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=== Dentition === |
=== Dentition === |
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=== Postcranial skeleton === |
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=== Footprints === |
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== Palaeobiology == |
== Palaeobiology == |
Revision as of 18:25, 5 July 2024
PrimalMustelid/sandbox Temporal range:
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P. magnum skeleton | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Perissodactyla |
Family: | †Palaeotheriidae |
Genus: | †Palaeotherium Cuvier, 1804 |
Type species | |
†Palaeotherium magnum Cuvier, 1804
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Other species | |
For subspecies suggested, see below. | |
Synonyms | |
Genus synonymy
Synonyms of P. magnum
Synonyms of P. medium
Synonyms of P. crassum
Synonyms of P. curtum
Synonyms of P. duvali
Synonyms of P. muehlbergi
Dubious species
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Palaeotherium (Ancient Greek for 'old beast') is an extinct genus of perissodactyl ungulate known from the Mid Eocene to earliest Oligocene of Europe. First described by French naturalist Georges Cuvier in 1804, Palaeotherium was among the first Paleogene mammals to be described.
Research history
First descriptions
In 1782, the French naturalist Robert de Lamanon described a fossil skull retaining the upper and lower jaws that was collected from the quarries of Montmartre, a large hill near Paris, France, that belonged to the nobleman Philippe-Laurent de Joubert. He recognized that the morphologies of its molars and incisors were roughly akin to those of ruminants but noted that its dentition still lacked any modern analogues. As a result, he hypothesized that the animal was extinct, had amphibious behaviors, and fed on both herbs and fish.[1][2]
Since 1796, the French naturalist Georges Cuvier innovated the idea of vanished worlds of extinct animals, but as his observations of fossils were mostly limited to drawings and very fragmentary fossil materials stored at the National Museum of Natural History, France, his palaeontological insight remained limited early on. The fossils of Montmartre were credited with great importance to the field of palaeontology, as the fossil taxa found near Paris were embedded in deeper and harder sediments, falling between the Pleistocene-aged mammals and the Cretaceous-aged reptiles.[2] In 1798, he documented fossils from Montmartre, suggesting initially that they could have belonged to the canid genus Canis based on its dentition.[3] Not long after in the same year, he changed his mind and thought that the fossil mammal instead would have been within the order of pachyderms, theorizing that it would have been closest to tapirs and that it would have had trunks like them. He also figured out that the animals of Montmartre comprised of multiple different species with different sizes and numbers of toes.[4][2]
Early taxonomy and depictions
In 1804, Georges Cuvier described the sets of fossils from the gypsum quarries of the outskirts of Paris (known as the Paris Basin). Describing the skull previously reported by de Lamanon, he confirmed that it belonged to a mammal, had a complete set of 44 total teeth, and had molar morphologies similar to those of rhinoceroses and hyraxes. The naturalist, recognizing that its separate affinities from other mammals, established the genus name Palaeotherium and established the first species name Palaeotherium medium.[5][6] The genus name means "ancient beast," for which the etymology is a compound of the Greek prefix παλαιός ('palaios') meaning 'old' or 'ancient' and the suffix θήρ ('thēr') meaning 'beast' or 'wild animal'.[7] He debunked Lamanon's hypothesis that Palaeotherium was an omnivorous amphibian and suspected that it had trunks akin to those of tapirs.[5][2]
Later, he wrote about a species that he deemed to have similar dentition to P. medium to the extent that it belonged to Palaeotherium. Cuvier observed that the species had larger-sized dentition compared to the other species based on imprints that he was provided. Thus, he established the species Palaeotherium magnum. He also erected the genus Anoplotherium in recognition of its differing dentition from that of Palaeotherium.[8] He also gave mentions to the postcranial fossils of the genus and listed a newly recognized species named Palaeotherium minus.[9] In a later journal of the same year, Cuvier described a mostly complete skeleton from the French commune of Pantin that he determined to have belonged to P. minus.[10]
