ichnological record in this region is rapidly becoming of major importance for our knowledge of dinosaur faunas in south-western China.     

Phenotypically plastic responses of green frog embryos to conflicting predation risk

Predators have been shown to alter the timing of switch points between life history stages, but few studies have addressed switch point plasticity in prey exposed simultaneously to conflicting predation pressure. We tested hatching responses of green frog (Rana clamitans) embryos subject to perceived predation risk from chemical cues released by two stage-specific predators, predicting that these predators would elicit: (1) directional hatching responses when presented independently, and (2) intermediate phenotypic responses when presented simultaneously. R. clamitans embryos in outdoor exclosures were exposed to cues from an egg predator (freshwater leeches; Nephelopsis obscura), a larval predator (dragonfly nymphs, Aeschna canadensis), and both predators in a 2 × 2 factorial experiment, and changes in hatchling size, hatchling developmental stage, and hatching time were compared to those for control embryos. Leeches alone induced embryos to hatch at a smaller size and an earlier developmental stage than controls, while dragonfly nymphs elicited a delay in egg hatching time that was associated with larger size and later developmental stage at hatching. Embryos failed to respond to simultaneous exposure to both predators, implying that responses to each occurred concurrently and were therefore dampened. Our results indicate that prey under threat from conflicting predators may manifest intermediate defensive phenotypes. Such intermediate responses may result in elevated rates of prey mortality with possible consequences at the population level.     

New Data on Therangospodus oncalensis from the Berriasian Fuentesalvo Tracksite (Villar del Río,Soria, Spain): An Example of Gregarious Behaviour in Theropod Dinosaurs

Therangospodus oncalensis is a Berriasian theropod ichnotaxon from the Cameros Basin (Soria Province, Spain). We discovered new trackways assigned to this ichnotaxon during cleaning and conservation works at the type locality (Fuentesalvo), enabling us to describe it more precisely. Therangospodus oncalensis was made by a medium-sized theropod dinosaur and is characterized by a single tapering pad on each toe, a rounded heel impression and a narrow trackway. In addition to having similar morphometric characteristics, most of the trackways at the Fuentesalvo site show comparable size and biometric features. This indicates that, in terms of size and age, a homogeneous population of a single theropod species could have produced these tracks. Moreover, the similar orientation (lying within a range of only 15°) and estimated speed suggested by most of the trackways, lend weight to the hypothesis that the trackmaker of Therangospodus oncalensis was able, at least at times, to move in structured packs displaying gregarious behaviour. Furthermore, this is the first structured pack of theropods to be described.     

Vertebrate Tracksites in the Middle Jurassic-Upper Cretaceous of South Tunisia

Four vertebrate tracksites from the Middle Jurassic and Upper Cretaceous in the Tataouine basin of southern Tunisia are described. Approximately 130 tridactyl footprints distributed over an area of 200 square meters, preserved on Callovian beds exposed at the Beni Ghedir site, represent the oldest evidence of a dinosaur fauna in Tunisia. In addition, three tracksites—Chenini, Ksar Ayaat, and Jebel Boulouha—have been discovered in the Cretaceous beds of the upper Continental Intercalaire, previously considered as a strictly marine depositional sequence. In addition to dinosaur tracks, the Chenini tracksite (late Albian) includes poorly preserved crocodilian tracks, and footprints assigned to a pleurodiran turtle have been recovered at the Ksar Ayaat locality (early Cenomanian). The Jebel Boulouha tracksite is dominated by well-preserved tridactyl tracks referred to small-sized theropods. Depositional settings of each tracksite have been defined on stratigraphic and sedimentologic data, and tracks were ascribed to different ichnocoenoses in relation to their paleoenvironments. This new and differentiated track record gives important information on how the fossil vertebrate fauna changed in southern Tunisia during mid-Jurassic to mid-Cretaceous times. These data provide a unique and useful census of tetrapod associations along the southern margin of the peri-Mediterranean area.     

