Castorocauda

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Castorocauda
Temporal range: Middle Jurassic, 164  Ma
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Castorocauda BW.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Therapsida
Clade: Cynodontia
Clade: Mammaliaformes
Order:† Docodonta
Family:† Docodontidae
Genus:† Castorocauda
Ji et al., 2006
Type species
†Castorocauda lutrasimilis
Ji et al., 2006

Castorocauda is an extinct, semi-aquatic, superficially otter-like genus of docodont mammaliaforms with one species, C. lutrasimilis. It is part of the Yanliao Biota, found in the Daohugou Beds of Inner Mongolia, China dating to the Middle to Late Jurassic. It was part of an explosive Middle Jurassic radiation of Mammaliaformes moving into diverse habitats and niches. Its discovery in 2006, along with the discovery of other unusual mammaliaforms, disproves the previous hypothesis of Mammaliaformes remaining evolutionarily stagnant until the extinction of the non-avian dinosaurs at the end of the Mesozoic.

Contents

Weighing an estimated 500–800 g (1.1–1.8 lb), Castorocauda is the largest known Jurassic mammaliaform. It is the earliest known mammaliaform with aquatic adaptations or a fur pelt. It was also adapted for digging, and its teeth are similar to those of seals and Eocene whales, collectively suggesting it behaved similarly to the modern-day platypus and river otters and ate primarily fish. It lived in a wet, seasonal, cool temperate environment – which possibly had an average temperature not exceeding 15 °C (59 °F) – alongside salamanders, pterosaurs, birdlike dinosaurs, and other mammaliaforms.

Discovery and etymology

The holotype specimen, JZMP04117, was discovered in the Daohugou Beds of the Jiulongshan Formation in the Inner Mongolia region of China, which dates to about 159–164 million years ago (mya) in the Middle to Late Jurassic. [1] [2] It comprises a partial skeleton including an incomplete skull but well-preserved lower jaws, most of the ribs, the limbs (save for the right hind leg), the pelvis and the tail. The remains are so well preserved that there are elements of its soft anatomy and hair. [1]

The genus name Castorocauda derives from Latin Castor "beaver" and cauda "tail", in reference to its presumed beaver-like tail. The species name lutrasimilis derives from Latin lutra "otter" and similis "similar", because some aspects of its teeth and vertebrae are similar to modern otters. [1]

Description

Castorocauda was the largest of known docodonts. [3] The preserved length from head to tail is 425 mm (16.7 in), but in life it was much larger. Based on the dimensions of the platypus, the lower weight limit was estimated to be 518 g (1.1 lb) in life, and the upper 700 to 800 g (1.5 to 1.8 lb), making it the largest known Jurassic mammaliaform, surpassing the previous record of 500 g (1.1 lb) for Sinoconodon . [1]

It had specialized teeth that curve backwards to help it hold onto slippery fish, as seen in modern seals and also ancestral whales. [4] The first two molars have cusps in a straight row, and interlocked during biting. This feature is similar to the ancestral condition in Mammaliaformes (such as in triconodonts) but is a derived character (it was specially evolved instead of inherited) in Castorocauda. [1] [5] The lower jaw contained 4 incisors, 1 canine, 5 premolars and 6 molars. [1]

The forelimbs of Castorocauda are very similar to those of the modern platypus: the humerus widens towards the elbow; the forearm bones have hypertrophied (large) epicondyles (where the joint attaches); the radial and ulnar joints are widely separated; the ulna has a massive olecranon (where it attaches to the elbow); the wrist bones are block-like; and the finger bones are robust. Docodontans were likely burrowing creatures and had a sprawling gait, and Castorocauda may have also used its arms for rowing, similar to the platypus. There are traces of soft tissue between the toes, suggesting webbed hind feet. [1] It likely also had claws, [6] and the holotype shows a spur on the hind ankle, which, in male platypuses, is venomous. [1]

Castorocauda likely had 14 thoracic, 7 lumbar, 3 sacral and 25 tail vertebrae. Like some mammals, it had plated ribs, and the ribs extended into the lumbar vertebrae. Plating occurred on the proximal margins (the part of the rib closest to the vertebra), and, in Castorocauda, they may have served to increase the insertion area (the part of a muscle which moves while contracting) of the iliocostalis muscle on the back, which would interlock nearby ribs and better support the torso of the animal. [1] Plated ribs are present in arboreal (tree-dwelling) and fossorial (burrowing) xenarthrans (sloths, anteaters, armadillos and relatives). The tail vertebrae are flattened dorsoventrally (shortened vertically and widened more horizontally); and each centrum has two pairs of transverse processes (which jut out diagonally from the centrum) on the headward side and another on the tailward side, making the centrum appear somewhat like the letter H from the top-view looking down. This tail anatomy is similar to beavers and otters, which use their tails for paddling and propulsion. [1] [5]

