Instruct: Given a question, retrieve relevant passages that answer the question.
Query: When was evidence of the Catopsbaatar first found?
Catopsbaatar is a genus of multituberculate, an extinct order of rodent-like mammals. It lived in what is now Mongolia during the late Campanian age of the Late Cretaceous Period, about 72 million years ago. The first fossils were collected in the early 1970s, and the animal was named as a new species of the genus Djadochtatherium in 1974, D. catopsaloides. The specific name refers to the animal's similarity to the genus Catopsalis. The species was moved to the genus Catopsalis in 1979, and received its own genus (Catopsbaatar, Greek and Mongolian for "visible hero") in 1994. Five skulls, one molar, and one skeleton with a skull are known; the latter is the genus' most complete specimen. Catopsbaatar was a member of the family Djadochtatheriidae.
The intermediate zygomatic ridge on the squamosal bone (also for jaw-muscle attachment) was much smaller and lower than the anterior zygomatic ridge in front of it. Catopsbaatar differed from other djadochtatheriids in that the intermediate ridge contacted the anterior ridge with its front edge. The posterior zygomatic ridge at the lower back of the squamosal bone was the weakest of the three ridges, and only marked by a depression. The postorbital processes behind the orbit on the parietal bone were very long, and the parietal ridges extended from the hind margin of the postorbital processes towards each other (but without reaching). The nuchal crest was very prominent, and extended to the sides to form "wings". The occipital plate was almost vertical and slightly concave, and was obscured by the nuchal crest when the skull was viewed from above. Though incompletely preserved, it is assumed that the occipital plate sloped back and upwards from the occipital condyles (as in some relatives).[3] The orbito-temporal vascular system inside the skull of Catopsbaatar did not differ much from those of related genera.[15]
It is unclear exactly when horses were first ridden because early domestication did not create noticeable physical changes in the horse. However, there is strong circumstantial evidence that horse were ridden by people of the Botai culture during the copper age, circa 3600-3100 BCE. The earliest evidence suggesting horses were ridden dates to about 3500 BCE, where evidence from horse skulls found at site in Kazakhstan indicated that they had worn some type of bit. Wear facets of 3 mm or more were found on seven horse premolars in two sites, Botai and Kozhai 1, dated about 3500–3000 BCE. It is theorized that people herding animals first rode horses for this purpose, presumably bareback, and probably used soft materials such as rope or possibly bone to create rudimentary bridles and hackamores. However, the earliest definitive evidence of horses being ridden dates to art and textual evidence dating to about 2000-1500 BCE.
The only Catopsbaatar specimen which preserves the postcranial skeleton is PM120/107, which is fragmentary. It includes elements which are unknown, or incompletely preserved, in other multituberculates. One lumbar vertebra (the fifth or sixth, from between the rib cage and the pelvis) had a spinous process which was stout in side view and long when seen from above. The clavicle was slightly less curved than that of Kryptobaatar (resembling a bent rod which widened at each end), and measured about 24.8mm (0.98in). The upper part of the scapulocoracoid was relatively wide in side view, and the lower part was very narrow. It was probably about 60mm (2.4in) long when complete. The preserved part of the humerus (upper arm bone) was about 37.5mm (1.48in) long. Its shaft was triangular in cross-section, relatively narrow when seen from above, and most of its width was occupied by the intertubercular groove. This groove was delimited on the side by the crest of the greater tubercle, whose middle part formed the deltopectoral crest. The ulnar condyle, where the ulna of the lower arm articulated with the humerus, was more prominent than the radial condyle (where the radius articulated), oval, and delimited from the radial condyle by a groove. The radius was about 26mm (1.0in) long, with a prominent head. Its shaft was smooth, compressed from top to bottom, and oval in cross-section. The ulna was compressed sideways, flatter than the radius, and about 40mm (1.6in) long as preserved.[12]
