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Based on morphology alone, what type of claw does the Tyrannosaur have?

Based on morphology alone, what type of claw does the Tyrannosaur have?


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I understand that the exact use of Tyrannosaur Rex's claw is a mystery, or at least debated. I also understand that claws can be used for a variety of different purposes, selective pressure adapts their morphological to be better optimized for certain tasks. Some are used for hunting and killing, like large cats' claws, some are used for digging, like bear's claws, while others are used for climbing, like squirrel's claws.

My question is, all other clues and bits of information about if and how T. Rex might have used its arms, what 'type' of claws did it possess? I'm not asking, in this question, about its arm, the arm's musculature, its mouth or jaw, or about any of the rest of the animal. Based on the morphology of the claw alone, and comparing it to known uses of similarly shaped claws, what types of tasks were the claws geared towards?


unfortunately they have fairly generic claws, they are not specialized enough to point to a use. They are curved enough to be used to grasp something but that is true for their ancestral line too so there is no sign of a specific directional selection.


Fungus Identification

Monika Novak Babič , . Nina Gunde-Cimerman , in Reference Module in Life Sciences , 2020

Correct and Uniform Identification of Fungi

Another problem is fungal identification to the species level. Microscopic observations are rarely correct ( De Hoog et al., 2019 ). Even after visible growth on a common cultivation media, not all fungi form distinctive macro- and micro-morphological features. Some genera may thus not be identified correctly in laboratories equipped with only the basic equipment ( Novak Babič et al., 2017 ). The molecular approach is a good alternative to microscope identification, but sometimes even the internal transcribed spacer (ITS) used as a general fungal barcode fails to identify fungi to the genus or species level, prolonging identification time and increasing costs ( Irinyi et al., 2015 ). Lastly, correct and uniform identification of fungi may represent an issue for laboratories lacking people trained in mycology, since fungal taxonomy is currently facing a huge restructuring, known under the term “one fungus – one name”, frequently resulting in old names being recently replaced or combined based on recognition of both sexual and asexual forms ( Taylor, 2011 ).


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Vol 329, Issue 5998
17 September 2010

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By Stephen L. Brusatte , Mark A. Norell , Thomas D. Carr , Gregory M. Erickson , John R. Hutchinson , Amy M. Balanoff , Gabe S. Bever , Jonah N. Choiniere , Peter J. Makovicky , Xing Xu

Science 17 Sep 2010 : 1481-1485


Contents

The first remains of tyrannosaurids were uncovered during expeditions led by the Geological Survey of Canada, which located numerous scattered teeth. These distinctive dinosaur teeth were given the name Deinodon ("terrible tooth") by Joseph Leidy in 1856. The first good specimens of a tyrannosaurid were found in the Horseshoe Canyon Formation of Alberta, and consisted of nearly complete skulls with partial skeletons. These remains were first studied by Edward Drinker Cope in 1876, who considered them a species of the eastern tyrannosauroid Dryptosaurus. In 1905, Henry Fairfield Osborn recognized that the Alberta remains differed considerably from Dryptosaurus, and coined a new name for them: Albertosaurus sarcophagus ("flesh-eating Alberta lizard"). [5] Cope described more tyrannosaur material in 1892, in the form of isolated vertebrae, and gave this animal the name Manospondylus gigas. This discovery was mostly overlooked for over a century, and caused controversy in the early 2000s when it was discovered that this material actually belonged to, and had name priority over, Tyrannosaurus rex. [6]

In his 1905 paper naming Albertosaurus, Osborn described two additional tyrannosaur specimens that had been collected from in Montana and Wyoming during a 1902 expedition of the American Museum of Natural History, led by Barnum Brown. Initially, Osborn considered these to be distinct species. The first, he named Dynamosaurus imperiosus ("emperor power lizard"), and the second, Tyrannosaurus rex ("king tyrant lizard"). A year later, Osborn recognized that these two specimens actually came from the same species. Despite the fact that Dynamosaurus had been found first, the name Tyrannosaurus had appeared one page earlier in his original article describing both specimens. Therefore, according to the International Code of Zoological Nomenclature (ICZN), the name Tyrannosaurus was used. [7]

Barnum Brown went on to collect several more tyrannosaurid specimens from Alberta, including the first to preserve the shortened, two-fingered forelimbs characteristic of the group (which Lawrence Lambe named Gorgosaurus libratus, "balanced fierce lizard", in 1914). A second significant find attributed to Gorgosaurus was made in 1942, in the form of a well-preserved, though unusually small, complete skull. The specimen waited until after the end of World War II to be studied by Charles W. Gilmore, who named it Gorgosaurus lancesnis. [5] This skull was re-studied by Robert T. Bakker, Phil Currie, and Michael Williams in 1988, and assigned to the new genus Nanotyrannus. [8] It was also in 1946 that paleontologists from the Soviet Union began expeditions into Mongolia, and uncovered the first tyrannosaur remains from Asia. Evgeny Maleev described new Mongolian species of Tyrannosaurus and Gorgosaurus in 1955, and one new genus: Tarbosaurus ("terrifying lizard"). Subsequent studies, however, showed that all of Maleev's tyrannosaur species were actually one species of Tarbosaurus at different stages of growth. A second species of Mongolian tyrannosaurid was found later, described by Sergei Kurzanov in 1976, and given the name Alioramus remotus ("remote different branch"), though its status as a true tyrannosaurid and not a more primitive tyrannosaur is still controversial. [9] [5]

The tyrannosaurids were all large animals, with all species capable of weighing at least 1 metric ton. [10] A single specimen of Alioramus of an individual estimated at between 5 and 6 metres (16 and 20 ft) long has been discovered, [9] although it is considered by some experts to be a juvenile. [10] [11] Albertosaurus, Gorgosaurus and Daspletosaurus all measured between 8 and 10 metres (26 and 33 ft) long, [12] while Tarbosaurus reached lengths of 12 metres (39 ft) from snout to tail. [13] The massive Tyrannosaurus reached 12.3 metres (40 ft) in one of the largest specimens, FMNH PR2081. [2]

Tyrannosaurid skull anatomy is well understood as complete skulls are known for all genera but Alioramus, which is known only from partial skull remains. [14] Tyrannosaurus, Tarbosaurus, and Daspletosaurus had skulls that exceeded 1 m (3.3 ft) in length. [12] Adult tyrannosaurids had tall, massive skulls, with many bones fused and reinforced for strength. At the same time, hollow chambers within many skull bones and large openings (fenestrae) between those bones helped to reduce skull weight. Many features of tyrannosaurid skulls were also found in their immediate ancestors, including tall premaxillae and fused nasal bones. [10]

Tyrannosaurid skulls had many unique characteristics, including fused parietal bones with a prominent sagittal crest, which ran longitudinally along the sagittal suture and separated the two supratemporal fenestrae on the skull roof. Behind these fenestrae, tyrannosaurids had a characteristically tall nuchal crest, which also arose from the parietals but ran along a transverse plane rather than longitudinally. The nuchal crest was especially well-developed in Tyrannosaurus, Tarbosaurus and Alioramus. Albertosaurus, Daspletosaurus and Gorgosaurus had tall crests in front of the eyes on the lacrimal bones, while Tarbosaurus and Tyrannosaurus had extremely thickened postorbital bones forming crescent-shaped crests behind the eyes. Alioramus had a row of six bony crests on top of its snout, arising from the nasal bones lower crests have been reported on some specimens of Daspletosaurus and Tarbosaurus, as well as the more basal tyrannosauroid Appalachiosaurus. [11] [15]

The snout and other parts of the skull also sported numerous foramina. According to the 2017 study which described D. horneri, scaly integument as well as tactile sensitivity was correlated with the multiple rows of neurovascular foramina seen in crocodylians and tyrannosaurids. [16]

The skull was perched at the end of a thick, S-shaped neck, and a long, heavy tail acted as a counterweight to balance out the head and torso, with the center of mass over the hips. Tyrannosaurids are known for their proportionately very small two-fingered forelimbs, although remnants of a vestigial third digit are sometimes found. [10] [17] Tarbosaurus had the shortest forelimbs compared to its body size, while Daspletosaurus had the longest.

Tyrannosaurids walked exclusively on their hindlimbs, so their leg bones were massive. In contrast to the forelimbs, the hindlimbs were longer compared to body size than almost any other theropods. Juveniles and even some smaller adults, like more basal tyrannosauroids, had longer tibiae than femora, a characteristic of fast-running dinosaurs like ornithomimids. Larger adults had leg proportions characteristic of slower-moving animals, but not to the extent seen in other large theropods like abelisaurids or carnosaurs. The third metatarsals of tyrannosaurids were pinched between the second and fourth metatarsals, forming a structure known as the arctometatarsus. [10]

It is unclear when the arctometatarsus first evolved it was not present in the earliest tyrannosauroids like Dilong, [18] but was found in the later Appalachiosaurus. [15] This structure also characterized troodontids, ornithomimids and caenagnathids, [19] but its absence in the earliest tyrannosauroids indicates that it was acquired by convergent evolution. [18]

Teeth Edit

Tyrannosaurids, like their tyrannosauroid ancestors, were heterodont, with premaxillary teeth D-shaped in cross section and smaller than the rest. Unlike earlier tyrannosauroids and most other theropods, the maxillary and mandibular teeth of mature tyrannosaurids are not blade-like but extremely thickened and often circular in cross-section, with some species having reduced serrations. [10] Tooth counts tend to be consistent within species, and larger species tend to have lower tooth counts than smaller ones. For example, Alioramus had 76 to 78 teeth in its jaws, while Tyrannosaurus had between 54 and 60. [20]