Fur was preserved on the holotype, and it is the earliest known pelt; [7] this showed that fur, with its many uses including heat retention and as a tactile sense, was an ancestral trait of mammals. Mammals preserved with fur from the Chinese Yixian Formation show little hair on the tail, whereas the fur outline preserved on the Castorocauda tail was 50% wider than the pelvis. The first quarter is covered by guard hairs, the middle half by scales and little hair cover and the last quarter by scales with some guard hair. Beavers have a very similar tail. [1] Evidence of fur and assumed heightened tactile senses indicate it had a well-developed neocortex, a portion of the brain unique to mammals which, among other things, controls sensory perception. [7]

Taxonomy

Stem Mammaliaformes
Castorocauda with other mammaliaforms [9]

Castorocauda is a member of the order Docodonta, an extinct group of mammaliaforms. Mammaliaformes includes mammal-like creatures and the crown mammals (all descendants, living or extinct, of the last common ancestor of all living mammals). Docodonts are not crown mammals. When Castorocauda was first described in 2006, it was thought to be most closely related to the European Krusatodon and Simpsonodon . [1] In a 2010 review of docodonts, Docodonta was split into Docodontidae, Simpsonodontidae and Tegotheriidae, with Castorocauda considered incertae sedis with indeterminate affinities. [10] Simpsonodontidae is now considered to be paraphyletic and thus invalid, and Castorocauda appears to have been most closely related to Dsungarodon , [8] [6] which came from the Junggar Basin of China and probably ate plants and soft invertebrates. [11]

Castorocauda is part of a Middle Jurassic mammaliaform diversification event, wherein mammaliaforms radiated into a wide array of niches and evolved several modern traits, such as more modern mammalian teeth and middle ear bones. [9] It was previously thought that mammals were small and ground-dwelling until the Cretaceous–Paleogene boundary (K–Pg boundary) when dinosaurs went extinct. The discovery of Castorocauda, [5] and evidence for an explosive diversification in the Middle Jurassic – such as the appearance of eutriconodontans, multituberculates, australosphenidans, metatherians and eutherians, among others – disproves this notion. This may have been caused by the breakup of Pangaea, which started in the Early to Middle Jurassic and diversified habitats and niches, or modern traits that had been slowly accumulating since mammaliaforms evolved until reaching a critical point which allowed for a massive expansion into different habitats. [9]

Paleoecology

Castorocauda is the earliest known aquatic mammaliaform, [12] pushing back the first appearance of mammaliaform aquatic adaptations by over 100 million years. [5] The teeth interlocked while biting, suggesting that they were strictly used for gripping; the recurved molars were likely used to hold slippery prey; and the teeth shapes are convergent with seals and Eocene whales, suggesting a similar ecological standing. Based on these, its adaptations to swimming and digging and its large size, Castorocauda was probably comparable to the modern day platypus, river otters and similar semi-aquatic mammals in ecology and fed primarily on fish (piscivory). [1]

Map of the Middle Jurassic MiddleJurassicMap.jpg
Map of the Middle Jurassic

The Daohugou Beds also include several salamanders, numerous pterosaur species (of which many likely were piscivorous), [2] several insects, the clam shrimp Euestheria [1] and some birdlike dinosaurs. No fish are known from specifically the Daohugou Beds, but the related Linglongta locality contains undetermined ptycholepiformes. Other mammals include the flying-squirrel-like Volaticotherium , the burrowing Pseudotribos , the oldest known eutherian Juramaia . [2] the rat-like Megaconus [13] and the gliding Arboroharamiya . [14] The plant life of the Tiaojishan Formation was dominated by cycadeoids (mainly Nilssonia and Ctenis ), leptosporangiate ferns and ginkgophytes and has pollen remains predominantly from pteridophytes and gymnosperms, which indicate a cool temperate and wet climate with distinct wet and dry seasons, [15] [16] possibly with an annual temperature of below 15 °C (59 °F). [16]

See also

Related Research Articles

<span class="mw-page-title-main">Docodonta</span> Extinct order of mammaliaforms

Docodonta is an order of extinct Mesozoic mammaliaforms. They were among the most common mammaliaforms of their time, persisting from the Middle Jurassic to the Early Cretaceous across the continent of Laurasia. They are distinguished from other early mammaliaforms by their relatively complex molar teeth. Docodont teeth have been described as "pseudotribosphenic": a cusp on the inner half of the upper molar grinds into a basin on the front half of the lower molar, like a mortar-and-pestle. This is a case of convergent evolution with the tribosphenic teeth of therian mammals. There is much uncertainty for how docodont teeth developed from their simpler ancestors. Their closest relatives may have been certain Triassic "symmetrodonts", namely Woutersia, and Delsatia. The shuotheriids, another group of Jurassic mammaliaforms, also shared some dental characteristics with docodonts. One study has suggested that shuotheriids are closely related to docodonts, though others consider shuotheriids to be true mammals, perhaps related to monotremes.