In 1805, Cuvier described additional postcranial fossil bones of Palaeotherium. He noted that its forefeet consisted of three short toes and that no other animal had postcranial bones that closely resembled those of the extinct genus. Nonetheless, he also gave emphasis to some of the fossil foot bones resembling those of either tapirs or rhinoceroses. Based on the metacarpal bone shapes located on the front feet, he erected the species name P. crassum, mentioning that it was a distinct species from P. medium.[11]
In 1812, he examined more metacarpal bone material that he classified as belonging to Palaeotherium. He stated that the newer material was nearly the size of those of P. crassum but that it was shorter than even those of P. minus. Thus, he felt the need to establish another species P. curtum based on the fossils. He then made a review of the species he previously erected. According to Cuvier, P. magnum was the size of a horse. He then stated that both P. medium and P. crassum were both the size of a pig, the former having narrow and elongated feet and the having broader and shorter feet. P. curtum did not have any listed size analogue, but the naturalist said that it had the feet of a horse. Finally, he said that P. minus was the size of a sheep and had narrow feet. He also listed five new additional species from surrounding areas of France that he did not further elaborate on, such as the rhinoceros-sized P. giganteum, the ox-sized P. tapiroïdes, the pig-sized P. buxovillanum plus P. aurelianense, and the sheep-sized P. occitanicum.[12]
The naturalist also suggested palaeobiologies of the four species of Palaeotherium that he described from the gypsum quarries. He acknowledged that P. magnum had skull and limb bone material but lacked vertebra and rib fossils. Regardless, he was able to speculate based on available material that P. magnum would have resembled a tapir the size of a horse with bare amounts of hair. He also hypothesized that P. crassum would have resembled a tapir and been the size of one, which in theory would have caused people to confuse the two. P. medium, he suggested, would have also resembled a tapir but differed by higher legs and longer feet. He was able to construct a speculative skeletal reconstruction drawing of P. minus because of a previously found skeleton and hypothesized that it was smaller than a sheep and could have been cursorial with its slender legs and face. Finally, he theorized that P. curtum would have been the bulkiest species with lower legs compared to P. minus that were stocky like those of P. crassum. Cuvier also suggested that Palaeotherium as in the entire genus was tridactyl (or three-toed).[12][13]
In 1822, Cuvier recognized additional species of Palaeotherium based on postcranial materials. He also depicted a drawn reconstruction of the skeleton of P. magnum, outlining that it was the size of a Javan rhinoceros, was stocky in body build, and had a massive head.[14] Palaeotherium was also depicted in 1822 drawings by the French palaeontologist Charles Léopold Laurillard under the direction of Cuvier.[15] In 1824, he listed most species of Palaeotherium that he previously named and described, namely P. magnum, P. medium, P. crassum, P. latum, P. curtum, P. minus, and P. aureliense. He also recognized an additional species P. isselanum, but he did not describe its fossils.[16]
Palaeotherium magnum, Palaeotherium medium and "Plagiolophus minus" (= Plagiolophus) are notably depicted in the forms of one sculpture representing each species in the Crystal Palace Dinosaurs attraction in the Crystal Palace Park in the United Kingdom, open to the public since 1854 and constructed by English sculptor Benjamin Waterhouse Hawkins. The original large-sized P. magnum sculpture was lost at some point after 1958 and was replaced by a new replicated model in 2023. The other two palaeothere statues in the park represent the medium-sized P. medium and the small-sized "P. minus" (= Plagiolophus minor). Both the large P. magnum and P. medium were posed in standing positions whereas the smallest sitting statue was made to represent "P. minus". The models' resemblances to tapirs reflected early perceptions that the palaeothere species resembled them in body plan appearances. Despite this, the sculptures differ from living tapirs in several ways, such as shorter plus taller faces, higher eye positions, slender legs, longer tails, and the presence of three toes on the forelimbs unlike the four toes of the forelimbs of tapirs.[17][18]