William Abler observed in 2001 that Albertosaurus tooth serrations resemble a crack in the tooth ending in a round void called an ampulla. [21] Tyrannosaurid teeth were used as holdfasts for pulling meat off a body, so when a tyrannosaur would have pulled back on a piece of meat, the tension could cause a purely crack-like serration to spread through the tooth. [21] However, the presence of the ampulla would have distributed these forces over a larger surface area, and lessened the risk of damage to the tooth under strain. [21] The presence of incisions ending in voids has parallels in human engineering. Guitar makers use incisions ending in voids to, as Abler describes, "impart alternating regions of flexibility and rigidity" to the wood they work with. [21] The use of a drill to create an "ampulla" of sorts and prevent the propagation of cracks through material is also used to protect airplane surfaces. [21] Abler demonstrated that a plexiglass bar with incisions called "kerfs" and drilled holes was more than 25% stronger than one with only regularly placed incisions. [21] Unlike tyrannosaurs and other theropods, ancient predators like phytosaurs and Dimetrodon had no adaptations to prevent the crack-like serrations of their teeth from spreading when subjected to the forces of feeding. [21]

The name Deinodontidae was coined by Edward Drinker Cope in 1866 for this family, [22] and continued to be used in place of the newer name Tyrannosauridae through the 1960s. [23] The type genus of the Deinodontidae is Deinodon, which was named after isolated teeth from Montana. [24] However, in a 1970 review of North American tyrannosaurs, Dale Russell concluded that Deinodon was not a valid taxon, and used the name Tyrannosauridae in place of Deinodontidae, stating that this was in accordance with ICZN rules. [12] Therefore, Tyrannosauridae is preferred by modern experts. [5]

Tyrannosaurus was named by Henry Fairfield Osborn in 1905, along with the family Tyrannosauridae. [25] The name is derived from the Ancient Greek words τυραννος (tyrannos) ('tyrant') and σαυρος (sauros) ('lizard'). The very common suffix -idae is normally appended to zoological family names and is derived from the Greek suffix -ιδαι -idai, which indicates a plural noun. [26]

Taxonomy Edit

Tyrannosauridae is uncontroversially divided into two subfamilies. Albertosaurinae comprises the North American genera Albertosaurus and Gorgosaurus, while Tyrannosaurinae includes Daspletosaurus, Teratophoneus, Bistahieversor, Tarbosaurus, Nanuqsaurus, Zhuchengtyrannus, and Tyrannosaurus itself. [27] Some authors include the species Gorgosaurus libratus in the genus Albertosaurus and Tarbosaurus bataar in the genus Tyrannosaurus, [15] [5] [28] while others prefer to retain Gorgosaurus and Tarbosaurus as separate genera. [10] [11] Albertosaurines are characterized by more slender builds, lower skulls, and proportionately longer tibiae than tyrannosaurines. [10] In tyrannosaurines, the sagittal crest on the parietals continues forward onto the frontals. [11] In 2014, Lü Junchang et al. described the Alioramini as a tribe within the Tyrannosauridae containing the genera Alioramus and Qianzhousaurus. Their phylogenetic analysis indicated that the tribe was located at the base of the Tyrannosaurinae. [29] [30] Some authors, such as George Olshevsky and Tracy Ford, have created other subdivisions or tribes for various combinations of tyrannosaurids within the subfamilies. [31] [32] However, these have not been phylogenetically defined, and usually consisted of genera that are now considered synonymous with other genera or species. [20]

Additional subfamilies have been named for more fragmentary genera, including Aublysodontinae and Deinodontinae. However, the genera Aublysodon and Deinodon are usually considered nomina dubia, so they and their eponymous subfamilies are usually excluded from taxonomies of tyrannosaurids. An additional tyrannosaurid, Raptorex, was initially described as a more primitive tyrannosauroid, but likely represents a juvenile tyrannosaurine similar to Tarbosaurus. However, as it is known only from a juvenile specimen, it is also currently considered a nomen dubium. [33]

Phylogeny Edit

With the advent of phylogenetic taxonomy in vertebrate paleontology, Tyrannosauridae has been given several explicit definitions. The original was produced by Paul Sereno in 1998, and included all tyrannosauroids closer to Tyrannosaurus than to either Alectrosaurus, Aublysodon or Nanotyrannus. [34] However, Nanotyrannus is often considered to be a juvenile Tyrannosaurus rex, while Aublysodon is usually regarded as a nomen dubium unsuitable for use in the definition of a clade. [10] Definitions since then have been based on more well-established genera.

In 2001, Thomas R. Holtz Jr. published a cladistic analysis of the Tyrannosauridae. [35] He concluded that there were two subfamilies: the more primitive Aublysodontinae, characterized by unserrated premaxillary teeth and the Tyrannosaurinae. [35] The Aublysodontinae included Aublysodon, the "Kirtland Aublysodon", and Alectrosaurus. [35] Holtz also found that Siamotyrannus exhibited some of the synapomorphies of the tyrannosauridae, but lay "outside the [family] proper." [35]

Later in the same paper, he proposed that Tyrannosauridae be defined as "all descendants of the most recent common ancestor of Tyrannosaurus and Aublysodon". [35] He also criticized definitions previously proposed by other workers, like one proposed by Paul Sereno, that the Tyrannosauridae was "all taxa closer to "Tyrannosaurus" than to Alectrosaurus, Aublysodon, and Nanotyrannus". [35] Holtz observed that since Nanotyrannus was probably a misidentified T. rex juvenile, Sereno's proposed definition would have the family Tyrannosauridae as a subtaxon of the genus Tyrannosaurus. [35] Further, his proposed definition of the subfamily Tyrannosaurinae would also be limited to Tyrannosaurus. [35]

A 2003 attempt by Christopher Brochu included Albertosaurus, Alectrosaurus, Alioramus, Daspletosaurus, Gorgosaurus, Tarbosaurus and Tyrannosaurus in the definition. [36] Holtz redefined the clade in 2004 to use all of the above as specifiers except for Alioramus and Alectrosaurus, which his analysis could not place with certainty. However, in the same paper, Holtz also provided a completely different definition, including all theropods more closely related to Tyrannosaurus than to Eotyrannus. [10] The most recent definition is that of Sereno in 2005, which defined Tyrannosauridae as the least inclusive clade containing Albertosaurus, Gorgosaurus and Tyrannosaurus. [37]

Cladistic analyses of tyrannosaurid phylogeny often find Tarbosaurus and Tyrannosaurus to be sister taxa, with Daspletosaurus more basal than either. A close relationship between Tarbosaurus and Tyrannosaurus is supported by numerous skull features, including the pattern of sutures between certain bones, the presence of a crescent-shaped crest on the postorbital bone behind each eye, and a very deep maxilla with a noticeable downward curve on the lower edge, among others. [10] [15] An alternative hypothesis was presented in a 2003 study by Phil Currie and colleagues, which found weak support for Daspletosaurus as a basal member of a clade also including Tarbosaurus and Alioramus, both from Asia, based on the absence of a bony prong connecting the nasal and lacrimal bones. [20] Alioramus was found to be the closest relative of Tarbosaurus in this study, based on a similar pattern of stress distribution in the skull.

A related study also noted a locking mechanism in the lower jaw shared between the two genera. [38] In a separate paper, Currie noted the possibility that Alioramus might represent a juvenile Tarbosaurus, but stated that the much higher tooth count and more prominent nasal crests in Alioramus suggest it is a distinct genus. Similarly, Currie uses the high tooth count of Nanotyrannus to suggest that it may be a distinct genus, [11] rather than a juvenile Tyrannosaurus as most other experts believe. [10] [39] However, the discovery and description of Qianzhousaurus reveals that Alioramus is not a close relation to Tarbosaurus, instead belonging to a newly described tribe of tyrannosaurids the Alioramini. Qianzhousaurus further reveals that similar long-snouted tyrannosaurids were widely distributed throughout Asia and would have shared the same environment while avoiding competition with larger and more robust tyrannosaurines by hunting different prey. [40]

Growth Edit

Paleontologist Gregory Erickson and colleagues have studied the growth and life history of tyrannosaurids. Analysis of bone histology can determine the age of a specimen when it died. Growth rates can be examined when the age of various individuals are plotted against their size on a graph. Erickson has shown that after a long time as juveniles, tyrannosaurs underwent tremendous growth spurts for about four years midway through their lives. After the rapid growth phase ended with sexual maturity, growth slowed down considerably in adult animals. A tyrannosaurid growth curve is S-shaped, with the maximum growth rate of individuals around 14 years of age. [44]

The smallest known Tyrannosaurus rex individual (LACM 28471, the "Jordan theropod") is estimated to have weighed only 29.9 kilograms (66 lb) at only 2 years old, while the largest, such as FMNH PR2081 ("Sue"), most likely weighed about 5,654 kg (12,465 lb), estimated to have been 28 years old, an age which may have been close to the maximum for the species. [44] T. rex juveniles remained under 1,800 kg (4,000 lb) until approximately 14 years of age, when body size began to increase dramatically. During this rapid growth phase, a young T. rex would gain an average of 600 kg (1,300 lb) a year for the next four years. This slowed after 16 years, and at 18 years of age, the curve plateaus again, indicating that growth slowed dramatically. [45] For example, only 600 kg (1,300 lb) separated the 28-year-old "Sue" from a 22-year-old Canadian specimen (RTMP 81.12.1). [44] This sudden change in growth rate may indicate physical maturity, a hypothesis that is supported by the discovery of medullary tissue in the femur of an 18-year-old T. rex from Montana (MOR 1125, also known as "B-rex"). [46] Medullary tissue is found only in female birds during ovulation, indicating that "B-rex" was of reproductive age. [47]

Other tyrannosaurids exhibit extremely similar growth curves, although with lower growth rates corresponding to their lower adult sizes. [48] Compared to albertosaurines, Daspletosaurus showed a faster growth rate during the rapid growth period due to its higher adult weight. The maximum growth rate in Daspletosaurus was 180 kilograms (400 lb) per year, based on a mass estimate of 1,800 kg (4,000 lb) in adults. Other authors have suggested higher adult weights for Daspletosaurus this would change the magnitude of the growth rate, but not the overall pattern. [44] The youngest known Albertosaurus is a two-year-old discovered in the Dry Island bonebed, which would have weighed about 50 kg (110 lb) and measured slightly more than 2 metres (6.6 ft) in length. The 10-metre (33 ft) specimen from the same quarry is the oldest and largest known, at 28 years of age. The fastest growth rate is estimated to occur around 12–16 years of age, reaching 122 kg (269 lb) per year, based on a 1,300 kg (2,900 lb) adult, which is about a fifth of the rate for T.-rex. For Gorgosaurus, the calculated maximum growth rate is about 110 kilograms (240 lb) during the rapid growth phase, which is comparable to that of Albertosaurus. [44]