<span class="mw-page-title-main">Mammaliaformes</span> Clade of mammals and extinct relatives

Mammaliaformes is a clade of synapsid tetrapods that includes the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts during the Late Triassic. It is defined as the clade originating from the most recent common ancestor of Morganucodonta and the crown group mammals; the latter is the clade originating with the most recent common ancestor of extant Monotremata, Marsupialia and Placentalia. Besides Morganucodonta and the crown group mammals, Mammaliaformes also includes Docodonta and Hadrocodium.

Sinoconodon is an extinct genus of mammaliamorphs that appears in the fossil record of the Lufeng Formation of China in the Sinemurian stage of the Early Jurassic period, about 193 million years ago. While sharing many plesiomorphic traits with other non-mammaliaform cynodonts, it possessed a special, secondarily evolved jaw joint between the dentary and the squamosal bones, which in more derived taxa would replace the primitive tetrapod one between the articular and quadrate bones. The presence of a dentary-squamosal joint is a trait historically used to define mammals.

<i>Docodon</i> Extinct genus of mammaliaforms

Docodon is an extinct docodont mammaliaform from the Late Jurassic of western North America. It was the first docodont to be named.

<i>Volaticotherium</i> Extinct family of mammals

Volaticotherium antiquum is an extinct, gliding, insectivorous mammal that lived in Asia during the Jurassic period, around 164 mya. It is the only member of the genus Volaticotherium.

<span class="mw-page-title-main">Volaticotherini</span> Extinct clade of mammals

Volaticotherini is a clade of eutriconodont mammals from the Mesozoic. In addition to the type genus Volaticotherium, it includes the genera Argentoconodon, Ichthyoconodon, and potentially Triconolestes.

<span class="mw-page-title-main">Evolution of mammals</span> Derivation of mammals from a synapsid precursor, and the adaptive radiation of mammal species

The evolution of mammals has passed through many stages since the first appearance of their synapsid ancestors in the Pennsylvanian sub-period of the late Carboniferous period. By the mid-Triassic, there were many synapsid species that looked like mammals. The lineage leading to today's mammals split up in the Jurassic; synapsids from this period include Dryolestes, more closely related to extant placentals and marsupials than to monotremes, as well as Ambondro, more closely related to monotremes. Later on, the eutherian and metatherian lineages separated; the metatherians are the animals more closely related to the marsupials, while the eutherians are those more closely related to the placentals. Since Juramaia, the earliest known eutherian, lived 160 million years ago in the Jurassic, this divergence must have occurred in the same period.

<span class="mw-page-title-main">Haramiyida</span> Extinct order of mammaliaforms

Haramiyida is a possibly paraphyletic order of mammaliaform cynodonts or mammals of controversial taxonomic affinites. Their teeth, which are by far the most common remains, resemble those of the multituberculates. However, based on Haramiyavia, the jaw is less derived; and at the level of evolution of earlier basal mammals like Morganucodon and Kuehneotherium, with a groove for ear ossicles on the dentary. Some authors have placed them in a clade with Multituberculata dubbed Allotheria within Mammalia. Other studies have disputed this and suggested the Haramiyida were not crown mammals, but were part of an earlier offshoot of mammaliaformes instead. It is also disputed whether the Late Triassic species are closely related to the Jurassic and Cretaceous members belonging to Euharamiyida/Eleutherodontida, as some phylogenetic studies recover the two groups as unrelated, recovering the Triassic haramiyidians as non-mammalian cynodonts, while recovering the Euharamiyida as crown-group mammals closely related to multituberculates.

<span class="mw-page-title-main">Morganucodonta</span> Extinct order of mammaliaforms

Morganucodonta is an extinct order of basal Mammaliaformes, a group including crown-group mammals (Mammalia) and their close relatives. Their remains have been found in Southern Africa, Western Europe, North America, India and China. The morganucodontans were probably insectivorous and nocturnal, though like eutriconodonts some species attained large sizes and were carnivorous. Nocturnality is believed to have evolved in the earliest mammals in the Triassic as a specialisation that allowed them to exploit a safer, night-time niche, while most larger predators were likely to have been active during the day.