Kielan-Jaworowska and American palaeontologist Robert E. Sloan considered the genus Djadochtatherium a junior synonym of Catopsalis, and created the new combination C. catopsaloides in 1979.[5] American palaeontologists Nancy B. Simmons and Miao Desui conducted a 1986 cladistic analysis which indicated that Catopsalis was a paraphyletic taxon (an unnatural grouping of species), and C. catopsaloides required its own generic name.[6] Kielan-Jaworowska followed Simmons and Miao's suggestion, moving C. catopsaloides to its own monotypic genus (Catopsbaatar) in 1994. Catops is derived from the Greek word katoptos ("visible" or "evident"); baatar is Mongolian for "hero", and the name refers to Catopsbaatar's similarity to the genus Catopsalis (as is the case for the specific name).[7] The name Catopsalis itself consists of the Greek words for "visible" and "cutting shears" (psalis).[8][7]
The pelvic bones of Catopsbaatar specimen PM120/107 may not have been fused because fusion occurred late in development, because it was a sexually dimorphic feature only occurring in males (unfused pelvic bones might have enabled expansion of the birth canal in females), or pelvic fusion may be a taxonomic difference between Catopsbaatar and other multituberculates.[12] Unlike with other mammals, the pelvis of multituberculates was very narrow; in other genera where the pelvis is known, each half of the pubis and ischium were fused together, forming a keel. The length and rigidity of the keel indicate that the pelvis could not have spread during birth. Because there would be little space for the passage of an egg (egg-laying monotremes have wide ischial archs), Kielan-Jaworowska suggested in 1979 that multituberculates were viviparous (gave live birth) and that the newborns were extremely small—similar to those of marsupials.[4]
Catopsbaatar had a single lower pair of incisors, characteristic of multituberculates, which was very strong and compressed sideways. It had a sharply limited band of enamel, and grew continually. The p3 premolar was very small, and adhered entirely to the lower diastema under the larger p4. The blade-like p4 was roughly trapezoidal in side view, and had three cusps along the horizontal upper margin and one cusp on the outer back side. The p4 did not have the ridges on the outer and inner side, as are present in other multituberculates. The m1 molar was almost symmetrical, and its cusp formula was 4:4, the size of the cusps decreasing towards the back. The m2 had a cusp formula of 2−3:2, most specimens being 2:2. The cusps on the inner side were wider than those on the outer side, the inner row of cusps was shorter than the outer one, and the hind margin of the tooth was arranged obliquely.[3]
The limb posture of multituberculates has been debated. According to some researchers, they employed a parasagittal stance with the erect limbs under the body; others consider a sprawling stance more likely. Kielan-Jaworowska and Hurum supported the latter theory in 2006 based on the presence of hind-leg spurs, a feature they considered present only in sprawling mammals. They pointed out that all early mammals preserved in lacustrine (lake) deposits were compressed from top to bottom, suggesting a sprawling stance, whereas later mammals were preserved on their flanks. Earlier arguments for a sprawling stance include deep pelvises and features of the legs. They also suggested that the feet of multituberculates would have been plantigrade (the sole touching the ground) at rest, but digitigrade (the sole not touching the ground) when jumping and running quickly; they dismissed the idea that the forelimbs of multituberculates and other early mammals were more parasagittal than their hindlimbs. Kielan-Jaworowska and Hurum depicted Catopsbaatar with plantigrade, sprawling legs, with mobile spurs which pointed inward when preparing for attack.[18][4]
The most complete adult Catopsbaatar skull (specimen PM 120/107) is 63mm (2.5in) long and 55mm (2.2in) wide, with a 41mm-long (1.6in) lower jaw. By comparison, the skull of the juvenile holotype (ZPAL MgM−I/78) is about 53mm (2.1 inches) long and 56mm (2.2 inches) wide, with a 35mm-long (1.4in) lower jaw. The largest adult skull (ZPAL MgM−I/79) is 70mm (2.8in) long but, since it is incomplete, its other measurements are unknown. Catopsbaatar was larger than its relatives, Kryptobaatar and Djadochtatherium.[3] Multituberculates had relatively large skulls and short necks; their skulls were proportionally longer and wider than those of similarly sized rodents and marsupials.[4] The external appearance of their heads may have been similar to those of rodents.[11]