The discovery of an embryonic tyrannosaur of an as-yet-unknown genus suggests that tyrannosaurids developed their distinctive skeletal features while developing in the egg. Furthermore, the size of the specimen, a 1.1 in (2.8 cm) dentary from the lower jaw found in the Two Medicine Formation of Montana in 1983 and a foot claw found in the Horseshoe Canyon Formation in 2018 and described in 2020, suggests that neonate tyrannosaurids were born with skulls the size of a mouse or similarly sized rodents and may have been roughly the size of a small dog at birth. The jaw specimen is believed to have come from an animal roughly 2.5 ft (0.76 m) while the claw is believed to belong to a specimen measuring around 3 ft (0.91 m). While eggshells have not been found in association with either specimen, the location where these neonate tyrannosaurids were uncovered suggests these animals were using the same nest sites as other species they lived with and preyed upon. [49] The lack of eggshells associated with these specimens has also opened up speculation to the possibility that tyrannosaurids laid soft-shelled eggs as the genera Mussaurus and Protoceratops are believed to have done. [50]

Fossil footprints from the Wapiti Formation suggest that as tyrannosaurids grew, the feet became wider with thicker toes to support their weight. The broader feet suggest that adult tyrannosaurids were slower-moving than their offspring. [51] [52]

Life history Edit

The end of the rapid growth phase suggests the onset of sexual maturity in Albertosaurus, although growth continued at a slower rate throughout the animals' lives. [44] [48] Sexual maturation while still actively growing appears to be a shared trait among small [53] and large [54] dinosaurs as well as in large mammals, such as humans and elephants. [54] This pattern of relatively early sexual maturation differs strikingly from the pattern in birds, which delay their sexual maturity until after they have finished growing. [54] [55]

By tabulating the number of specimens of each age group, Erickson and his colleagues were able to draw conclusions about life history in tyranosauridae populations. Their analysis showed that while juveniles were rare in the fossil record, subadults in the rapid growth phase and adults were far more common. Over half of the known T. rex specimens appear to have died within six years of reaching sexual maturity, a pattern that is also seen in other tyrannosaurs and in some large, long-lived birds and mammals today. These species are characterized by high infant mortality rates, followed by relatively low mortality among juveniles. Mortality increases again following sexual maturity, partly due to the stresses of reproduction. While this could be due to preservation or collection biases, Erickson hypothesized that the difference was due to low mortality among juveniles over a certain size, which is also seen in some modern large mammals, like elephants. This low mortality may have resulted from a lack of predation, since tyrannosaurs surpassed all contemporaneous predators in size by the age of two. Paleontologists have not found enough Daspletosaurus remains for a similar analysis, but Erickson notes that the same general trend seems to apply. [48]

The tyrannosaurids spent as much as half its life in the juvenile phase before ballooning up to near-maximum size in only a few years. [44] This, along with the complete lack of predators intermediate in size between huge adult tyrannosaurids and other small theropods, suggests these niches may have been filled by juvenile tyrannosaurids. This is seen in modern Komodo dragons, where hatchlings start off as tree-dwelling insectivores and slowly mature into massive apex predators capable of taking down large vertebrates. [10] For example, Albertosaurus have been found in aggregations that some have suggested to represent mixed-age packs. [56] [57]

Locomotion Edit

Locomotion abilities are best studied for Tyrannosaurus, and there are two main issues concerning this: how well it could turn and what its maximum straight-line speed was likely to have been. Tyrannosaurus may have been slow to turn, possibly taking one to two seconds to turn only 45° – an amount that humans, being vertically oriented and tail-less, can spin in a fraction of a second. [58] The cause of the difficulty is rotational inertia, since much of Tyrannosaurus ' s mass was some distance from its center of gravity, like a human carrying a heavy timber. [59]

Scientists have produced a wide range of maximum speed estimates, mostly around 11 metres per second (25 mph), but a few as low as 5–11 metres per second (11–25 mph), and a few as high as 20 metres per second (45 mph). Researchers have to rely on various estimating techniques because, while there are many tracks of very large theropods walking, so far none have been found of very large theropods running—and this absence may indicate that they did not run. [60]

Jack Horner and Don Lessem argued in 1993 that Tyrannosaurus was slow and probably could not run (no airborne phase in mid-stride). [61] However, Holtz (1998) concluded that tyrannosaurids and their close relatives were the fastest large theropods. [62] Christiansen (1998) estimated that the leg bones of Tyrannosaurus were not significantly stronger than those of elephants, which are relatively limited in their top speed and never actually run (there is no airborne phase), and hence proposed that the dinosaur's maximum speed would have been about 11 metres per second (25 mph), which is about the speed of a human sprinter. [63] Farlow and colleagues (1995) have argued that a 6- to 8-ton Tyrannosaurus would have been critically or even fatally injured if it had fallen while moving quickly, since its torso would have slammed into the ground at a deceleration of 6 g (six times the acceleration due to gravity, or about 60 metres/s 2 ) and its tiny arms could not have reduced the impact. [64] [65] However, giraffes have been known to gallop at 50 km/h (31 mph), despite the risk that they might break a leg or worse, which can be fatal even in a "safe" environment such as a zoo. [66] [67] Thus it is quite possible that Tyrannosaurus also moved fast when necessary and had to accept such risks this scenario has been studied for Allosaurus too. [68] [69] Most recent research on Tyrannosaurus locomotion does not narrow down speeds further than a range from 17 to 40 km/h (11 to 25 mph), i.e. from walking or slow running to moderate-speed running. [60] [70] [71] A computer model study in 2007 estimated running speeds, based on data taken directly from fossils, and claimed that T. rex had a top running speed of 8 metres per second (18 mph). [72] [73] (Probably a juvenile individual. [74] )

Studies by Eric Snively et al., published in 2019 indicate that tyrannosaurids such as Tarbosaurus and Tyrannosaurus itself were more maneuverable than allosauroids of comparable size due to low rotational inertia compared to their body mass combined with large leg muscles. As a result, it is hypothesized that tyrannosaurids were capable of making relatively quick turns and could likely pivot their bodies more quickly when close to their prey, or that while turning, they could "pirouette" on a single planted foot while the alternating leg was held out in a suspended swing during pursuit. The results of this study potentially could shed light on how agility could have contributed to the success of tyrannosaurid evolution. [75]

Additionally, a 2020 study indicates that tyrannosaurids were exceptionally efficient walkers. Studies by Dececchi et al., compared the leg proportions, body mass, and the gaits of more than 70 species of theropod dinosaurs including tyrannosaurids. The research team then applied a variety of methods to estimate each dinosaur's top speed when running as well as how much energy each dinosaur expended while moving at more relaxed speeds such as when walking. Among smaller to medium-sized species such as dromaeosaurids, longer legs appear to be an adaptation for faster running, in line with previous results by other researchers. But for theropods weighing over 1,000 kg (2,200 lb), top running speed is limited by body size, so longer legs instead were found to have correlated with low-energy walking. The results of the study further indicated that smaller theropods evolved long legs for speed as a means to both aid in hunting and escape from larger predators while larger predatory theropods that evolved long legs did so to reduce the energy costs and increase foraging efficiency, as they were freed from the demands of predation pressure due to their role as apex predators. Compared to more basal groups of theropods in the study, tyrannosaurids showed a marked increase in foraging efficiency due to reduced energy expenditures during hunting and scavenging. This likely resulted in tyrannosaurs having a reduced need for hunting forays and requiring less food to sustain themselves as a result. Additionally, the research, in conjunction with studies that show tyrannosaurs were more agile than other large bodied-theropods, indicates they were quite well-adapted to a long-distance stalking approach followed by a quick burst of speed to go for the kill. Analogies can be noted between tyrannosaurids and modern wolves as a result, supported by evidence that at least some tyrannosaurids such as Albertosaurus were hunting in group settings. [76] [77]

Integument Edit

An ongoing debate in the paleontological community surrounds the extent and nature of tyrannosaurid integumentary covering. Long filamentous structures have been preserved along with skeletal remains of numerous coelurosaurs from the Early Cretaceous Yixian Formation and other nearby geological formations from Liaoning, China. [78] These filaments have usually been interpreted as "protofeathers," homologous with the branched feathers found in birds and some non-avian theropods, [79] [80] although other hypotheses have been proposed. [81] A skeleton of Dilong was described in 2004 that included the first example of "protofeathers" in a tyrannosauroid. Similarly to down feathers of modern birds, the "protofeathers" found in Dilong were branched but not pennaceous, and may have been used for insulation. [18] The discovery and description of the 9-metre (30 ft) feathered tyrannosauroid Yutyrannus in 2012 indicates the possibility large tyrannosaurids were also feathered as adults. [82]

Based on the principle of phylogenetic bracketing, it was predicted that tyrannosaurids might also possess such feathering. However, a study in 2017 published by a team of researchers in Biology Letters described tyrannosaurid skin impressions collected in Alberta, Montana and Mongolia, which came from five genera (Tyrannosaurus, Albertosaurus, Gorgosaurus, Daspletosaurus and Tarbosaurus). [83] Although the skin impressions are small, they are widely dispersed across the post-cranium, being collectively located on the abdomen, thoracic region, ilium, pelvis, tail and neck. They show a tight pattern of fine, non-overlapping pebbly scales (which co-author Scott Persons compared to those seen on the flanks of a crocodile [84] ) and preserve no hints of feathering. The basic texture is composed of tiny "basement scales" approximately 1 to 2 mm in diameter, with some impressions showing 7 mm "feature scales" interspersed between them. Additional scales can be seen in tyrannosaurid footprints [85] and potential ostelogical correlates for scales are present on the skull. [86]