<span class="mw-page-title-main">Tiaojishan Formation</span> Geological formation in China

The Tiaojishan Formation is a geological formation in Hebei and Liaoning, People's Republic of China, dating to the middle-late Jurassic period. It is known for its exceptionally preserved fossils, including those of plants, insects and vertebrates. It is made up mainly of pyroclastic rock interspersed with basic volcanic and sedimentary rocks. Previously, the Tiaojishan Formation was grouped together with the underlying Haifanggou Formation as a single "Lanqi Formation." The Tiaojishan Formation forms a key part of the Yanliao Biota assemblage, alongside the Haifanggou Formation.

Paritatodon is an extinct mammaliaform which existed in Kyrgyzstan and England during the Jurassic period. It was originally the holotype specimen of Shuotherium kermacki, but Martin and Averianov (2010) argued that it resembled the genus Itatodon (Docodonta) and so renamed it Paritatodon.

<i>Megaconus</i> Extinct genus of mammaliaforms

Megaconus is an extinct genus of allotherian mammal from the Middle Jurassic Tiaojishan Formation of Inner Mongolia, China. The type and only species, Megaconus mammaliaformis was first described in the journal Nature in 2013. Megaconus is thought to have been a herbivore that lived on the ground, having a similar posture to modern-day armadillos and rock hyraxes. Megaconus was in its initial description found to be member of a group called Haramiyida. A phylogenetic analysis published along its description suggested that haramiyidans originated before the appearance of true mammals, but in contrast, the later description of the haramiyidan Arboroharamiya in the same issue of Nature indicated that haramyidans were true mammals. If haramiyidans are not mammals, Megaconus would be one of the most basal ("primitive") mammaliaforms to possess fur, and an indicator that fur evolved in the ancestors of mammals and not the mammals themselves. However, later studies cast doubt on the euharamiyidan intrepretation, instead finding it to be a basal allotherian mammal.

<i>Haldanodon</i> Extinct genus of mammaliaforms

Haldanodon is an extinct docodont mammaliaform which lived in the Upper Jurassic. Its fossil remains have been found in Portugal, in the well-known fossil locality of Guimarota, which is in the Alcobaça Formation. It may have been a semi-aquatic burrowing insectivore, similar in habits to desmans and the platypus. Several specimens are known, include a partial skeleton and well-preserved skulls.

<i>Shenshou</i> Extinct genus of mammaliaforms

Shenshou is a genus of haramiyidan dating from the Oxfordian stage of the Late Jurassic, approximately 160 million years ago. Fossils were recovered from the Tiaojishan Formation in the Liaoning province of China.

<i>Agilodocodon</i> Extinct genus of mammaliaforms

Agilodocodon was a genus of shrew-sized docodont from the Middle Jurassic, believed to be the earliest known tree-climbing mammaliaform. It contains one species, A. scansorius.

<i>Docofossor</i> Extinct genus of mammaliaforms

Docofossor is an extinct mammaliaform from the Jurassic period. Its remains have been recovered in China from 160 million years old rocks. It appears to have been the earliest-known subterranean mammaliaform, with adaptations remarkably similar to the modern Chrysochloridae, the golden moles.

<i>Ichthyoconodon</i> Extinct family of mammals

Ichthyoconodon is an extinct genus of eutriconodont mammal from the Lower Cretaceous of Morocco. It is notable for having been found in a unique marine location, and the shape of its teeth suggests an unusual, potentially fish-eating ecological niche. Analysis suggests it is part of a group of gliding mammals that includes Volaticotherium.

<i>Borealestes</i> Extinct genus of mammaliaforms

Borealestes is a genus of docodontan from the Middle Jurassic of Britain, first discovered on the Isle of Skye near the village of Elgol. It was the earliest mammaliaform from the Mesozoic found and named in Scotland. A second species and was later found in other Middle Jurassic sites in England, but is now shown to be a different genus. A new species, B. cuillinensis was named in 2021, also from Skye.

<i>Vilevolodon</i> Extinct genus of mammaliaforms

Vilevolodon is an extinct, monotypic genus of volant, arboreal euharamiyids from the Oxfordian age of the Late Jurassic of China. The type species is Vilevolodon diplomylos. The genus name Vilevolodon references its gliding capabilities, Vilevol, while don is a common suffix for mammalian taxon titles. The species name diplomylos refers to the dual mortar-and-pestle occlusion of upper and lower molars observed in the holotype; diplo, mylos.

Dobunnodon its an extinct genus of docodont from the Middle Jurassic (Bathonian) Forest Marble Formation of England, first discovered in Oxfordshire near the village of Kirtlington. The type species, D. mussettae, was originally named as a species of Borealestes in 2003.

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