Hurum, Zhe-Xi Luo, and Kielan-Jaworowska suggested in 2006 that the spurs on the ankles of Mesozoic mammals (such as Catopsbaatar) were homologous with those of monotremes, and were a basal (or "primitive") feature lost by later therian mammals. The male platypus uses the spur to deliver venom from a gland, but it is unknown if the extinct groups were venomous as well. Mesozoic mammals were mostly small (with exceptions such as the fox-sized Repenomamus) and, although they were too small to be prey for large theropod dinosaurs, smaller theropods, large lizards, crocodiles, and birds could have fed on them. For example, mammal jaws have been found in the abdomen of a specimen of the small theropod Sinosauropteryx; the jaws belonged to Zhangheotherium, which also had spurs, and the multituberculate Sinobaatar. Since dinosaurs dominated Earth during the Mesozoic, this period has been called the "dark ages" of mammalian history. The spur, which would have been more effective if venomous, was probably used as a defensive weapon by small, early mammals. It could also have been used during intraspecific competition or predation.[16]
Evidence for the manufacture of Deritend ware in Birmingham was first discovered during the widening of High Street, Deritend in 1953, when misfired fragments of pottery were found to the south of the main road. Further evidence has since been found at the Old Crown Inn, the Custard Factory and underneath the site of Selfridges in the Bull Ring.
In 2008, Kielan-Jaworowska and Hurum suggested that the long spinous process on a Catopsbaatar vertebra and the long transverse processes in Nemegtbaatar may indicate that some multituberculates were saltatorial (had the ability to jump). Catopsbaatar probably had strong muscles attaching to the tuber calcanei, which further supports the jumping hypothesis.[12][19] Although it has been suggested that multituberculates were arboreal (lived in trees), most Asian taxa were probably terrestrial; some others were fossorial, digging and living underground.[4]
Red cedar was used extensively wherever it was found along the northwest coast (British Columbia, Washington state, Parts of Alaska). Evidence of this use is found in CMTs (Culturally Modified Trees) that are found throughout the coast. When First Nations people removed the bark from cedars, it left a scar - which is considered a CMT. Other types of harvest (for planks, tinder, and other uses) leave different types of evidence of cultural modification.
All specimens of Catopsbaatar are known from the Red Beds of Hermiin Tsav formation, except for one molar from the Barun Goyot Formation. The formations are considered coeval (of the same geological age), contain many of the same animals, and probably date to the late Campanian age of the Late Cretaceous Period (about 72 million years ago).[20] The rock facies of the Red Beds of Hermiin Tsav formation consist of orange-coloured, thick-bedded sandstone, with a thin interbedding of light-coloured silt stones and claystones.[21][3] The rock facies of the Barun Goyot Formation are considered to be the result of an arid or semi-arid environment, with aeolian (deposited by wind) beds.[22][23]
The dental formula (the number of teeth of each type in the tooth row of a mammal) of Catopsbaatar was 2.0.3.21.0.2.2 (two incisors, no canines, three premolars and two molars in half of the upper tooth row, and one incisor, no canines, two premolars and two molars in half of the lower). By comparison, the dental formula of humans is 2.1.2.2–32.1.2.2–3. Each tooth in a mammal is designated with a letter and number by position (I for incisor, C for canine, P for premolar, M for molar); the letters are capitalised for the teeth of the upper jaw, but not for those in the lower jaw. The cusp formula shows the arrangement and number of cusps in consecutive rows of a tooth, from the outer to the inner side; each row is separated by a colon.[3][4][10]
Catopsbaatar belonged to the order Multituberculata, a group within Allotheria (an infraclass of mammals outside Theria, the group that contains modern placentals and marsupials). Multituberculates are characterized by having premolars and molars with multiple low cusps, arranged in longitudal rows. Multituberculates are the best-known group of mammals from the Mesozoic Era, when the dinosaurs dominated; although the earliest multituberculate remains are from the Jurassic Period, the group is known as recently as the Eocene Epoch (thereby surviving the Cretaceous–Palaeogene extinction event). The group may have become extinct due to competition with eutherian mammals, such as rodents. Multituberculates were mainly known from teeth and jaws until the 1920s, when more complete specimens were discovered—first in Asia, and then elsewhere.[4] Postcranial bones (the rest of the skeleton, other than the skull) remain rare.[12]