Bell et al. performed an ancestral character reconstruction based on what is known about integument distribution in tyrannosauroids. Despite an 89% probability that tyrannosauroids started out with feathers, they determined that scaly tyrannosaurids have a 97% probability of being true. The data "provides compelling evidence of an entirely squamous covering in Tyrannosaurus," the team wrote, although they conceded that plumage may have still been present on the dorsal region where skin impressions haven't been found yet. [83]

The paper has been questioned by palaeontologists such as Andrea Cau and Thomas Holtz, who point out that feathers in theropod dinosaurs can grow in the same parts of the body as scales (as seen in Juravenator), and thus the presence of scales does not actually eliminate the presence of feathers from that part of the body. [87] Furthermore, feathers are delicate structures which can be very easily lost due to taphonomic factors. Paleontologist Mark Witton was warmer to the study, suggesting "we have to concede a scalier appearance than many of us thought likely", while also highlighting that much remains to be determined about tyrannosaurid life appearance. Though acknowledging the role that taphonomy plays in interpreting dinosaur skin, Witton points out that the scale impressions are high quality, consistent in form across all patches, and show no obvious evidence of fibre impressions or spaces for filament attachment, despite the preservation of submillimeter scales. More variation and evidence of deformation might be expected if taphonomy had distorted the specimens in a significant way. [88]

It has yet to be determined why such an integumentary change might have occurred. A precedent for feather loss can be seen in other dinosaur groups such as ornithischians, in which filamentous structures were lost, and scales reappeared. [89] Although gigantism has been suggested as a mechanism, Phil R. Bell, who co-authored the study, noted that the feathered Yutyrannus overlapped in size with Gorgosaurus and Albertosaurus. "The problem here is that we have big tyrannosaurs, some with feathers, some without that live in pretty similar climates. So what's the reason for this difference? We really don't know." [90]

Vision Edit

The eye-sockets of Tyrannosaurus are positioned so that the eyes would point forward, giving them binocular vision slightly better than that of modern hawks. While predatory theropods in general had binocular vision directly in front of their skull, tyrannosaurs had a significantly larger area of overlap. Jack Horner also pointed out that the tyrannosaur lineage had a history of steadily improving binocular vision. It is hard to see how natural selection would have favored this long-term trend if tyrannosaurs had been pure scavengers, which would not have needed the advanced depth perception that stereoscopic vision provides. [91] [92] In modern animals, binocular vision is found mainly in predators (the principal exceptions are primates, which need it for leaping from branch to branch). Unlike Tyrannosaurus, Tarbosaurus had a narrower skull more typical of other tyrannosaurids in which the eyes faced primarily sideways. All of this suggests that Tarbosaurus relied more on its senses of smell and hearing than on its eyesight. [93] In Gorgosaurus specimens, the eye socket was circular rather than oval or keyhole-shaped as in other tyrannosaurid genera. [11] In Daspletosaurus, this was a tall oval, somewhere in between the circular shape seen in Gorgosaurus and the 'keyhole' shape of Tyrannosaurus. [10] [11] [39]

Facial soft tissue Edit

Based on comparisons of bone texture of Daspletosaurus with extant crocodilians, a detailed study in 2017 by Thomas D. Carr et al. found that tyrannosaurs had large, flat scales on their snouts. [94] [95] At the center of these scales were small keratinised patches. In crocodilians, such patches cover bundles of sensory neurons that can detect mechanical, thermal and chemical stimuli. [96] [97] They proposed that tyrannosaurs probably also had bundles of sensory neurons under their facial scales and may have used them to identify objects, measure the temperature of their nests and gently pick-up eggs and hatchlings. [94] A 2018 study however did not agree with this and rather suggested a lip condition. Extant crocodillians don't have scales but rather cracked skin. They were analyzing the rugosity of tyrannosaurids and found a hummocky rugosity which would favor squamose-like scales that appeared in life. [98] [99]

Bony crests Edit

Bony crests are found on the skulls of many theropods, including many tyrannosaurids. Alioramus, a possible tyrannosaurid from Mongolia, bears a single row of five prominent bony bumps on the nasal bones a similar row of much lower bumps is present on the skull of Appalachiosaurus, as well as some specimens of Daspletosaurus, Albertosaurus, and Tarbosaurus. [15] In Albertosaurus, Gorgosaurus and Daspletosaurus, there is a prominent horn in front of each eye on the lacrimal bone. The lacrimal horn is absent in Tarbosaurus and Tyrannosaurus, which instead have a crescent-shaped crest behind each eye on the postorbital bone. These head crests may have been used for display, perhaps for species recognition or courtship behavior. [10]

Thermoregulation Edit

Tyrannosaurus, like most dinosaurs, was long thought to have an ectothermic ("cold-blooded") reptilian metabolism but was challenged by scientists like Robert T. Bakker and John Ostrom in the early years of the "Dinosaur Renaissance", beginning in the late 1960s. [100] [101] Tyrannosaurus rex itself was claimed to have been endothermic ("warm-blooded"), implying a very active lifestyle. [102] Since then, several paleontologists have sought to determine the ability of Tyrannosaurus to regulate its body temperature. Histological evidence of high growth rates in young T. rex, comparable to those of mammals and birds, may support the hypothesis of a high metabolism. Growth curves indicate that, as in mammals and birds, T. rex growth was limited mostly to immature animals, rather than the indeterminate growth seen in most other vertebrates. [45] It has been indicated that the temperature difference may have been no more than 4 to 5 °C (7 to 9 °F) between the vertebrae of the torso and the tibia of the lower leg. This small temperature range between the body core and the extremities was claimed by paleontologist Reese Barrick and geochemist William Showers to indicate that T. rex maintained a constant internal body temperature (homeothermy) and that it enjoyed a metabolism somewhere between ectothermic reptiles and endothermic mammals. [103] Later they found similar results in Giganotosaurus specimens, who lived on a different continent and tens of millions of years earlier in time. [104] Even if Tyrannosaurus rex does exhibit evidence of homeothermy, it does not necessarily mean that it was endothermic. Such thermoregulation may also be explained by gigantothermy, as in some living sea turtles. [105] [106] [107]

Coexistence of Daspletosaurus and Gorgosaurus Edit

In the Dinosaur Park Formation, Gorgosaurus lived alongside a rarer species of the tyrannosaurine Daspletosaurus. This is one of the few examples of two tyrannosaur genera coexisting. Similarly sized predators in modern predator guilds are separated into different ecological niches by anatomical, behavioral or geographical differences that limit competition. Niche differentiation between the Dinosaur Park tyrannosaurids is not well understood. [108] In 1970, Dale Russell hypothesized that the more common Gorgosaurus actively hunted fleet-footed hadrosaurs, while the rarer and more troublesome ceratopsians and ankylosaurians (horned and heavily armoured dinosaurs) were left to the more heavily built Daspletosaurus. [12] However, a specimen of Daspletosaurus (OTM 200) from the contemporaneous Two Medicine Formation of Montana preserves the digested remains of a juvenile hadrosaur in its gut region. [109] Unlike some other groups of dinosaurs, neither genus was more common at higher or lower elevations than the other. [108] However, Gorgosaurus appears more common in northern formations like the Dinosaur Park, with species of Daspletosaurus more abundant to the south. The same pattern is seen in other groups of dinosaurs. Chasmosaurine ceratopsians and hadrosaurine hadrosaurs are also more common in the Two Medicine Formation of Montana and in southwestern North America during the Campanian, while centrosaurines and lambeosaurines dominate in northern latitudes. Holtz has suggested that this pattern indicates shared ecological preferences between tyrannosaurines, chasmosaurines and hadrosaurines. At the end of the later Maastrichtian stage, tyrannosaurines like Tyrannosaurus rex, hadrosaurines like Edmontosaurus and chasmosaurines like Triceratops were widespread throughout western North America, while albertosaurines and centrosaurines became extinct, and lambeosaurines were rare. [10]

Social behavior Edit

There is limited evidence of social behavior among the tyrannosaurids. Researchers reported that a subadult and a juvenile skeleton were found in the same quarry as the "Sue" specimen, which has been used to support the hypothesis that tyrannosaurs may have lived in social groups of some kind. [110] While there is no evidence of gregarious behavior in Gorgosaurus, [56] [57] there is evidence of some pack behavior for Albertosaurus and Daspletosaurus.

A young specimen of the Dinosaur Park Daspletosaurus species (TMP 94.143.1) shows bite marks on the face that were inflicted by another tyrannosaur. The bite marks are healed over, indicating that the animal survived the bite. A full-grown Dinosaur Park Daspletosaurus (TMP 85.62.1) also exhibits tyrannosaur bite marks, showing that attacks to the face were not limited to younger animals. While it is possible that the bites were attributable to other species, intraspecific aggression, including facial biting, is very common among predators. Facial bites are seen in other tyrannosaurs like Gorgosaurus and Tyrannosaurus, as well as in other theropod genera like Sinraptor and Saurornitholestes. Darren Tanke and Phil Currie hypothesize that the bites are due to intraspecific competition for territory or resources, or for dominance within a social group. [56]

Evidence that Daspletosaurus lived in social groups comes from a bonebed found in the Two Medicine Formation of Montana. The bonebed includes the remains of three Daspletosaurus, including a large adult, a small juvenile, and another individual of intermediate size. At least five hadrosaurs are preserved at the same location. Geologic evidence indicates that the remains were not brought together by river currents but that all of the animals were buried simultaneously at the same location. The hadrosaur remains are scattered and bear many marks from tyrannosaur teeth, indicating that the Daspletosaurus were feeding on the hadrosaurs at the time of death. The cause of death is unknown. Currie speculates that the daspletosaurs formed a pack, although this cannot be stated with certainty. [57] Other scientists are skeptical of the evidence for social groups in Daspletosaurus and other large theropods [111] Brian Roach and Daniel Brinkman have suggested that Daspletosaurus social interaction would have more closely resembled the modern Komodo dragon, where non-cooperative individuals mob carcasses, frequently attacking and even cannibalizing each other in the process. [112]