山东莱阳晚白垩世恐龙与恐龙蛋研究历史和新进展

简要回顾了莱阳恐龙和恐龙蛋化石群的研究历史和以杨钟健为代表的老一代地质古生物学者对莱阳恐龙和恐龙蛋研究发现的杰出贡献,并介绍了莱阳恐龙和恐龙蛋的最新发现和若干研究进展.在近年来对莱阳周边地层进行的大规模考察中,发现了一系列发育在平原上的恐龙峡谷群,以及其中蕴含的十几个恐龙和恐龙蛋新地点和新层位.2010年开始,在对莱阳金岗口村附近的2个化石地点的发掘中,发现了以鸭嘴龙科为主的大量脊椎动物化石和蛋化石,包括一类新的栉龙亚科成员,一些大型兽脚类恐龙化石,以及一新的龟鳖类蛋化石等.2号地点化石富集层具有典型的泥石流沉积特征和骨骼埋藏特征.对棘鼻青岛龙的特殊头饰进行了CT扫描和三维重建,发现其头饰是实心结构,但其他骨骼特征证明棘鼻青岛龙属于具有头饰的赖氏龙亚科是确定无疑的,所以目前发现的头饰应不是其真实状态或根本不属于其头部骨骼.此外,对谭氏龙1属3种的重新观察研究得出以下结论:中国谭氏龙和金刚口谭氏龙应是有效属种,但部分骨骼还有疑问,还需要进一步研究,而莱阳谭氏龙为无效属种.     

Dinosaur Tracks from the Cedar Mountain Formation (Lower Cretaceous), Arches National Park,Utah

The seventh and largest known dinosaur tracksite from the Cedar Mountain Formation is reported from two important stratigraphic levels in the Ruby Ranch Member within the boundaries of Arches National Park. Previous reports of sites with a few isolated tracks are of limited utility in indicating the fauna represented by track makers. The Arches site reveals evidence of several theropod morphotypes, including a possible match for the coelurosaur Nedcolbertia and an apparently didactyl Utahraptor-like dromeosaurid. Sauropod tracks indicate a wide-gauge morphotype (cf. Brontopodus). Ornithischian tracks suggest the presence of an iguandontid-like ornithopod and a large ankylosaur. Dinosaur track diversity is high in comparison with other early Cretaceous vertebrate ichnofaunas, and it correlates well with faunal lists derived from skeletal remains, thus providing a convincing census of the known fauna.     

Shape variability in tridactyl dinosaur footprints: the significance of size and function

The functional anatomy of the hindlimb of bipedal dinosaurs has been intensively studied. Yet, surprisingly little work has been done concerning functional adaptation of digits for terrestrial locomotion. While complete and articulated pes skeletons are scarce, pes shape is abundantly recorded by fossil footprints. We elucidate the significance of footprint shape and size for locomotion using a large sample (n = 303) of tridactyl dinosaur footprints from a broad range of geographical localities and time slots. Size and shape variation are characterized separately for theropods and ornithischians, the two principal trackmaker taxa. At smaller sizes, theropod footprints are best discriminated from ornithischian footprints by their smaller interdigital angle and larger projection of digit III; at larger sizes digital widths are effective discriminants. Ornithischian footprints increase in size from the Early Jurassic to the Late Cretaceous, a trend not observed in theropod footprints. Size and function are argued to be important determinants of footprint shape, and an attempt made to infer function from shape. Digit III projection and length-to-width ratio of the footprints are negatively correlated with size in both groups; digit impression width is positively correlated with size only in ornithischians. Digit III projection appears to be positively correlated with cursorial ability. Increased interdigital angles are associated with a decrease in digital width, possibly an adaptation for stability. Weak digit III projection and increased digital width are interpreted as adaptations for graviportality. Footprints yield great potential for the understanding of the functional morphology of dinosaur feet.     

Vertebrate Ichnopathology: Pathologies Inferred from Dinosaur Tracks and Trackways from the Mesozoic

Literature concerning dinosaur footprints or trackways exhibiting abnormal gait or morphology reflecting pathology (ichnopathology) is rare. We report on a number of Jurassic and Cretaceous occurrences of theropod footprints from western North America with unusual morphologies interpreted herein as examples of inferred pathologies, or ichnopathologies. The majority of ichnopathologies are primarily manifested in the digit impressions and include examples of swelling, extreme curvature, dislocation or fracture, and amputation. A number of occurrences are single tracks on ex situ blocks with substantial deformation (inferred dislocation or fracture), or absence of a single digit impression. Two occurrences are from in situ natural mould trackways, one of which is a lengthy trackway of a presumed allosauroid with no noticeable deformation of the digits or feet but with strong inward rotation of the left footprint toward the midline and a pronounced, waddling limp. The other is a tyrannosaurid trackway consisting of three footprints (one right, two left) with the two left prints exhibiting repetitive ichnopathology of a partially missing Digit II impression.     