According to Gambaryan and Kielan-Jaworowska, the adaptation for crushing hard seeds sometimes—as in Catopsbaatar—opposed the benefit of a low condylar process (which discourages mandibular dislocation). The anterior and intermediate zygomatic ridges of the skull were the origin of the superficial masseter muscle, which facilitates chewing. The separation of the origin of this muscle into two parts and the rounded muscle scars left by them are unique among mammals to multituberculates. The masticatory muscles of multituberculates independently evolved features shared with rodents and small herbivorous marsupials. As with rodents, multituberculates may have been capable of bilateral mastication—where both rows of teeth in the mandible perform the same function simultaneously—and unilateral mastication (where the rows on one side are used).[3][11]
The maxilla (the main part of the upper jaw) was extensive, and formed most of the side of the snout. It contained all the upper teeth, except the incisors. The infraorbital foramina (openings at the lower front of the maxilla) were slit-like in some specimens and rounded in others, and varied in number from one to three. One of the most characteristic features of the face of Catopsbaatar was the very large anterior zygomatic ridge on the sides of the upper jaw (a site for jaw-muscle attachment). It was much higher than in other djadochtatheriids except for Djadochtatherium, from which it differed in that the ridge was semicircular rather than roughly trapezoidal (other genera have elliptical ridges). The front margin of the anterior zygomatic ridge was thickened, which produced a bulge on the side margin of the snout when viewed from the side and above. The lower part of the suture between the maxilla and the squamosal bone extended along the hind border of the anterior zygomatic ridge. The palatal processes of the maxilla formed most of the palate. The major palatine foramina had shallow grooves extending forward. The postpalatine torus (a bony protrusion on the palate) of Catopsbaatar was less prominent than was that of Tombaatar. The frontal bone was large, and formed most of the cranial roof. The suture between the frontal and parietal bones formed a U-shaped part in the middle which pointed backwards—similar to the condition in Kryptobaatar but less deep, with smaller U-shaped structures at the sides. This differed from Djadochtatherium, which had a narrower V-shaped suture between the frontal and parietal bones.[3]
Evidence of Roman religious beliefs in York have been found including altars to Mars, Hercules, Jupiter and Fortune, while phallic amulets are the most commonly found type of good luck charm. In terms of number of reference the most popular deities were the spiritual representation ("genius") of York and the Mother Goddess; there is also evidence of local or regional deities. There was also a Christian community in York although it is not known when it was first formed and there is virtually no archaeological record of it. The first evidence of this community is a document noting the attendance of Bishop Eborius of Eboracum at the Council of Arles (314), and bishops also attended the First Council of Nicaea in 325, the Council of Sardica, and the Council of Ariminum.
The skull of Catopsbaatar was up to 70mm (2.8in) long and, as in other multituberculates, proportionally large. The external appearance of their heads may have been similar to those of rodents. The skull was heavy-set and wide, with the zygomatic arches strongly expanded to the sides. The eye sockets were smaller and placed further back than in its relatives, and the snout was more elongated. Catopsbaatar had semicircular ridges on the side of the skull, to which the jaw muscles were attached. The mandible was strong and very elongated. It had very robust incisors, and cheek teeth with multiple cusps (for which multituberculates are named). The pelvic bones differed from those of other multituberculates in that they were not fused to each other. Catopsbaatar had spurs on its ankles, like those of the male platypus and echidna, without evidence of a venom canal (present in the former).