The Dry Island bonebed discovered by Barnum Brown and his crew contains the remains of 22 Albertosaurus, the most individuals found in one locality of any Cretaceous theropod, and the second-most of any large theropod dinosaur behind the Allosaurus assemblage at the Cleveland-Lloyd Dinosaur Quarry in Utah. The group seems to be composed of one very old adult eight adults between 17 and 23 years old seven sub-adults undergoing their rapid growth phases at between 12 and 16 years old and six juveniles between the ages of 2 and 11 years, who had not yet reached the growth phase. [48] The near-absence of herbivore remains and the similar state of preservation between the many individuals at the Albertosaurus bonebed quarry led Phil Currie to conclude that the locality was not a predator trap like the La Brea Tar Pits in California, and that all of the preserved animals died at the same time. Currie claims this as evidence of pack behavior. [113] Other scientists are skeptical, observing that the animals may have been driven together by drought, flood or for other reasons. [48] [111] [114]

While it generally remains controversial, evidence does exist that supports the theory that at least some tyrannosaurids were social. In British Columbia's Wapiti Formation, a trackway composed of the footprints of three individual tyrannosaurids (named as the ichnogenus Bellatoripes fredlundi) was discovered by a local outfitter named Aaron Fredlund and described in the journal PLOS One by Richard McCrea et al. An examination of the trackway found no evidence of one trackway being left long after another had been made, further supporting the hypothesis that three individual tyrannosaurs were traveling together as a group. Further research revealed the animals were traveling at a speed of between 3.9 and 5.2 mph (6.3 and 8.4 km/h) and likely had a hip height of around 7 to 9 feet. As three different genera of tyrannosaurids (Gorgosaurus, Daspletosaurus, and Albertosaurus, respectively) are known from the formation, it is unknown which genus was the maker of the trackway. [115] [116] [117]

Feeding Edit

Tyrannosaur tooth marks are the most commonly preserved feeding traces of carnivorous dinosaurs. [118] They have been reported from ceratopsians, hadrosaurs and other tyrannosaurs. [118] Tyrannosaurid bones with tooth marks represent about 2% of known fossils with preserved tooth marks. [118] Tyrannosaurid teeth were used as holdfasts for pulling meat off a body, rather than knife-like cutting functions. [119] Tooth wear patterns hint that complex head shaking behaviors may have been involved in tyrannosaur feeding. [119]

Speculation on the pack-hunting habits of Albertosaurus were made by a few researchers who suggest that the younger members of the pack may have been responsible for driving their prey towards the adults, who were larger and more powerful, but also slower. [113] Juveniles may also have had different lifestyles than adults, filling predator niches between those of the enormous adults and the smaller contemporaneous theropods, the largest of which were two orders of magnitude smaller than an adult Albertosaurus in mass. [10] However, as the preservation of behavior in the fossil record is exceedingly rare, these ideas cannot readily be tested. Phil Currie speculates that the Daspletosaurus formed packs to hunt, although this cannot be stated with certainty. [57] There is no evidence of such gregarious behavior in Gorgosaurus. [56] [57]

The debate about whether Tyrannosaurus was a predator or a pure scavenger is as old as the debate about its locomotion. Lambe (1917) described a good skeleton of Tyrannosaurus ' s close relative Gorgosaurus and concluded that it and therefore also Tyrannosaurus was a pure scavenger, because the Gorgosaurus ' s teeth showed hardly any wear. [120] This argument is no longer taken seriously, because theropods replaced their teeth quite rapidly. Ever since the first discovery of Tyrannosaurus most scientists have agreed that it was a predator, although like modern large predators it would have been happy to scavenge or steal another predator's kill if it had the opportunity. [121] [122]

Noted hadrosaur expert Jack Horner is currently the major advocate of the idea that Tyrannosaurus was exclusively a scavenger and did not engage in active hunting at all. [61] [123] [124] Horner has presented several arguments to support the pure scavenger hypothesis. The presence of large olfactory bulbs and olfactory nerves suggests a highly developed sense of smell for sniffing out carcasses over great distances. The teeth could crush bone, and therefore could extract as much food (bone marrow) as possible from carcass remnants, usually the least nutritious parts. At least some of its potential prey could move quickly, while evidence suggests that Tyrannosaurus walked instead of ran. [123] [125]

Other evidence suggests hunting behavior in Tyrannosaurus. The eye-sockets of tyrannosaurs are positioned so that the eyes would point forward, giving them binocular vision slightly better than that of modern hawks. Tyrannosaur-inflicted damage has been found on skeletons of hadrosaurs and Triceratops that seemed to have survived initial attacks. [126] [127] [128] Some researchers argue that if Tyrannosaurus were a scavenger, another dinosaur had to be the top predator in the Amerasian Upper Cretaceous. The top prey were the larger marginocephalians and ornithopods. The other tyrannosaurids share so many characteristics with Tyrannosaurus that only small dromaeosaurs remain as feasible top predators. In this light, scavenger hypothesis adherents have suggested that the size and power of tyrannosaurs allowed them to steal kills from smaller predators. [125]

Cannibalism Edit

Evidence also strongly suggests that tyrannosaurids were at least occasionally cannibalistic. Tyrannosaurus itself has strong evidence pointing towards it having been cannibalistic in at least a scavenging capacity based on tooth marks on the foot bones, humerus, and metatarsals of one specimen. [129] Fossils from the Fruitland Formation, Kirtland Formation (both Campanian in age) and the Maastichtian aged Ojo Alamo Formation suggest that cannibalism was present in various tyrannosaurid genera of the San Juan Basin. The evidence gathered from the specimens suggests opportunistic feeding behavior in tyrannosaurids that cannibalized members of their own species. [130]

Distribution Edit

While earlier tyrannosauroids are found on all three northern continents, tyrannosaurid fossils are known only from North America and Asia. Sometimes fragmentary remains uncovered in the Southern Hemisphere have been reported as "Southern Hemisphere tyrannosaurids," although these seem to have been misidentified abelisaurid fossils. [131] The exact time and place of origin of the family remain unknown due to the poor fossil record in the middle part of the Cretaceous on both continents, although the earliest confirmed tyrannosaurids lived in the early Campanian stage in western North America. [10]

Tyrannosaurid remains have never been recovered from eastern North America, while more basal tyrannosauroids, like Dryptosaurus and Appalachiosaurus, persisted there until the end of the Cretaceous, indicating that tyrannosaurids must have evolved in or dispersed into western North America after the continent was divided in half by the Western Interior Seaway in the middle of the Cretaceous. [15] Tyrannosaurid fossils have been found in Alaska, which may have provided a route for dispersal between North America and Asia. [132] Alioramus and Tarbosaurus are found to be related in one cladistic analysis, forming a unique Asian branch of the family. [20] This was later disproven with the discovery of Qianzhousaurus and the description of the tyrannosaur family Alioramini. Tyrannosaurid teeth from a large species of unknown variety were discovered in the Nagasaki Peninsula by researchers from the Fukui Prefectural Dinosaur Museum, further expanding the range of the group. The teeth were estimated to be 81 million years old (Campanian Age). [133]

Of the two subfamilies, tyrannosaurines appear to have been more widespread. Albertosaurines are unknown in Asia, which was home to the tyrannosaurines, such as Tarbosaurus and Zhuchengtyrannus, and Qianzhousaurus and Alioramus of the Alioramini. Both the Tyrannosaurinae and Albertosaurinae subfamilies were present in the Campanian and early Maastrichtian stages of North America, with tyrannosaurines like Daspletosaurus ranging throughout the Western Interior, while the albertosaurines Albertosaurus and Gorgosaurus are currently known only from the northwestern part of the continent. [134]

By the late Maastrichtian, albertosaurines appear to have gone extinct, while the tyrannosaurine Tyrannosaurus roamed from Saskatchewan to Texas. This pattern is mirrored in other North American dinosaur taxa. During the Campanian and early Maastrichtian, lambeosaurine hadrosaurs and centrosaurine ceratopsians are common in the northwest, while hadrosaurines and chasmosaurines were more common to the south. By the end of the Cretaceous, centrosaurines are unknown and lambeosaurines are rare, while hadrosaurines and chasmosaurines were common throughout the Western Interior. [10] A study published in the journal Scientific Reports on February 2, 2016 by Steve Brusatte, Thomas Carr et al. indicates that during the later Maastrichtian, Tyrannosaurus itself might have been partially responsible for the extinction of the other tyrannosaurids in most of western North America. The study indicates that Tyrannosaurus might have been an immigrant from Asia as opposed to having evolved in North America (possibly a descendant of the closely related Tarbosaurus) that supplanted and outcompeted other tyrannosaurids. This theory is further supported by the fact that few to no other types of tyrannosaurid are found within Tyrannosaurus' known range. [135]


Introduction

Predation is known to be an important biological factor with direct (e.g. prey mortality) and indirect effects (e.g. effects on organisms associated with the prey species) on population dynamics and community structure of marine ecosystems (e.g. Hughes, 1980, Menge et al., 1986). The rocky intertidal is recognised as a particularly tractable system to examine predator–prey interactions since predation effects are thought to be stronger here than in freshwater and terrestrial habitats (Sih et al., 1985). Predation is also known to interact with other biological (e.g. competition, Menge and Sutherland, 1976) and physical factors (e.g. wave exposure, Menge, 1978a), influencing its effects on prey populations (Menge, 1991, Menge, 2000, Menge and Sutherland, 1987). Hence, predator foraging and therefore predation pressure, are modulated by biological constraints and environmental factors, which subsequently determine realized foraging patterns (Lawton and Zimmerfaust, 1992, Thompson et al., 2004).

