Periptychus
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''Periptychus'' is an extinct genus of mammal belonging to the family
Periptychidae Periptychidae is a family of Paleocene placental mammals, known definitively only from North America. The family is part of a radiation of early herbivorous and omnivorous mammals formerly classified in the extinct order " Condylarthra", which ...
. It lived from the Early to Late
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pal ...
and its fossil remains have been found in North America.


Description

This animal was of medium size and could exceed one meter in overall length; ''Periptychus'' is supposed to have weighed about 23 kilograms. ''Periptychus'' was an unusual mammal that combined a number of rather specialized dental, cranial, and postcranial features with a relatively generalized skeletal structure.


Skull

The shape of the skull of ''Periptychus'' was almost identical to that of early
Eutheria Eutheria (; from Greek , 'good, right' and , 'beast'; ) is the clade consisting of all therian mammals that are more closely related to placentals than to marsupials. Eutherians are distinguished from noneutherians by various phenotypic tra ...
, although it was more robust. The snout of ''Periptychus'' was moderately elongated and tall, and tapered anteriorly without a rostral constriction. The
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
of the rostrum of this animal was very similar to that found in related genera such as '' Carsioptychus'' or '' Ectoconus''; the snout was not elongated as in other
condylarth Condylarthra is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene epochs. They are considered early, primitive ungulates. It is now largely considered to be a waste ...
s such as ''
Arctocyon ''Arctocyon'' ('bear dog') is an extinct genus of ungulate mammals. ''Arctocyon'' was a "ground dwelling omnivore", that lived from 61.3-56.8 Ma. Synonyms of ''Arctocyon'' include ''Claenodon'', and ''Neoclaenodon''. ''Arctocyon'' was likely plant ...
'', but it was longer than that of
pantodonts Pantodonta is an extinct suborder (or, according to some, an order) of eutherian mammals. These herbivorous mammals were one of the first groups of large mammals to evolve (around 66 million years ago) after the end of the Cretaceous. The l ...
such as '' Pantolambda''. In lateral view, the dorsal surface of the skull of ''Periptychus'' was relatively flat compared to the more domed morphology found in ''Pantolambda''. The zygomatic arches of ''Periptychus'' were wide and protruded laterally. The skull was small and low, with well-developed nuchal and sagittal ridges. The latter, in particular, was robust and provided a substantial insertion point for the temporal muscles along the entire length of the cranial box. The paired nasal bones were very elongated and went to form a wide, flat "roof" over the rostrum. The
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. T ...
was characterized by an unusual pitted surface, just above the tooth row; these holes, concentrated mainly above the premolars and around the infraorbital foramen, varied in shape and size: some were circular and others ovoid, and did not exceed 0.7 millimeters in diameter. In the living animal, these holes probably housed a network of capillaries that supply blood to the jaws and were connected to the vibrissae. The
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
of ''Periptychus'' was particularly robust; the paired dental bones were co-ossified toward the end, and went to form a high mandibular
symphysis A symphysis (, pl. symphyses) is a fibrocartilaginous fusion between two bones. It is a type of cartilaginous joint, specifically a secondary cartilaginous joint. # A symphysis is an amphiarthrosis, a slightly movable joint. # A growing togethe ...
that extended from the
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
s to the ventral limit of the mandible, at the level of the anterior margin of the second inferior premolar. The ascending ramus of the mandible was rather high in relation to the mandibular body, and went to form a rounded, high and wide coronoid process. The dentition of ''Periptychus'' was distinctive: the premolars were larger than the
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
and possessed highly expanded central cusps, while the molars were highly
bunodont The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
and possessed bulbous cusps compressed at the apex. The dental enamel was strongly crenulated with well-aligned ridges, typical of all periptychids but reaching their maximum development in ''Periptychus.'' The canine was relatively small and single-rooted.


Postcranial skeleton

The postcranial skeleton of ''Periptychus'' was that of a robust, strong-footed animal with a plantigrade gait. The
lumbar vertebrae The lumbar vertebrae are, in human anatomy, the five vertebrae between the rib cage and the pelvis. They are the largest segments of the vertebral column and are characterized by the absence of the foramen transversarium within the transverse p ...
were robust and dorsoventrally compressed. The caudal vertebrae were also robust, rod-like; the tail must have been rather long and heavy, although not as long as that of ''Ectoconus.'' The forelimbs were mainly characterized by a humerus that was rather short compared to
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
and
radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...
, with a hemispherical head equipped with large low tuberosities; the humerus also had a wide and elongated deltopectoral region and a region for the insertion of the ''teres major'' muscle that was rather small. Expanded medial and lateral entepicondyles were also present, and the humerus-radial joint was open. The ulna was rather straight and gracile, with a posterior bulge of the
diaphysis The diaphysis is the main or midsection (shaft) of a long bone. It is made up of cortical bone and usually contains bone marrow and adipose tissue (fat). It is a middle tubular part composed of compact bone which surrounds a central marrow cavit ...
, while the
olecranon The olecranon (, ), is a large, thick, curved bony eminence of the ulna, a long bone in the forearm that projects behind the elbow. It forms the most pointed portion of the elbow and is opposite to the cubital fossa or elbow pit. The olecranon ...
was massive and formed an almost rectangular projection at the proximal part of the ulna. The radius was robust compared to the ulna. The
carpus In human anatomy, the wrist is variously defined as (1) the carpus or carpal bones, the complex of eight bones forming the proximal skeletal segment of the hand; "The wrist contains eight bones, roughly aligned in two rows, known as the carpal ...
was wide and had an enlarged central bone. The hand had five fingers, and the number of phalanges was the generalist number for mammals (2-3-3-3). The bones of the fingers were rather short and compact, and ended in nail-like
hoof The hoof (plural: hooves) is the tip of a toe of an ungulate mammal, which is covered and strengthened with a thick and horny keratin covering. Artiodactyls are even-toed ungulates, species whose feet have an even number of digits, yet the rum ...
phalanges. The ilium of ''Periptychus'' was slender and elongated, and in dorsal view was concave in the central area. The
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
was typical of a robustly built animal, and it had strong
trochanter A trochanter is a tubercle of the femur near its joint with the hip bone. In humans and most mammals, the trochanters serve as important muscle attachment sites. Humans are known to have three trochanters, though the anatomic "normal" includes ...
s; a third trochanter was present. The greater trochanter was high but did not extend beyond the head of the femur. The
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
was also strong, but somewhat elongated, while the
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity i ...
was small. The foot of ''Periptychus'' was plantigrade, pentadactylous and paraxonic in structure, with wide toes well spaced apart, much like those of the hand. The
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
was dorsoventrally compressed, and its joint was wedged between the tibia and fibula, allowing the latter to make contact with the calcaneus. A tibial bone was present, sandwiched between the tendons of the posterior tibial muscle. The phalanges of the tarsus were almost identical to those of the hand, but were slightly larger.


