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Pterodon (mammal)

Pterodon is an extinct genus of hyaenodont in the family Hyainailouridae, containing two species. The type species Pterodon dasyuroides is known exclusively from the late Eocene to the earliest Oligocene of western Europe. The genus was first erected by the French zoologist Henri Marie Ducrotay de Blainville in 1839, who said that Georges Cuvier presented one of its fossils to a conference in 1828 but died before he could make a formal description of it. It was the second hyaenodont genus with taxonomic validity after Hyaenodon, but this resulted in taxonomic confusion over the validities of the two genera by other taxonomists. Although the taxonomic status of Pterodon was revised during the late 19th and early 20th centuries, it became a wastebasket taxon for other hyaenodont species found in Africa and Asia. Today, only the type species is recognized as belonging to the genus while just one is pending reassessment to another genus.

Taxonomy
Research history Early history In 1838, French zoologist Henri Marie Ducrotay de Blainville made a review of palaeontological history and taxonomy as built upon from previous decades. Blainville recognized the importance of dentition in determining the affinities of fossil animals but criticized the overreliance of dental systems as automatically indicating taxonomic affinities. He then went on to examine reported fossil "didelphids" (the taxonomic group now known as "marsupials") in European land. He conducted reviews of fossils as previously described by Laizer and Parieu, concluding that Hyaenodon is a valid genus based on its dentition but rejected the idea of it being a didelph based on its dental system and its molars being closer in affinity to modern carnivorans. Blainville also mentioned a fossil of an upper jaw that the late Georges Cuvier found and previously thought was close in affinity to the Tasmanian devil (Sarcophilus harrisii) of New Holland (Australia). Previously, Cuvier presented the fossil to the French Academy of Sciences in 1828 and thought that it was a large species of thylacine (Thylacinus cynocephalus) but died before he could make a formal description of it. Of note is that although the article of Hyaenodon by Blainville was first written and published in 1838, it was not until a year later in 1839 when it was republished that Blainville added a footnote to the paragraph about the upper jaw. In it, he said that he restudied that fossil from the Paris Basin and again was certain that it was a "monodelphian" (today placental) predator, which he named Pterodon dasyuroides. The genus name means "wing tooth" and is a combination of the Ancient Greek words "" (wing) and "" (tooth). The etymology of the type species name derives from the Australian marsupial genus Dasyurus and the Greek suffix "" meaning "like" due to apparent initial confusion of the genus affinity by Blainville. In 1841-1842, Blainville mentioned the genus Pterodon but replaced the previous species name with P. parisiensis. He also stated that despite thinking that the mammal did not have close affinities with Dasyurus, he did not have sufficient skull material to prove what mammal group it was closest to. The species name is in reference to Paris where its first fossil was found. In 1848-1852, however, he reclassified the species as Hyaenodon requieni (having recognized the validity of the genus) and listed P. dasyuroides as the only species of Pterodon. In 1853, Pomel changed his position by recognizing the validity of Hyaenodon, restoring the taxonomic affinities of species previously classified as belonging to it and therefore establishing that they no are no longer classified under Pterodon (H. leptorhynchus and H. brachyrhynchus). He also reclassified Taxotherium as a junior synonym of Hyaenodon instead of Pterodon. Within the genus Pterodon, he recognized the species name P. dasyuroides instead of P. parisiensis and created two additional species based on dentition shapes and sizes: P. cuvieri and P. coquandi. European hyaenodont revisions Gervais erected the species P. exiguum in 1873 based on dentition with some similarities to both Pterodon and Hyaenodon but noted that it may constitute a new genus once he has more fossil material. The palaeontologist then corrected himself in 1876 by stating that the species belongs to Hyaenodon as H. exiguum, not Pterodon. In 1876, Filhol recognized only P. dasyuroides among all species previously erected for the genus and named a new species P. biincisivus from the phosphorite deposits of Escamps, France. He, in addition to confirming the taxonomic validities of the two species, erected a third named P. quercyi in 1882. The French naturalist named a new species Oxyaena galliæ based on dental remains being apparently similar to that of species of Oxyaena previously described by Edward Drinker Cope in the Eocene deposits of New Mexico, United States. in The same year, Filhol also reported that an individual whose last name was Pradines recently discovered anterior portions of the skull of P. dasyuroides from the phosphate deposits of Limogne-en-Quercy. English naturalist Richard Lydekker made a review of known pan-carnivoran genera in 1884, classifying them within the order Carnivora and rejecting