Crabs are highly mobile and are known to have an important role in structuring assemblages on rocky shores worldwide (e.g. Burrows et al., 1999, Ebling et al., 1964, Rilov and Schiel, 2006, Robles, 1987, Silva et al., 2008). It is therefore important to better understand the mechanisms of crab predation at the individual level in order to better understand their effects on prey populations. Traditionally, studies on predator–prey interactions have focused either on lethal effects on prey such as mortality (e.g. Rilov and Schiel, 2006, Silva et al., 2004, Silva et al., 2008, Thompson et al., 2000), or on non-lethal effects such as morphology (e.g. Dalziel and Boulding, 2005, Trussell, 1996, Vermeij, 1978), physiology (e.g. Moller and Beress, 1975), life history or behaviour (e.g. Phillips, 1976, Sih, 1987). As for specific predator–prey studies involving hard-shelled prey, durophagous crabs have been observed to crush and/or peel molluscan shells (Shoup, 1968, Takeda and Suga, 1979, Vermeij, 1977). Many crabs such as Calappa spp. crabs, are capable of accessing prey by peeling the shell aperture and thus avoid amore time and energy-consuming behaviour such as shell crushing (e.g. Shoup, 1968). This specialization of crab feeding behaviour reflects also on morphology, with changes in claw dentition developing to meet the demands of peeling the prey shell (Yamada and Boulding, 1998). Despite such a specialization in feeding behaviour of crabs towards their molluscan prey, there are also reports of generalization in crab feeding, such as a widespread preference to feed on small-sized molluscan prey, which are presumably easier to crush and so offer higher net energy gain (Juanes, 1992).

Studies on the predator–prey arms race between crabs and their gastropod prey are abundant (e.g. Bertness and Cunningham, 1981, Boulding, 1984, Brookes and Rochette, 2007, Cotton et al., 2004, Lowell, 1986), and crabs have been reported to have larger claws in locations where the most consumed prey are shelled molluscs (e.g. Yamada and Boulding, 1998). These studies have, however, mainly focused on consequences in terms of defences developed by prey, and predator responses to the interaction with prey often are overlooked. Hence, there is limited information on the cheliped morphological responses to the crab foraging behaviour, taking into consideration variations in the physical environment and in prey availability (interpreted here as abundance).

Phenotypic plasticity is the ability of a particular genotype to produce different phenotypes in response to environmental variation (DeWitt and Scheiner, 2004). Studies on crab flexibility have, however, mainly been investigated in terms of behaviour (e.g. Briffa et al., 1998, Hazlett, 1995), and there is little information on phenotypic flexibility according to differing environmental conditions (but see for example Freire et al., 1996, Hughes, 2000, Yamada and Boulding, 1996) and associated functional responses. The cheliped of crabs are important for reproduction (Hughes, 2000, Lee, 1995) and are crucial for feeding, being used to crush or detach molluscan prey (Hughes and Elner, 1979, Iwasaki, 1993, Silva et al., 2008, Yamada and Boulding, 1998). Cheliped morphology is known to vary according to both diet (Brown et al., 1979, Elner, 1978, Freire et al., 1996, Hughes, 2000, Smith and Palmer, 1994) and mating interactions (Lee and Seed, 1992). To our knowledge, there is no evidence that chelipeds are a key feature in the reproduction for Eriphia verrucosa. Little has been done, however, to describe relationships between external physical factors and variation in the cheliped size and shape (but see Takeda and Murai, 2003), and the mechanisms underlying phenotypic responses to the environment are not well understood (DeWitt and Scheiner, 2004). Specifically, differences in predator morphological traits in relation to wave action have not previously been examined. Such information is important because patterns of claw morphology can have direct consequences for prey populations and potentially also have evolutionary consequences for their morphology (see Vermeij, 1982, Vermeij et al., 1981).

In the intertidal, prey composition and abundance vary along environmental gradients (Lewis, 1964, Raffaelli and Hawkins, 1996, Stephenson and Stephenson, 1949) thus it becomes important to examine differences in predator foraging characteristics at this scale. Here we focus on the role of exposure to wave action, an important environmental gradient in intertidal habitats, on the foraging pattern of the xanthid crab E. verrucosa. This species was used as a model organism because it is ubiquitous on exposed and sheltered shores in southern Europe (for a Portuguese reference see Flores and Paula, 2001). We examined natural variations in stomach content composition of E. verrucosa and hypothesized that crabs from sheltered locations would differ in cheliped form and size from those at exposed locations, and that this would be related to differences in prey abundance and consumption. At the same time, we aimed to describe the abundance, stomach content composition and population structure of E. verrucosa on shores of differing exposure. The sex of predators was included in the analysis since this can influence predator distribution, behaviour and diet (e.g. Bishop and Wear, 2005, Brousseau et al., 2001, Buck et al., 2003, Mascaró and Seed, 2001, Spooner et al., 2007). Patterns of prey abundance were also examined between shores of differing exposure and related to stomach content composition of E. verrucosa.

The following specific null hypothesis were examined in relation to shores of differing exposure: (1) there are no differences in prey abundance (2) there are no differences in the abundance and population structure of E. verrucosa (3) there are no differences in stomach content composition (4) there are no differences in claw size or shape between sexes (5) claw shape and/or size do not explain possible differences in stomach content composition (6) there is no relationship between crab size (carapace width and claw size) and the percentage of hard-shelled prey found in their stomachs.


Materials and methods

Animals and surfaces used in experiments

Beetles (Pachnoda marginata Drury, Scarabaeoidea) were obtained in the larval stage from a supplier. After pupation and hatching, adults were kept under normal room conditions (20-24 °C). Beetles were individually weighted prior to the experiments (mass=1.002±0.233 g, mean ±S.D., N=9). Intact and broken claws were air-dried for 4 weeks,sputter-coated with gold-palladium (10 nm) and examined in a Hitachi S-800 scanning electron microscope (SEM) at 20 kV. A SEM study of the claw material was also carried out with a freshly fixed, dehydrated and critical-point-dried claw. There was no significant difference in the structure of this stiff material from claws that had been air-dried or critical-point-dried.

The sandpaper used for the experiments is covered by Al2O3 particles (Wirtz-Buehler GmbH, Düsseldorf,Germany). The grit number, particle diameter and surface roughness(Ra) are listed in Table 1. Surface profile was measured using the perthometer M1 (Mahr GmbH, Göttingen, Germany). Ra was defined as the square root value of the difference between heights to its average height (see Appendix A). Rawas not measured for the sandpaper types P60 and P100, because their roughness was beyond the measuring range of the perthometer.

Variables of the sandpaper used in experiments

. Type of sandpaper . . . . . .
Variables . P60 † . P120 † . P280 * . P400 * . P0 * . 12 μm † .
Mean diameter (μm) 269 125 52.2±2.0 35.0±1.5 18.3±1.0 12
Roughness Ra (μm) - - 8.464 7.999 5.996 2.408
F/W from experiments 38 22 26 19 3 0.9
. Type of sandpaper . . . . . .
Variables . P60 † . P120 † . P280 * . P400 * . P0 * . 12 μm † .
Mean diameter (μm) 269 125 52.2±2.0 35.0±1.5 18.3±1.0 12
Roughness Ra (μm) - - 8.464 7.999 5.996 2.408
F/W from experiments 38 22 26 19 3 0.9

Values are means ± S.D. * N=10 † roughness data obtained from supplier.

F/W, force ratio (see Fig. 7).

Force measurements of beetles walking on different textures

To measure forces generated by walking beetles on different textures, a force sensor (load cell force transducer, 10 g) was used (Biopac System Inc. USA), mounted on a stand connected to the platform(Fig. 1). A metallic cross beam was used to transmit the beetle force to the sensor. One side of the crossbeam was connected to the sensor and another supported by the platform. The height of the beam was adjustable in a vertical direction. The distance between the beam and the sandpaper surface was adjusted to a suitable height (D)corresponding to the beetle's height (5-7 mm) plus 1-3 mm. The force sensor was attached to an amplifier and computer-based data-acquisition and processing system. The beetle's position was monitored by a video camera mounted on a binocular microscope. Experiments were video-recorded, and geometrical parameters were obtained from single video-frames.

Force measurement system of the beetle Pachnoda marginata. The system consists of a platform covered by sandpaper, force sensor, and videorecorder. 100 mm long crossbeam was connected to the force sensor on one side and supported on the other side. The height of the beam (D) relative to the sandpaper plane was adjusted to be 1-3 mm higher than the dorsal surface of the walking beetle. The force generated by the beetle was transferred by the beam to the sensor and monitored by the load cell force transducer and the signal amplified by the MP-100 system (Biopac system Inc. USA). The data were finally sampled and processed with the aid of a computer. A binocular microscope equipped with a video camera connected to a videorecorder was used to collect images of the beetle during the force measurements. From these images, the distance from the beetle to the sensor (X) was obtained. Force generated by the beetle FB was calculated as FB=100FS/(100-X), where FS is the force measured with the sensor.

Force measurement system of the beetle Pachnoda marginata. The system consists of a platform covered by sandpaper, force sensor, and videorecorder. 100 mm long crossbeam was connected to the force sensor on one side and supported on the other side. The height of the beam (D) relative to the sandpaper plane was adjusted to be 1-3 mm higher than the dorsal surface of the walking beetle. The force generated by the beetle was transferred by the beam to the sensor and monitored by the load cell force transducer and the signal amplified by the MP-100 system (Biopac system Inc. USA). The data were finally sampled and processed with the aid of a computer. A binocular microscope equipped with a video camera connected to a videorecorder was used to collect images of the beetle during the force measurements. From these images, the distance from the beetle to the sensor (X) was obtained. Force generated by the beetle FB was calculated as FB=100FS/(100-X), where FS is the force measured with the sensor.

The sensor was calibrated at the point where the crossbeam was connected before and after an experiment (sensitivity 10 μN). 3-5 beetles with three repetitions per individual were used for the experiment with each sandpaper.

Forces of the freely moving legs

To evaluate forces generated by a freely moving single leg, the beetle was fixed to a micromanipulator (World Precision Instruments Inc.), enabling adjustment of the beetle position relative to the sensor. Whenever the beetle grasped and pulled the sensor tip, the force was monitored and recorded. Five repetitions each for the forelegs, midlegs and hindlegs were done in three individual beetles.

Mechanical strength of the claw

The mechanical properties of the claw were tested on a Biotester Basalt-01(Tetra GmbH, Ilmenau, Germany) (for details, see Gorb et al., 2000). The claw of a freshly killed beetle was glued with cyanacrylat glue (5925 Universal, S. Kisling & Cie AG Zürich, Switzerland) to the platform. The metal spring was moved downwards, pressing with its tip against the claw tip until the claw was broken (Fig. 4A). The deflection of the spring tip was monitored by the fiber-optical sensor. Knowing the spring constant, the deflection was recalculated in the force. The maximum force of force—distance curves was used for calculations of braking stress. Since claw geometry of the fore-, mid- and hindlegs is constant (Fig. 2A), seven claws from different legs were tested.