The following cladogram shows the placement of Catopsbaatar among other multituberculates according to Kielan-Jaworowska and Hurum, 1997:[13]
Although multituberculates were thought to have been carnivores or herbivores, since American palaeontologist William A. Clemens and Kielan-Jaworowska suggested modern rat kangaroos as analogues for the group in 1979 they have been considered omnivores (feeding on both plants and animals).[4] Uniquely among mammals, multituberculates employed a backward chewing stroke which resulted in the masticatory muscles—the muscles which move the mandible—being inserted more to the front than in other groups (including rodents). Gambaryan and Kielan-Jaworowska reconstructed the masticatory musculature of various multituberculates in 1995, and found that Catopsbaatar and its relatives had very powerful masticatory musculature, due to their high zygomatic arches and large anterior and intermediate zygomatic ridges and coronoid processes. Their powerful incisors, with limited bands of enamel, would have been well adapted to gnawing and to cutting hard seeds (similar to rodents). Since it was larger than some other multituberculates, Catopsbaatar would only have to open its mouth 25 degrees to crush hard seeds 12–14mm (0.47–0.55in) in diameter; a 40-degree gape would have caused dislocation. After the incisors cut, the premolars and molars would begin to grind with a "power stroke".[11]
The spurs of Catopsbaatar and other Mesozoic mammals may have been used for protection against theropod dinosaurs and other predators. Multituberculates are thought to have given live birth, and the fact that they had hair indicates that they were homeothermic ("warmblooded"). Multituberculates would have been omnivorous; Catopsbaatar had powerful jaw muscles, and its incisors were well adapted for gnawing hard seeds, using a backwards chewing stroke. Multituberculates are thought to have had a sprawling posture, and Catopsbaatar may have been able to jump. Catopsbaatar is known from the Red Beds of Hermiin Tsav and the Barun Goyot Formation in Mongolia, which are thought to be the same geologic age (about 72 million years old).
Unlike most other multituberculates and other mammals, the calcaneus bone at the back of the foot had a short tuber calcanei (like some tree kangaroos), with an expanded, anvil-shaped proximal process strongly bent downwards and to the side. Catopsbaatar had an os calcaris bone on the inner side of its ankle, a feature also seen in modern male monotremes (the platypus and the echidna) and other Mesozoic mammals. The os calcaris bone was plate-like and rectangular in outline; as in monotremes, it supported the cornu calcaris in forming a spur on the outer side of the tarsus (cluster of foot bones). Unlike with other Mesozoic mammals, these two elements were not fused together in multituberculates. The cornu calcaris was triangular, with a concavity in the middle, and was 13mm (0.51in) long. The spur was flattened, and was thicker at its conjunction with the os calcaris (where they connected via several ridges). As the spur of PM120/107 may have been moved from its original position, it is unknown whether it faced inwards (like in the platypus). Unlike in the platypus, there was no impression of a canal for venom. The cornu calcaris of Catopsbaatar was ossified (turned into bone) and would have been covered in keratin (the horny covering seen in nails and hoofs). The cornu calcaris of the platypus consists only of keratin, and is hollow.[12][16]
Being a cimolodont, Catopsbaatar did not have I1 incisors.[13] The I2 upper-front incisors of Catopsbaatar were very robust and had a sharply limited band of enamel. The two incisors converged slightly towards the middle, touching each other. The smaller I3 incisor behind was cone-shaped. The alveolus (tooth socket) of Catopsbaatar's I3 incisor was formed by the premaxilla, rather than the premaxilla and maxilla (unlike in Tombaatar). The front upper premolars P1 and P3 were only present in juveniles (deciduous), disappearing (with their alveoli) in older individuals. P1 appears to have had two cusps, was single-rooted, and had a cone-like, blunt crown. P3 was single-rooted and smaller than P1. The cusp formula of the P4 premolar was 5−4:1, the central cusp being the largest. The P4 of Catopsbaatar was almost trapezoidal in shape (unlike in Djadochtatherium and Kryptobaatar, where it is crescent-shaped), smaller, and lacking ridges. Catopsbaatar also differed by only having three upper premolars, lacking the P2 (a feature shared with Tombaatar). Other mammals usually evolve the loss of teeth at the beginning or end of a tooth row, not in the middle (as in multituberculates). The cusp formula of the M1 molar was 5−6:5−6:4, with the inner ridge extending about 75 percent of the tooth's length. The cusps of M1 were sharp and unworn in juveniles, but worn and concave in older animals. The cusp formula of the M2 molar was 2:2−3:2−3.[3][4]