Cope's classification of the members into the suborder Creodonta within the order Bunotheria. While Cope originally assigned Hyaenodon as the sole member of Hyaenodontidae and Pterodon plus Oxyaena into Oxyaenidae, Lydekker felt that Pterodon was close in affinity to Hyaenodon and therefore belonged in Hyaenodontidae. For H. brachyhynchus, he listed P. brachyhynchus, P. requieni, and H. requieni as junior synonyms. He also listed P. leptorhynchus as a junior synonym of H. leptorhynchus as well as H. exiguus and P. exiguus as synonyms of H. vulpinus. For Pterodon, he recognized P. dasyuroides as the main valid species of the genus and listed P. parisiensis as a definite synonym as well as P. cuvieri and P. coquandi as possible synonyms, although he did not invalidate P. biincisivus. He also listed the species Oxyæna galliæ but thought that the genus could be merged into Pterodon due to minor dental differences and similarities to P. biincisivus. Swiss palaeontologist Ludwig Rütimeyer erected another species named P. magnus in 1891 based on the larger dentition sizes compared to typical species of Pterodon. However, in 1906, German scientist Rudolf Martin said that he wanted to synonymize P. magnum and the questionable P. quercyi with P. dasyuroides. He also listed P. biincisivus as a synonym of P. dasyuroides and stated that only one species is valid within Pterodon. Additionally, he revalidated the species O. galliæ but created the genus Paroxyaena for it, arguing that because oxyaenids are very weakly represented in Europe compared to North America and that therefore the species' similarities to oxyaenids may have an instance of convergent evolution. In 1979, Brigitte Lange-Badré made a systematic review of known hyaenodonts from Europe including Pterodon. She listed P. parisiensis, P. cuvieri, P. coquandi, P. biincisivus, and P. quercyi as synonyms of the only European species P. dasyuroides. In addition, she listed P. magnum as a synonym of Paroxyaena galliae and listed Hyaenodon exiguus as taking taxonomic priority over H. vulpinus, therefore making the latter name and P. exiguum synonyms. Many of the species classified or formerly classified to Pterodon were of African or Asian origins. Akhnatenavus leptognathus, "Hyainailouros" bugtiensis, Orienspterodon dahkoensis, and Neoparapterodon rechetovi. Several species names previously assigned to Pterodon were later considered to be synonyms of Hyaenodon species, namely "P. exploratus" (= H. incertus), "P. californicus" (= H. vetus), and "P. mongoliensis" (= H. mongoliensis). Additionally, Hemipsalodon was made a synonym of Pterodon by Robert Joseph Gay Savage in 1965 while Metapterodon was synonymized with it by Leigh M. Van Valen in 1967, but the synonymies were unsupported by later authors. As a result of the synonymies, only one species assigned to Pterodon remains pending reassessment to another genus: P. hyaenoides, which is classified as belonging to the Hyaenodontinae and is located in Asia. In 1999, A.V. Lavrov mentioned in his dissertation paper a genus he erected named "Epipterodon" for which "Pterodon" hyaenoides would have been reclassified to. However, as the genus name has not yet been referenced and taxonomically validated in any peer-reviewed source, its name currently remains invalid. Classification horridus'' of North America. The Hyaenodontidae was recently determined to have been a separate lineage from the Hyainailouridae. Pterodon has historically been classified undisputedly as at least being within the clade of hyaenodonts within the later 20th century, later being included within the subfamily Hyainailourinae. In 2016, Matthew Borths et al. suggested that the Hyainailouridae had a close relationship with the Teratodontinae and therefore arranged the superfamily Hyainailouroidea to include them and include the Proviverrinae, Hyaenodontidae, and North American hyaenodont groups. Compared to the proviverrines which never exceeded , the hyainailourines were much larger in size. This is because P. dasyuroides does not form a natural clade with non-European species classified within the genus, therefore meaning that they are pending reassessments to other genera. Pterodon sensu stricto (in a strict sense) made its appearance in western Europe by MP18 (late Eocene) in the form of P. dasyuroides and lasted up to MP20. P. dasyuroides was likely part of a ghost lineage of dispersing hyainailourines from Africa, and Kerberos did not appear to have descended into Pterodon or Parapterodon. }} == Description==
Description