Measurements of the mechanical strength of the claw. (A) Principle of force measurements using the force tester Basalt-01 (Tetra GmbH, Ilmenau, Germany). The claw (CL) was glued to the platform (PF). The metal spring (MS) was driven downwards by the piezo-drive until the claw was broken. Displacement of the spring tip equipped with a mirror M was detected in the vertical direction by the fiber optical sensor (FOS). (B) Force—distance curve obtained in experiments with the claw and used to calculate breaking stress of the claw.

Measurements of the mechanical strength of the claw. (A) Principle of force measurements using the force tester Basalt-01 (Tetra GmbH, Ilmenau, Germany). The claw (CL) was glued to the platform (PF). The metal spring (MS) was driven downwards by the piezo-drive until the claw was broken. Displacement of the spring tip equipped with a mirror M was detected in the vertical direction by the fiber optical sensor (FOS). (B) Force—distance curve obtained in experiments with the claw and used to calculate breaking stress of the claw.

Claw geometry. (A) Claw shape of the hind-, mid- and forelegs (drawings are based on SEM micrographs). (B) Five arcs used for quantitative description of the claw geometry (see Table 2). (C) Cross-section model of the claw for stress calculation. The model is based on SEM data (see Fig. 3E,F). The claw consists of three parts in its cross section: (1)a dense layer of the exocuticle (gray), (2) a loosely packed endocuticle, and the claw lumen. Semicircular (A′) and rectangular (B′) parts of the claw were calculated separately. XA, XB and XT are bending centers of sections A, B and A+B, respectively. For an explanation of other symbols, see text, Table 3 and Appendix B.

Claw geometry. (A) Claw shape of the hind-, mid- and forelegs (drawings are based on SEM micrographs). (B) Five arcs used for quantitative description of the claw geometry (see Table 2). (C) Cross-section model of the claw for stress calculation. The model is based on SEM data (see Fig. 3E,F). The claw consists of three parts in its cross section: (1)a dense layer of the exocuticle (gray), (2) a loosely packed endocuticle, and the claw lumen. Semicircular (A′) and rectangular (B′) parts of the claw were calculated separately. XA, XB and XT are bending centers of sections A, B and A+B, respectively. For an explanation of other symbols, see text, Table 3 and Appendix B.


Geochemical analysis

Stable isotopes

Methods: Pedogenic calcium carbonate (calcite) nodules both from in situ (unit 3) and ex situ (unit 4) sources were analyzed for stable carbon (δ 13 CCO3) and oxygen (δ 18 OCO3) isotopes in order to characterize the site’s paleohydrology and determine if the unit 4 pedogenic nodules were locally derived from unit 3 or related strata. Pedogenic nodules from unit 3 consisted of both micrite and fracture-fill calcite spar components that were analyzed separately and together in bulk. Nodules from unit 4 were homogeneous micrite and were analyzed in bulk. All carbonate samples were digested in phosphoric acid at room temperature and analyzed via a gas bench II attached to a Thermo Advance Plus isotope ratio mass spectrometer (IRMS) at the University of Arkansas Stable Isotope Laboratory (UASIL).

Shells of five different turtle taxa randomly sampled from units 4–5 (a large trionychid = Aspideretoides, a small trionychid = Gilmoremys gettyspherensis, a medium-sized baenid = cf. Boremys), a large baenid = Neurankylus utahensis and the giant panchelonioid) were also analyzed for δ 18 Op to determine their ecological (aquatic) fingerprints specifically to try and falsify our assumption they were all derived locally from the same facies/strata. Silver phosphate samples from turtle specimens were reacted with glassy carbon chips at 1400 °C on a thermo high temperature conversion elemental analyzer (TC/EA) attached to a continuous flow Advance Plus IRMS at UASIL.

Isotopic composition of water was calculated from turtle remains using the equation of Barrick, Fischer & Showers (1999) revised by Coulson et al. (2008):

δ 18 Ow = 1.08δ 18 Op – 22.3, while isotopic composition of water derived from carbonate nodules was based on temperature estimates using clumped isotopes (Burgener et al., 2019) and plant Leaf Margin Analysis (LMA) (Miller et al., 2013). We used the temperature dependent isotopic fractionation factor of O’Neil, Clayton & Mayeda (1969), to estimate isotopic composition of meteoric water:

1000 lna = 3.23(10 6 /T 2 ) − 3.5

δ 18 Ow = (δ 18 Oc + 1000)/a α − 1000

All carbonate values are reported as ‰VPDB, while all phosphate values are ‰VSMOW.

Pedogenic Carbonate.—Results are given in Table 1, Table S2 and Fig. 11A. Bulk (spar + micrite) mean δ 13 C, for the unit 3 calcite nodules is −9.26‰ (n = 14, SD = 0.76). The mean δ 13 C for the micritic component of the nodules is −9.06‰ (n = 8, SD = 0.28) while the mean for the sparry calcite is −9.54‰ (n = 6, SD = 0.59). Bulk (spar + micrite) mean δ 18 OCO3 for unit 3 calcite nodules is −7.37‰ (n = 14, SD = 0.70). The mean δ 18 O for the micritic component is −7.44‰ (n = 8, SD = 0.53) while that for the sparry calcite is −7.27‰ (n = 6, SD = 0.88). For conglomerate-derived micrite nodules (unit 4), mean δ 13 C is −7.43‰ (n = 2, SD = 0.47), and mean δ 18 O is −7.73‰ (n = 2, SD = 0.16).

Nodule type Unit n δ 18 OCO3 VPDB 1σ δ 18 OCO3 δ 13 CCO3 1σ δ 13 CCO3
Bulk (micrtic + sparry mixed) 3 14 −7.37 0.7 −9.26 0.76
Micritic 3 8 −7.44 0.53 −9.06 0.28
Sparry 3 6 −7.27 0.88 −9.54 0.59
Micritic (conglomerate) 4 2 −7.73 0.16 −7.43 0.47

Figure 11: Plots of stable isotope data.

Turtles.— A summary of data is shown in Table 2 and Fig. 11B. The δ 18 OP for the turtles ranges between 12.8 ± 1.3 (1σ) for the giant panchelonioid to 11.0 ± 1.4‰ for an unnamed baenid. Neurankylus utahensis averages δ 18 OP = 11.3 ± 2.4‰, while Aspideretoides δ 18 OP = 11.3 ± 0.7‰, and Gilmoremys gettyspherensis δ 18 OP = 11.1 ± 2.3‰. The four non-panchelonioid taxa produced δ 18 OP values statistically indistinguishable from each other, while the panchelonioid is slightly heavier (Fig. 11B).

Taxon n Median δ 18 OP VSMOW Avg. δ 18 OP VSMOW 1σ δ 18 OP
Aspideretoides 8 11.4 11.3 0.7
Gilmoremys 4 11.2 11.1 2.3
Baenid 4 10.8 11.0 1.4
Neurankylus utahensis 3 11.2 11.3 2.4
Large panchelonioid 23 12.7 12.8 1.3

Water Isotopic Composition.—Water isotopic composition was calculated from both turtles and pedogenic carbonate nodules (Table 3) to determine if the meteoric water source for the different carbonate nodules in units 3 and 4 was the same and to compare them to water that turtles consumed and lived in. For carbonate nodules, temperature must be assumed, and we used temperature averages derived from both clumped isotope analysis of Burgener et al. (2019) (35 ± 4 °C), and LMA of Miller et al. (2013) (20 ± 1 °C). Given the lack of significant difference between the different calcite phases of unit 3 carbonate nodules we use a bulk δ 18 Oco3 value of −7.37‰. With these average values, the isotopic composition of water that precipitated unit 3 nodules at 35 °C was −6.40‰ and the isotopic composition of water that precipitated nodules re-worked into unit 4 was −6.76‰. Using 20 °C, water isotopic composition was −9.83‰ for bulk unit 3 nodules and −10.18‰ for unit 4 transported nodules. Given the low variability of different turtle shell δ 18 Op we averaged all turtle δ 18 Op values. The δ 18 Ow that turtles lived in was −11.21‰.

Mineral Unit Temp α δ 18 Ow
VSMOW
Bulk unit 3 3 35 1.029901 −6.40
Bulk unit 3 3 20 1.03347 −9.83
Unit 4 4 35 1.029901 −6.76
Unit 4 4 20 1.03347 −10.19
Average large panchelonioid 5 −9.4
Average turtles 5 −11.2

Results are based on MAT data from Miller et al. (2013) and clumped isotope work of Burgener et al. (2019).

Discussion: The independent analysis of spar and micrite components in unit 3 nodules shows that they formed under different conditions, almost certainly at different times. The tight cluster of micrite values for unit 3 nodules, which we interpret as the primary Cretaceous age pedogenic calcite, indicates they all formed from waters with nearly identical composition. Much more widely scattered results from the sparry fraction suggest remobilization of the micritic calcite and random mixing with a slightly heavier fraction during later diagenesis or possibly even Holocene weathering. The oxygen isotopic values from unit 4 nodules, which do not contain sparry fill, are nearly identical to those of unit 3 micrite, indicating they are all from the same pedogenic process and meteoric water sources. However, the lighter C-isotopic values of the micrite component of unit 3 nodules argue they formed lower in the soil profile with a greater proportion of plant root respired CO2 and restricted input from atmospheric CO2 than unit 4 a finding consistent with our interpretation that most lag material in the unit 4 bonebed was locally derived from a slightly higher stratigraphic horizon than unit 3 (Fig. 12).

Figure 12: Pre-erosional soil profile showing hypothetical δ 18 O and δ 13 C isotopic patterns for unit 3 and overlying strata (now removed).