Of the three sculptures, P. medium most closely resembles a tapir, and it has remained mostly intact but suffered from damages that were eventually repaired. P. medium was depicted as having durable skin and a slender face with a trunk, representing archaic perceptions that it was a slow animal that lived in closed habitats. The original P. magnum sculpture was last known from a 1958 photograph of it that reveals that it was the largest sculpture of the three species and that it had a highly robust form with large and deep eyes, a proportionally large head, bulky legs, and a muscular-looking body plan. The trunk of the model appears to start from the upper section of the skull and descends down to the lower lip. The overall anatomy of the sculpture appears to have been based off of elephants compared to the other two palaeothere statues.[18]
Later 19th century taxonomy
Throughout much of the 19th century, many species were classified under Palaeotherium, some of which were eventually reclassified under different genera.[19] For instance, "P." aurelianense was reclassified to its own genus Anchitherium by Christian Erich Hermann von Meyer in 1844. In an 1839–1864 osteography, the French naturalist Henri Marie Ducrotay de Blainville relisted "P." tapiroides, "P." buxovillanum and "P." occitanicum as species belonging to Lophiodon,[20] but the latter two were eventually reclassified to Paralophiodon and Lophiaspis, respectively in the 20th century.[21][22] In 1862, Swiss zoologist Ludwig Ruetimeyer defined the previously recognized genera Plagiolophus and Propalaeotherium as distinct from Palaeotherium and containing the species P. minor and P. isselanum, respectively.[23]
In 1853, Pomel erected the species P. duvali based on fossil limbs that he thought to have been less stocky compared to those of P. curtum.[24] In an 1839–1864 osteography, Blainville listed Palaeotherium species previously recognized by other taxonomists and erected P. girondicum.[20] In 1863, the French naturalist Jean-Baptiste Noulet created the species P. castrense based on an incomplete mandible from the commune of Viviers-lès-Montagnes, where it was later studied in Castres.[25] In 1869, Pictet and Humbert erected the species Plagiolophus siderolithicus using fossil molars from a museum collection whose form is similar to that of P. minor but differs mainly by the dimensions.[26] The same year, German palaeontologist Oscar Fraas erected P. suevicum based on teeth that he thought to have distinct enamel.[27] Gervais in 1875 described fossil bones and teeth from the French commune of Dampleux, noting that the particular species was smaller than others classified to Palaeotherium and that the dental measurements were similar to those of Plagiolophus minor. He assigned the fossils to the newly erected species P. eocaenum.[28]
Palaeotherium skeletons
For much of paleontological history, Palaeotherium was not known by any complete skeleton since its initial description by Cuvier.[29] This changed when in 1873, the French geologist Gaston Casimir Vasseur uncovered the first complete skeleton of P. magnum from a gypsum quarry in the commune of Vitry-sur-Seine.[30] The quarry was owned by the civil engineer Fuchs, who donated the skeleton to the National Museum of Natural History, France.[31][32] The skeleton was first described by Gervais in an academic journal the same year, who noted that it allows for more accurate confirmations of the species' anatomical traits. He pointed out that the skeleton had a skull measures 0.5 m (1 ft 8 in) long, a longer neck than previously expected, and a less stocky build compared to tapirs and rhinoceroses. The naturalist said that the extraction process was difficult but completed by multiple skillful workers.[31] Since then, it has been displayed at the Gallery of Paleontology and Comparative Anatomy exhibit of the museum, where it had been noted as an important and famous component of the gallery to the modern day.[32][30]