Skull The order Hyaenodonta is diagnosed as having an elongated and narrow skull with a narrow cranial base (basicranium), a high and narrow occipital bone, and a transversely constricted interorbital region. Within the order, members of the Hyainailouridae share traits including a proportionally massive skull with an absence of any suture between the parietal bone and frontal bone, and a weak postorbital process (or projection), an extended pterygoid bone in its underside and side areas, the presence of a preglenoid crest, and side expansions of the squamosal relative to the back position of the zygomatic arch. and it is similar to that of Kerberos in being elongated and having a long rostrum and neurocranium. It is one of the two only known Eocene hyainailourine genera to be known by skull material, the other being Kerberos. The natural endocast is stored at the National Museum of Natural History, France with no catalogue number. It is estimated to have an endocast volume of . Only limited morphologies can be observed as that of the surface was not well-preserved. The cerebellum is bulky compared to the front portion of the cerebrum because the former's cavity is not as large as the cerebral fossa and lack of coverage by the neocortex. The cerebellum appears higher in position than the cerebrum, and the cerebellar vermis strongly projects between the cerebellum's two hemispheres. The primary fissure of the cerebellum, located on the paleocerebellum-neocerebellum boundary, is in a backwards position, more so than certain "condylarths" such as Arctocyon and Pleuraspidotherium. There is no other known transverse furrow on the cerebellar vermis. == Palaeobiology ==
Palaeobiology
The order Hyaenodonta occupied a wide range of body sizes/body masses and ranged from mesocarnivorous to hypercarnivorous diets. The Hyainailourinae, which includes P. dasyuroides, was one hyaenodont lineage that gained hypercarnivorous adaptations given its various specific dental configurations. Due to an overall lack of postcranial remains known for P. dasyuroides in the modern day, the locomotion of the Paleogene hyainailourid is unknown. In comparison, adequate postcranial remains are known for Hyainailouros sulzeri, Kerberos langebadreae, and Simbakubwa kutokaafrika, allowing for determinations of the locomotion methods of the hyainailourids. The elbow of H. sulzeri reveals incapability of flexible pronation-supination movements compared to typical cursorial mammals. The angulations and lengths of the fingers in relation to the metapodial bones, and the relation of the radius to the ulna suggest digitigrade movements of the forelimbs. The hindlimbs indicate similar results of digitigrade movement but, according to Ginsburg, have remnant traits of ancestral plantigrade movement. He theorized that it may have been semi-digitigrade overall with capabilities of leaping and occasional plantigrade movement. Similar traits of semi-digitigrady were also observed in S. kutokaafrika, with no capability of full digitigrade movement. Such adaptations of Miocene hyainailourines were likely the result of responses to more open environments. In comparison, however, the Paleogene hyainailourid Kerberos shows plantigrade stances and terrestrial locomotion based on known postcranial evidence. The locomotion method made it differ from the more cursorial hyaenodontid Hyaenodon as well as hyaenids and borophagine canids, which all also displayed degrees of ossiphageous (bone-crushing) adaptations. The locomotion method of Kerberos suggests that plantigrady was a primitive trait of the order Hyaenodonta, including the hyainailourids, while digitigrady/semi-digitigrady adaptations were derived traits within the order. Kerberos is thought to have been an active predator and opportunistic scavenger based on its fossil evidence. == Palaeoecology ==
Palaeoecology
Early pre–Grande Coupure Europe of Europe and Asia during the middle Eocene with possible artiodactyl and perissodactyl dispersal routes. For much of the Eocene, the world's environments were shaped by warm and humid climates, with subtropical to tropical closed forests being the dominant habitats. Multiple carnivorous mammal groups arose in Europe, Asia, Afro-Arabia, and North America, namely mesonychians, hyaenodonts, oxyaenids, and carnivoramorphs, dispersing between the continents. Land-based connections to the north of the developing Atlantic Ocean were interrupted around 53 Ma, meaning that North America and Greenland were no longer well-connected to western Europe. From the early Eocene up until the Grande Coupure extinction event (56 Ma to 33.9 Ma), the western Eurasian continent was separated into three landmasses, the former two of which were isolated by seaways: western Europe (an archipelago), Balkanatolia, and eastern Eurasia (Balkanatolia was in between the Paratethys Sea of the north and the Neotethys Ocean of the south). The Holarctic mammalian faunas of western Europe were therefore mostly isolated from other continents including Greenland, Africa, and eastern Eurasia, allowing for endemism to occur within western Europe. By MP16, a faunal turnover event occurred that marked the extinctions of lophiodonts, European tapiroids, and all crocodylomorphs except for the alligatoroid Diplocynodon. The causes of the faunal turnover have been attributed to a shift from humid and highly tropical environments to drier and more temperate forests with open areas and more abrasive vegetation. The surviving herbivorous faunas shifted their dentitions and dietary strategies accordingly to adapt to abrasive and seasonal vegetation. The environments were still subhumid and full of subtropical evergreen forests, however. The Palaeotheriidae was the sole remaining European perissodactyl group, and frugivorous-folivorous or purely folivorous artiodactyls became the dominant group in western Europe. During and after the faunal turnover event in western Europe, the Hyainailourinae made its first appearance in the continent by MP16 while the Hyaenodontinae appeared by MP17a and the carnivoran family