Turtle skeletal and carapace (phosphate) oxygen isotope composition (δ 18 Op) directly reflects their aqueous (δ 18 Ow) habitat (e.g., small pond vs. large stream) (Barrick, Fischer & Showers, 1999 Coulson et al., 2008). Given the calculated water isotope values from the RUQ turtles (Fig. 11B), excepting the giant panchelonioid, it is likely they all inhabited water with a similar isotopic composition during phosphate precipitation and therefore represent individuals derived from the same spatial/environmental/sedimentary context. The giant panchelonioid is statistically different being approximately 1.6‰ more enriched in 18 O (calculated δ 18 Ow average-9.4‰) relative to the other turtles, suggesting it lived in isotopically enriched ponded water, spent more time out of water subjecting itself to evaporative conditions, or spent time closer to the coast. Alternatively, the giant panchelonioid could have had a unique physiology that enriched its body water relative to smaller (or younger) counterparts, a phenomenon observed in other giant taxa (e.g., sauropods and very large theropods) (Suarez et al., 2014). There is no evidence to suggest it routinely inhabited marine environments as it would present much higher δ 18 Op and calculated δ 18 Ow (Cretaceous marine water = 0 to −1.2‰) (Suarez, González & Ludvigson, 2011 Ufnar et al., 2002). Simple mass balance modeling suggests that if the giant panchelonioid did spend time in marine water, it would only have been for

10% of its entire growth history.

Rare earth elements

Methods: Approximately 0.1 g samples were taken from tyrannosaur dentine and one vertebra, fish scales and one vertebra, dentine from Deinosuchus, four turtle shells, a carbonate nodule from unit 4, and a carbonate nodule embedded within a tyrannosaur vertebra. These were crushed or drilled, dissolved in 0.28 mL of 15.7 M trace grade nitric acid and diluted to 10 mL using DDI water, then analyzed for cerium through lutetium (lanthanum was not analyzed) on a Thermo iCap q inductively-coupled mass spectrometer at the Trace Element and Radiogenic Isotope Lab at the University of Arkansas. All samples were then normalized to the North American Shale Constant (NASC) using data in Gromet et al. (1984).

Results: Data is summarized in Table S3. Total rare earth element (REE) abundance ranged between 78.2 and 4,662.1 ppm for bone/dentine. The two carbonate nodules analyzed (one from within a bone cavity and one from unit 4) ranged between 33.6 and 214.4 ppm. NASC-normalized data show values for all taxa and the carbonate nodules are light REE (LREE) and middle REE (MREE)-enriched and heavy REE (HREE)-depleted. Fish and tyrannosaur material have NASC-normalized patterns that are indistinguishable from each other. Three of the four turtle elements have similar REE patterns to the tyrannosaur and fish material but are lower in overall REE concentration. Finally, the carbonate nodules’ REE patterns are also HREE-depleted.

Discussion: Rare earth elements (La 139 to Lu-175) are incorporated in fossil bones and teeth very early in their diagenetic history (Trueman & Tuross, 2002 Kohn & Moses, 2013 Suarez & Kohn, 2020) via a diffusion reaction process as fossilizing fluids (typically groundwater) trigger apatite recrystallization and replacement of (mostly) calcium cations in the crystal lattice (Millard & Hedges, 1996 Suarez & Kohn, 2020). The REE pattern, that is the relative proportions and abundance of LREEs, MREEs and HREEs are controlled by Redox, pH, dissolved colloids and ligands, and source rock and reflect the early diagenetic history of the geographic setting (Trueman, 1999). Thus, bones sharing the same fossilization history have closely similar REE patterns even though their absolute abundances differ. The similarity of REE patterns and ratios for all taxa analyzed (Fig. 13) demonstrate they share the same fossilization history, while similarities of the bone REE pattern to the carbonate nodule REE pattern (Fig. 13) indicate the overall signature was imparted during pedogenic carbonate formation (Fig. 12) i.e., during pedogenesis (Grandstaff & Terry, 2009 Metzger, Terry & Grandstaff, 2004 Suarez et al., 2007). This is corroborated by the HREE-depleted signature of RUQ fossils, which is typical of high suspension load or colloid-rich fluvial environments like floodplains (Herwartz et al., 2013 Rousseau et al., 2015), indicating REE infusion of fossils and peds was essentially complete prior to incorporation into the channel setting of units 4 and 5, which would have enriched the bones with HREEs (Trueman, 1999). The overall lower abundance of REE in the bones also suggests very little diagenetic alteration and rapid fossilization (Ullmann et al., 2020).

Figure 13: NASC-normalized REE patterns and ratios for RUQ fossils and carbonate.


Contents

Originally engineered before the Great War by the U.S. government as a cheap replacement for human troops during combat operations, deathclaws were derived from a mixed animal stock, primarily the popular Jackson's Chameleon. Although the project was successful in creating a ferocious predator capable of surviving on its own in the wild, no references exist of them ever being deployed on the battlefield. After the Great War, deathclaws escaped into the wild and quickly spread across the continent. The Enclave had no hand in the deathclaw's creation, so far as the Appalachian Enclave's research division knows, but they do note that the genetic manipulation looks similar to their own projects. ΐ] Eventually, they were refined by the Master through genetic manipulation and the Forced Evolutionary Virus. [Non-game 1] Because initial reports were limited to a series of isolated nests, deathclaws were viewed as legendary creatures by the various inhabitants of southern California. Α] However, the population in the Boneyard was keenly aware of their existence, as a single den mother and her offspring claimed the area between Downey and Norwalk around 2161, keeping the Gun Runners in a checkmate while terrorizing other communities in the region. Β] Also, a single deathclaw was found living near the outskirts of the Hub. Γ]

Their gradual spread throughout the wasteland raised awareness of their existence, until they entered common consciousness as a lethal predator. As stated above, the Enclave eventually continued their research project started before the war, developing intelligent deathclaws for use in hostile environments around 2235. [Non-game 2] On May 17, 2242, the first successful pack was dropped into Vault 13 to cloak the presence of the Enclave and their abduction of the dwellers within. Following their first combat test, the deathclaws broke free of their Enclave masters, becoming far more intelligent than anyone could foresee. [Non-game 3] They began developing a unique culture, as the first known non-humanoid sentient beings in history. Δ] However, their intelligence was discovered by Dr. Schreber of Navarro, whose report led to the extermination of intelligent deathclaws with extreme prejudice. Ε] Subsequent experimentation involved the aforementioned domestication units, although by the end of the 23 rd century. Ζ]


Scientists unearth first baby tyrannosaur fossils ever found

Reading Time: 3 minutes The fossilized lower jawbone of a Daspletosaurus horneri, one of the first baby tyrannosaurs ever discovered. Estimates based on this 75-million-year-old fossil suggest the dinosaur embryo measured about 71 centimetres long. (Photo: Greg Funston)

A University of Alberta student is part of a team of researchers who have just published an in-depth study of a stunning find: the first tyrannosaur embryo fossils ever discovered. The results shed new light on how the iconic dinosaurs grew and developed.

“Tyrannosaurs are represented by dozens of skeletons and thousands of isolated bones or partial skeletons,” said Mark Powers, second author on the study and PhD student in the Department of Biological Sciences. “But despite this wealth of data for tyrannosaur biology, the smallest identifiable individuals are aged three to four years old, much larger than when they would have hatched. No tyrannosaur eggs or embryos have been found even after 150 years of searching – until now.”

The study, led by U of A graduate Greg Funston, focused on two fossils of interest: a small toe claw of an Albertosaurus sarcophagus found near Morrin, Alta., and a small lower jawbone of a Daspletosaurus horneri found in Montana. The claw is roughly 71.5 million years old and the jawbone roughly 75 million years old.

“The discovery of embryonic material is a huge find in our efforts to understand how some of the most popular and charismatic dinosaurs began their life, and grew to immense sizes,” said Powers, who completed the research as a master’s student under the supervision of Philip Currie. “It provides a much-needed – and until now, missing – data point depicting the starting point for tyrannosaur growth.”

Powers said the unprecedented finds have a lot to teach researchers.

“There were two surprising results. The first is that the small tyrannosaur teeth were distinct from the teeth of older individuals – having not yet developed true serrations along the cutting edge of its teeth, as is iconic of the juveniles all the way through to adults,” he noted.

The second was the estimated size of the embryos. The specimen belonging to the toe claw was estimated to be about 110 cm long, while that of the jaw bone was about 71 cm.

“Amazingly, the size estimates match well with a hypothetical hatchling proposed by the late American-Canadian paleontologist Dale Russell back in 1970. This attests to Dale Russell’s insight into tyrannosaur development,” said Powers.

“This was an unexpected but surreal result, as the study was published in a special issue of the Canadian Journal of Earth Sciences honouring Dale Russell for his contributions to the field of paleontology.”

| By Andrew Lyle for © Troy Media

The views, opinions and positions expressed by columnists and contributors are the author’s alone. They do not inherently or expressly reflect the views, opinions and/or positions of our publication.

Scientists unearth first baby tyrannosaur fossils ever found added by University of Alberta on January 28, 2021
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Acknowledgements

We thank J. Moore, M. McDowell, L. Hatcher, G. Gully, A. Baynes, J. Shaw and P. McGuigan for help with fieldwork and A. Wood (Western Australian Department of Environment and Conservation) for the use of the “Wharncliffe Hilton”. We are grateful for the assistance of G. Johnston, G. Males, S. Washford, H. Howlett and W. Foster in the actualistic trials. Cheers to R. Correll for collecting the bark used in the opportunistic study. For assistance with the mock actualistic scratching trials we thank D. Stemmer and P. Blias (South Australian Museum) and M. Walters (Princess Margaret Hospital). We greatly appreciate loaning of surveying gear and help with surveying and GIS analysis provided by A. O’Flaherty and P. Connelley (TAFE SA). M. Siversson and H. Gore (Western Australian Museum) assisted with the bone modification study. The research was supported by an Australian Research Council Discovery Project grant (DP0772943) to G.J.P. This paper is dedicated to the memory of James Moore (1990–2014).


Watch the video: Tyrannosaurus Dinosaur Collection Jurassic World, T-Rex Heads, Dino Anatomy 티라노사우루스 공룡 (September 2022).


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