A new peculiar dinosaur egg,Sankofa pyrenaica oogen. nov. oosp. nov. from the Upper Cretaceous coastal deposits of the Aren Formation,south‐central Pyrenees,Lleida, Catalonia,Spain

Abstract: A new type of small, ovoid dinosaur egg, Sankofa pyrenaica oogen. nov. oosp. nov., with a prismatic type eggshell is described from upper Cretaceous (upper Campanian–Maastrichtian) deposits of the Montsec area, South Pyrenean Central Unit, Lleida, Catalonia, Spain. This egg type was sub‐vertically laid in only two rich monospecific sites of a single stratigraphic layer from coastal deposits of the Aren Formation, interpreted as an emerged beach ridge of a barrier island – lagoon depositional system. The size and shape of these eggs with their asymmetric poles are roughly similar to modern hen eggs, which is unusual in the Cretaceous fossil egg record. Its phylogenetic position clusters with bird and Troodontid eggs. A morphospace analysis of egg shapes shows the similarity of the new egg to a Campanian fossil bird egg from Argentina, both being intermediate between modern‐bird eggs and extinct nonavian theropod eggs. However, the eggshell microstructure of Sankofa pyrenaica differs from that of bird eggs in its incipient squamatic texture. It has a peculiar pattern of interlocking small crystals in the middle of the palisade layer, instead of the thick squamatic structure commonly present in modern avian eggshells. This new egg type is attributed to a small theropod, probably with a single oviduct like birds and whose mosaic distribution of features is a combination between that of birds and nonavian theropods. This enhances the arguments supporting the close phylogenetic relationships between both groups.     

Coprolite morphotypes from the Upper Cretaceous of Sweden: novel views on an ancient ecosystem and implications for coprolite taphonomy

Eriksson, M.E., Lindgren, J., Chin, K. & Månsby, U. 2011: Coprolite morphotypes from the Upper Cretaceous of Sweden: novel views on an ancient ecosystem and implications for coprolite taphonomy. Lethaia, Vol. 44, pp. 455–468. Coprolites (fossilized faeces) are common, yet previously unreported, elements in the Campanian (Upper Cretaceous) shallow‐marine strata of Åsen, southern Sweden. They are associated with a diverse vertebrate fauna and comprise at least seven different morphotypes that suggest a variety of source animals. Their faecal origin is corroborated by several lines of evidence, including chemical composition (primarily calcium phosphate), external morphology and nature of the inclusions. Preservation in a fossil coquina, interpreted as a taphocoenosis, suggests early lithification promoted by rapid entombment. This would have prevented disintegration of the faecal matter and facilitated transportation and introduction to the host sediment. The coprofabrics can generally be correlated to specific gross morphologies, supporting a morphology‐determined coprolite classification. Moreover, having been deposited under presumably comparable taphonomic conditions, variations in coprofabrics infer differences in diet and/or digestive efficiency of the host animal. Size and morphology of the coprolites imply that most, if not all, were produced by vertebrates and the largest specimens infer a host animal of considerable size. Two spiralled coprolite morphotypes yield bone fragments and scales of bony fish, suggesting that the producers were piscivorous sharks. Other coprolites contain inclusions interpreted as the remains of shelled invertebrates, thus indicating that they may have derived from durophagous predators and/or scavengers. The occurrence of small scrapes, tracks and traces on several specimens suggest manipulation of the faeces by other (presumably coprophagous) organisms after deposition. The collective data from the Åsen coprolites provide new insights into a shallow‐water Late Cretaceous marine ecosystem hitherto known solely from body fossils. □ Coprolites, vertebrates, coprofabrics, taphonomy, trophic levels, Upper Cretaceous, Sweden.     

Dinosaur nest ecology and predation during the Late Cretaceous: was there a relationship between upper Cretaceous extinction and nesting behavior?

Many hypotheses have been advanced to explain the K/Pg extinctions, yet none closely examines the likely interactions between dinosaurs and contemporary taxa within their communities. The diversity of predators of dinosaur nests and hatchlings increased toward the end of the Cretaceous. In addition to large snakes having been found fossilized in the act of foraging in dinosaur nests, mammals and birds had also evolved new forms potentially capable of exploiting this resource. The constraints on mammal size and niche diversity lessened prior to the K/Pg boundary. Using comparisons of predator/prey size ratios between extant species and known fossils, we demonstrate that mammalian and avian clades had members large enough to prey on dinosaur eggs and hatchlings. We argue that the reproductive strategy of obligatory nest defense was likely practiced by most non-avian dinosaur species. This strategy was highly susceptible to the increasing numbers of mammalian, avian, and reptilian predators, which rendered this strategy obsolete. Continued selection against large oviparous species in the Cenozoic has limited this life-history strategy to habitats that provide concealment – primarily grasslands, a habitat that did not exist until the Miocene. We urge the evaluation of multiple, perhaps synergistic, hypotheses when considering extinction events of this magnitude.