Amphicyonidae by MP18. Hyaenodonts remained the dominant carnivorous mammal group compared to the carnivoraforms. Late Eocene P. dasyuroides first appears in the western European fossil record by MP18, with a significant ghost lineage of probably Afro-Arabian origins that makes its exact evolutionary history unknown. non-endemic artiodactyls (Dichobunidae, Tapirulidae, and Anthracotheriidae), perissodactyls (Palaeotheriidae), primates (Adapoidea and Omomyoidea), eulipotyphlans (Nyctitheriidae), chiropterans, and endemic rodents (Pseudosciuridae, Theridomyidae, and Gliridae). The alligatoroid Diplocynodon, present only in Europe since the upper Paleocene, coexisted with pre-Grande Coupure faunas as well, likely consuming insects, fish, frogs, and eggs due to prey partitioning previously with other crocodylomorphs that had since died out by the late Eocene. In addition to snakes, frogs, and salamandrids, rich assemblage of lizards are known in western Europe as well from MP16-MP20, representing the Iguanidae, Lacertidae, Gekkonidae, Agamidae, Scincidae, Helodermatidae, and Varanoidea, most of which were able to thrive in the warm temperatures of western Europe. The MP19 locality of Escamps, for instance, indicates that P. dasyuroides coexisted with the herpetotheriids Peratherium and Amphiperatherium, pseudorhyncocyonid Pseudorhyncocyon, bats (Hipposideros, Vaylatsia, Vespertiliavus, Stehlinia), primates (Microchoerus, Palaeolemur), rodents (Blainvillimys, Theridomys, Plesiarctomys, Glamys), hyaenodont Hyaenodon, amphicyonid Cynodictis, palaeotheres Palaeotherium and Plagiolophus, dichobunid Dichobune, anoplotheriids Anoplotherium and Diplobune, cainothere Paroxacron and Oxacron, xiphodonts (Xiphodon, Dichodon, Haplomeryx), and amphimerycid Amphimeryx. == Extinction ==
Extinction
in the Isle of Wight, from which Anoplotherium material has been collected. The stratigraphy of it and the Bouldnor Formation led to better understandings of faunal chronologies from the Late Eocene up to the Grande Coupure. The Grande Coupure extinction and faunal turnover event of western Europe, dating back to the earliest Oligocene (MP20-MP21), is one of the largest and most abrupt faunal events in the Cenozoic record, which is coincident with climate forcing events of cooler and more seasonal climates. The result of the event was a 60% extinction rate of western European mammalian lineages while Asian faunal immigrants replaced them. The Grande Coupure is often marked by palaeontologists as part of the Eocene-Oligocene boundary as a result at 33.9 Ma, although some estimate that the event began 33.6-33.4 Ma. The event correlates directly with or after the Eocene-Oligocene transition, an abrupt shift from a greenhouse world characterizing much of the Paleogene to a coolhouse/icehouse world of the early Oligocene onwards. The massive drop in temperatures stems from the first major expansion of the Antarctic ice sheets that caused drastic pCO2 decreases and an estimated drop of ~ in sea level. The seaway dynamics separating western Europe from other landmasses to strong extents but allowing for some levels of dispersals prior to the Grande Coupure are complicated and contentious, but many palaeontologists agreed that glaciation and the resulting drops in sea level played major roles in the drying of the seaways previously acting as major barriers to eastern migrants from Balkanatolia and western Europe. The Turgai Strait is often proposed as the main European seaway barrier prior to the Grande Coupure, but some researchers challenged this perception recently, arguing that it completely receded already 37 Ma, long before the Eocene-Oligocene transition. Alexis Licht et al suggested that the Grande Coupure could have possibly been synchronous with the Oi-1 glaciation (33.5 Ma), which records a decline in atmospheric CO2, boosting the Antarctic glaciation that already started by the Eocene-Oligocene transition. The Oi-1 glaciation, similar to the first glaciation event, caused large drops in sea level and pushed the global climate towards a coolhouse/icehouse environment. The Grande Coupure event also marked a large faunal turnover marking the arrivals of later anthracotheres, entelodonts, ruminants (Gelocidae, Lophiomerycidae), rhinocerotoids (Rhinocerotidae, Amynodontidae, Eggysodontidae), carnivorans (later Amphicyonidae, Amphicynodontidae, Nimravidae, and Ursidae), eastern Eurasian rodents (Eomyidae, Cricetidae, and Castoridae), and eulipotyphlans (Erinaceidae). MP20 also marks the last known appearance of Hyaenodon requieni while Hyaenodon gervaisi and Cynodictis extended beyond the extinction event. The extinction of Pterodon and survival of Hyaenodon by the Grande Coupure extinction event are notable but have no clear explanation available. The extinction causes have been attributed to climate deterioration (subsequent losses of suitable habitats and food), competition with dispersing carnivorans, or some combination of the two. However, carnivorans playing a role in its extinction has been questioned as there’s no strong evidence of carnivorans competitively displacing hyaenodonts. Additionally, the newcomers didn’t appear in Europe until the MP21, while hyainailourines were last record in MP19-20. ==References==
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