During the 20th century, a second complete skeleton of P. magnum was excavated from the plasters within the French commune of Mormoiron. It was sent to the geological department of the University of Lyon and described after preparation by the Austrian geologist Frédéric Roman in 1922. Roman depicted a drawing of a reconstruction of the skeleton of P. magnum based on the Mormoiron skeleton within his 1922 monography. According to Austrian palaeontologist Othenio Abel in 1923, it was the most complete skeleton of Palaeotherium to have been found and amongst the most complete of early Cenozoic mammal skeletons, missing only a few ribs and a left femur.[29][33]
20th century revisions
In 1904, Swiss palaeontologist Hans Georg Stehlin first created the species name P. lautricense based on an upper jaw from a collection at the Muséum de Toulouse that originated from sandstone deposits at Castres. He also wrote about two somewhat crushed skulls of the species, producing a sketched reconstruction of it based on the first one.[34] In his monography for palaeotheres, published the same year, Stehlin considered most species within Palaeotherium to be potentially valid and created more of them, but he noted that most taxonomists were cautious invalidating species erected by Cuvier. Stehlin also revised P. girondicum as P. magnum var. girondicum, or P. magnum girondicum. He established the subspecies name P. curtum var. perrealense, or P. curtum perrealense, based on jaw fragments from La Débruge. He erected P. Mühlbergi based on dental material in the Swiss municipality of Obergösgen that Ruetimeyer examined back in 1862. He also stated that the recent excavations at Mormont from Natural History Museum of Basel had yielded fossils that he classified along with a mandible identified by Pictert in 1869 under the new species name P. Renevieri. Finally, he also determined that an additional species Palaeotherium Rütimeyeri, which he described as having primitive premolars, was present in the municipality of Egerkingen.[35]
In 1917, French palaeontologist Charles Depéret recognized two additional species of Palaeotherium called P. Euzetense and P. Stehlini.[36] German palaeontologist Wilhelm Otto Dietrich named the German species P. Kleini in 1922, basing it off of fossils from the locality of Mähringen and mentioning that it would have been the size of P. curtum and P. Heimi.[37]
In 1968, upcoming German palaeontologist Jens Lorenz Franzen, then a graduate student, made major revisions of Palaeotherium within his dissertation. He synonymized or rendered dubious statuses many species of Palaeotherium that were erected throughout the 19th and early 20th centuries, including those named by Cuvier. He also erected P. pomeli using fossils from a locality in Castres and reclassified "Plagiolophus" siderolithicum into Palaeotherium. Furthermore, Franzen converted some species into subspecies, namely P. magnum girondicum, P. magnum stehlini, P. medium suevicum, and P. medium euzetense. In addition, he named the following subspecies that he named in his thesis: P. castrense robiacense, P. crassum robustum, P. curtum villerealense, P. curtum frohnstettense, P. muehlbergi praecursum, and P. duvali priscum. Not all species within Palaeotherium had any recognized subspecies in them.[19]
In 1975, Spanish palaeontologist María Lourdes Casanovas-Cladellas erected the species P. crusafonti from a left maxilla with dentition from the Spanish site of Roc de Santa.[38] In 1980, both she and José-Vicente Santafé-Llopis established the second Iberian species P. franzeni, taking into account morphological differences of the dental fossils from the Spanish municipality of Sossís.[39] In 1985, the French palaeontologist Jean-Albert Remy named a subspecies P. muehlbergi thaleri in honor of fellow palaeontologist Louis Thaler, having documented that its fossils were from the commune of Saint-Étienne-de-l'Olm and that both the holotype and paratype each consist of a skull with a mandible.[40]
In 1991, Casanovas-Cladellas and Santafé-Llopis erected P. llamaquiquense from partial jaw material from the Spanish locality of Llamaquique in the city of Oviedo, where the name derived from.[41] The next year in 1992, Remy proposed the creation of two subgenera of Palaeotherium based on cranial characteristics. The first subgenus he listed was Palaeotherium, which includes the type species P. magnum along with P. medium, P. crassum, P. curtum, P. castrense, P. siderolithicum, and P. muehlbergi. The second subgenus name that he proposed was Franzenitherium, which includes the type species P. lautricense plus P. duvali and was named in honor of Franzen's review of Palaeotherium.[42] The Spanish palaeontologist Miguel Ángel Cuesta Ruiz-Colmenares established the species P. giganteum using dentition from the Mazaterón site in the Duero Basin in 1993, considering it to be the largest species of Palaeotherium known.[43] In 1998, Casanovas-Cladellas et al. formally recognized P. crassum sossissense from a fragmented right maxilla with dentition from Sossís in Spain. They also invalidated the previously named P. franzeni due to reassigning the material to P. magnum stehlini.[44]
Classification
Palaeotherium is the type genus of the Palaeotheriidae, largely considered to be one of two major hippomorph families in the superfamily Equoidea, the other being the Equidae. Alternatively, some authors have proposed that equids are more closely related to the Tapiromorpha than to the Palaeotheriidae. It is also usually thought to consist of two families, the Palaeotheriinae and Pachynolophinae; not all authors agree on the latter as a palaeotheriid subfamily, however.[45] Some authors have also considered the Plagiolophinae to be a separate subfamily, while others group its genera into the Palaeotheriinae.[46] The geographic range of the palaeotheres were in contrast to equids, which are generally thought to have been an endemic radiation in North America. Some of the most basal equoids of the European landmass are of uncertain affinities, with some genera being thought to potentially belong to the Equidae.[47] Palaeotheriids are well-known for having lived in western Europe during much of the Palaeogene but were also present in eastern Europe, possibly the Middle East, and, in the case of pachynolophines (or pachynolophs), Asia.[45][46]
The Perissodactyla makes its earliest known appearance in the European landmass in MP7 of the Mammal Palaeogene zones. During the temporal unit, many genera of basal equoids such as Hyracotherium, Pliolophus, Cymbalophus, and Hallensia made their first appearances there. A majority of the genera persisted to the MP8-MP10 units, and "pachynolophines" (probably true palaeotheres) such as Propalaeotherium and Orolophus arose by MP10.[47][48] The MP13 unit saw the appearances of later pachynolophines such as Pachynolophus and Anchilophus along with definite records of the first palaeotheriines such as Palaeotherium and Paraplagiolophus.[49] The palaeotheriine Plagiolophus has been suggested to have potentially made an appearance by MP12. It was by MP14 that the subfamily proceeded to diversify,[50] and the plagiolophines were generally replaced but still reached the late Eocene. In addition to more widespread palaeothere genera such as Plagiolophus, Palaeotherium, and Leptolophus, some of their species reaching medium to large sizes, various other palaeothere genera that were endemic to the Iberian penninsula, such as Cantabrotherium, Franzenium and Iberolophus, appeared by the middle Eocene.[49]
The phylogenetic tree for several members of the family Palaeotheriidae within the order Perissodactyla (including three outgroups) as created by Remy in 2017 and followed by Remy et al. in 2019 is defined below:[51][50]
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As shown in the above phylogeny, the Palaeotheriidae is defined as a monophyletic clade, meaning that it did not leave any derived descendant groups in its evolutionary history. Hyracotherium sensu stricto (in a strict sense) is defined as amongst the first offshoots of the family and a member of the Pachynolophinae. "H." remyi, formerly part of the now-invalid genus Propachynolophus, is defined as a sister taxon to more derived palaeotheriids. Both Pachynolophus and Lophiotherium, defined as pachynolophines, are defined as monophyletic genera. The other pachynolophines Eurohippus and Propalaeotherium consistute a paraphyletic clade in relation to members of the derived and monophyletic subfamily Palaeotheriinae (Leptolophus, Plagiolophus, and Palaeotherium), thus making Pachynolophinae a paraphyletic subfamily clade.[51]
List of lineages
Since 1968, many species of Palaeotherium have multiple defined subspecies due to taxonomic revisions conducted by Franzen involving new species plus subspecies erections and conversions of some species into subspecies that were accepted by subsequent authors. From his dissertation was he able to justify the subspecies by proof of various intraspecific variations.[19][52] The following table defines the species and subspecies of Palaeotherium and additional information about them:
Lineage | Proposed subgenus | MP unit(s) | Author(s) of taxon | Taxon publication year |
---|---|---|---|---|
P. castrense castrense | Palaeotherium | 14, 16 | Noulet | 1863 |
P. castrense robiacense | Palaeotherium | 16 | Franzen | 1968 |
P. crassum sossisense | Palaeotherium | 17 | Casanovas-Cladellas, Checa, and Santafé-Llopis | 1998 |
P. crassum crassum | Palaeotherium | 19 | Cuvier | 1805 |
P. crusafonti | Incertae sedis | 17 | Casanovas-Cladellas | 1975 |
P. curtum villerealense | Palaeotherium | 17, 18 | Franzen | 1968 |
P. curtum curtum | Palaeotherium | 19 | Cuvier | 1812 |
P. curtum frohnstettense | Palaeotherium | 20 | Franzen | 1968 |
P. duvali priscum | Franzenitherium | 17 | Franzen | 1968 |
P. duvali duvali | Franzenitherium | 19 | Pomel | 1853 |
P. eocaenum | Incertae sedis | 13, 14 | Gervais | 1875 |
P. giganteum | Incertae sedis | 16 | Cuesta | 1993 |
P. lautricense | Franzenitherium | 16 | Stehlin | 1904 |
P. llamaquiquense | Incertae sedis | 16 | Casanovas-Cladellas & Santafé Llopis | 1991 |
P. magnum stehlini | Palaeotherium | 17 | Depéret | 1917 |
P. magnum girondicum | Palaeotherium | 18 | Blainville | 1846 |
P. magnum magnum | Palaeotherium | 19, 20 | Cuvier | 1804 |
P. medium euzetense | Palaeotherium | 17 | Depéret | 1917 |
P. medium perrealense | Palaeotherium | 18 | Stehlin | 1904 |
P. medium medium | Palaeotherium | 19 | Cuvier | 1804 |
P. medium suevicum | Palaeotherium | 20, 21 | Fraas | 1869 |
P. muehlbergi praecursum | Palaeotherium | 17 | Franzen | 1968 |
P. muehlbergi thaleri | Palaeotherium | 18 | Remy | 1985 |
P. muehlbergi muehlbergi | Palaeotherium | 19, 20 | Stehlin | 1904 |
P. pomeli | Incertae sedis | 16 | Franzen | 1968 |
P. renevieri | Incertae sedis | 19 | Stehlin | 1904 |
P. ruetimeyeri | Incertae sedis | 14, 16 | Stehlin | 1904 |
P. siderolithicum | Palaeotherium | 16, 17, 18, 19 | Pictet & Humbert | 1869 |
Description
Skull
The Palaeotheriidae is diagnosed in part as generally having orbits that are wide open in the back area and are located in the middle of the skull or in a slight frontal area of it. The nasal bones are slightly extensive to very extensive in depth. Palaeotherium is characterized as having calvaria that range in base length from 150 mm (5.9 in) to 520 mm (20 in) depending on the species. The pterygoid crest, which is located on the pterygoid processes of the sphenoid bone, does not cover the optic foramen, which is separated from other cranial cavities at the temporal fossa. The zygomatic process of the squamosal bone is elongated and extends to the maxilla at a back angle of the orbit. The genus is also diagnosed by the presence of an anastomosis (anatomical connection between two passageways) roughly at the sphenoid bone and prominent temporalis muscle developments.[19][42]
According to Remy, the subgenus Palaeotherium is the more specialized one of the two, characterized by the orbit being located more in front of the skull's middle length. The optic foramen is separated by a bony wall, and there are two optic canals in total. The cranium is constricted in its front area behind the postorbital processes and close to the suture for the frontal bone and parietal bone. The other subgenus Franzenitherium in comparison has more generalized skull traits, its orbit being aligned within the middle length of the head. It has a front constriction of the cranium near the postorbital processes, and the optic foramen crosses through the skull from one side to the other. Not all species are placed in any subgenus due to having inadequate skull evidence for further analyses.[42]
The height and weight proportions of the skull of Palaeotherium are roughly equivalent with those of other taxa within the Equoidea. In comparison to other equoids where the skull's maximum width extends above the front root of the parallel zygomatic arches, those of Palaeotherium and most other palaeotheres (except Leptolophus) extend back to the joint of the squamosal bone and mandible. Unlike that of Plagiolophus, the maxillary hiatus, or an opening of the maxillary sinus, in Palaeotherium is wide, diamond-shaped, and has oblique back edges. Palaeotherium differs from most other palaeotheres by the nasal opening stretching up to the P3 tooth at minimum (noticeable in P. duvali and P. siderolithicum) or up to the front edge of the orbit above M3 in the case of P. magnum. Similar to other palaeotheres, the back process (tissue projection) of the premaxilla is reduced, but its morphology can vary. The maxilla can extend to the nasal opening but can also vary in proportions. While the shapes and proportions of the nasal bones vary by species, they extend beyond P1 in adults and sometimes even the canine like in equines. The nasomaxillary suture, which unites the maxilla and nasal bones, is short and strongly curved.[42]
The jugal bone and lacrimal bone, both located in front of the orbit, are weak in development. The latter bone is elongated in its back and touches the posterolateral process of the nasal bones. Members of the Equoidea have relatively shortened front areas of the face. The orbits of Palaeotherium, unlike those of other equoids, are proportionally smaller and are situated somewhat in front of the skull's mid-length area; they might be more forwards in the case of P. medium. Similar to other Palaeogene equoids, the front edge of the orbit is aligned with M1 or M2 while the back area is wide. In most species, the infraorbital foramen in adults is located above P4 or M1; in P. curtum frohnstettense, it extends to above M2. Each zygomatic arch is wide, and its uneven narrowing in the front area under the orbit may be the result of either species traits or sexual differentiation. The squamosal process of the postorbital is elongated and reaches the maxilla at a back angle of the orbital floor, the roof of the maxillary sinus. The orbit is shallow within its front area, its back opening of the infraorbital canal having a small distance of 10 mm (0.39 in) to 15 mm (0.59 in). The canal itself is smaller than those of most other palaeotheriines except for that of Leptolophus, and it has a tendency to shorten in later species.[42]
The side wall of the snout is usually concave but may be interrupted by other local concavities that are normally poorly distinguished. The palatine bone has a well-developed front area, which in the case of P. curtum villerealense can rise near the orbit's top. The sphenopalatine foramen, large and oval-shaped, is above the back of M4. The optic canal is small, has a primitive form of opening in front of the pterygoid crest, and is separated from it by a thick bony wall except in the cases of P. lautricense and P. duvali. Whereas there are two optic canals in most species that are nearly parallel of each other and separate in the end, there is only one long one in P. lautricense and possibly P. duvali. The sphenoid bone's wings are well-developed in their backs, and a suture for the parietal and sphenoid bones separates the frontal bone from the squamosal bone. The postorbital processes of the frontal bone are not very elongated. The cranial vault is broad, domed, and wider than the overall skull. The maximum front narrowing of the cranium, with the exceptions of those of P. lautricense and P. duvali, is set far back to roughly where the frontoparietal suture occurs. The skull's top peaks at the far back area, although this is not observed in P. lautricense. The sagittal crest can be prominent and depends on the age and sex of the individual for development. The nuchal crest, where the neck attaches to the head, is prominent and, except in P. lautricense, extends outwards plus backwards past the occipital condyles. The temporal fossae are large but vary in proportion. In Palaeotherium and not Plagiolophus, the overall basicranium's axis is thick plus wide.[42]
The horizontal ramus of the mandible is overall thick plus tall and has an elongated mandibular symphysis, but the width and lower area morphology vary by species. It is wide in both the front and back areas and low compared to equines. The joint for the squamosal and mandible of Palaeotherium is low compared to those of Plagiolophus and Leptolophus. The angular process, located above the angle of the mandible, is blocked from further expansion by the mandibular notch and is well-developed in its rear like in Palaeogene equids. The coronoid process of the mandible, an upper eminence, is both broad in the front plus back and stocky. The condyloid process, an upper process of the mandible, is transversely elongated and cylindrical in shape. Both the zygomatico-mandibular fossa and pterygoid fossa are prominent.[42]
Dentition
Postcranial skeleton
Footprints
Palaeobiology
Palaeoecology
Extinction
Notes
References
- ^ de Lamanon, Robert de Paul (1782). "Description de divers fossiles trouvés dans les carrières de Montmartre près Paris, & vues générales sur la formation des pierres gypseuses". Introduction aux Observations sur la Physique, sur l'Histoire Naturelle et sur les Arts. 19: 173–194.
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