Classification

''Periptychus'' was first described by Edward Drinker Cope in
1881 Events January–March * January 1– 24 – Siege of Geok Tepe: Russian troops under General Mikhail Skobelev defeat the Turkomans. * January 13 – War of the Pacific – Battle of San Juan and Chorrillos: The ...
, based on fossil remains found in
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ke ...
in deposits dating to the Middle Paleocene. The fossils were very fragmentary including a part of a jaw with teeth and belonging to a juvenile specimen, which Cope did not recognize as such and assigned it to
Creodonta Creodonta ("meat teeth") is a former order of extinct carnivorous placental mammals that lived from the early Paleocene to the late Miocene epochs in North America, Europe, Asia and Africa. Originally thought to be a single group of animals ance ...
. In the same year Cope described a jawbone, also from New Mexico, and described it as ''Catathleus rhabdodon''. Only later, through the discovery of more complete remains, did it become clear that the two forms were synonymous and belonged to the same species. To the genus ''Periptychus'' was then also ascribed by Cope himself the species ''P. coarctatus,'' which is slightly older and perhaps ancestral to the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
. Fossils of ''Periptychus'' have since also been found in
Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest U.S. state by ...
,
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
and
Wyoming Wyoming () is a state in the Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the southwest, and Colorado to the s ...
. ''Periptychus'' is the eponymous genus of the family
Periptychidae Periptychidae is a family of Paleocene placental mammals, known definitively only from North America. The family is part of a radiation of early herbivorous and omnivorous mammals formerly classified in the extinct order " Condylarthra", which ...
, a family of archaic mammals that developed immediately after the Cretaceous-Tertiary extinction event, and produced a variety of forms of different sizes. In particular, ''Periptychus'' would appear to be a derived member of the group.


Paleobiology

''Periptychus'' was endowed with many features such as the broad facial region, high sagittal and nuchal ridges, dentition with enlarged premolars and highly developed enamel that indicate a
durophagous Durophagy is the eating behavior of animals that consume hard-shelled or exoskeleton bearing organisms, such as corals, shelled mollusks, or crabs. It is mostly used to describe fish, but is also used when describing reptiles, including fossil tu ...
diet, based on dense, fibrous plants. The postcranial skeleton of ''Periptychus'' indicates that this animal was plantigrade, adapted to move slowly over the ground; it was also capable, on occasion, of making rapid movements. Despite its robust structure, ''Periptychus'' retained a fairly high degree of movement capacity in its legs: they were very powerful but did not make very rapid movements. The flexor and extensor muscles in the hands and fingers were highly developed, indicating that this animal was probably capable of climbing and digging.


References

*E. D. Cope. 1881a. Mammalia of the lower Eocene beds. American Naturalist 15:337-338 *E. D. Cope 1881b. On some mammals of the Lower Eocene beds of New Mexico. Proc Am Philos Soc. 19: 484-495. *E. D. Cope. 1883. On some fossils of the Puerco Formation. Proc Acad Nat Sci;35: 168-170. *O. C. Marsh. 1894. Description of artiodactyls of the Tertiary. American Journal of Science 48(285):259-275. *G. G. Simpson. 1936. Additions to the fauna of the Puerco, Lower Paleocene. Am Mus Novit;849:1-12. *W. D. Matthew. 1937. Paleocene faunas of the San Juan Basin, New Mexico. Transactions of the American Philosophical Society 30:1-510. *J. A. Schiebout. 1974. Vertebrate paleontology and paleoecology of the Black Peaks Paleocene Formation, Big Bend National Park, Texas. Bulletin of the Texas Memorial Museum 24:1-88. *L. M. Van Valen. 1978. The beginning of the mammalian age. Evolutionary Theory 4:45-80. *J. K. Rigby. 1980. Swain Quarry of the Fort Union Formation, Middle Paleocene (Torrejonian), Carbon County, Wyoming: geologic setting and mammalian fauna. Evolutionary Monographs 3:1-178 *T. E. Williamson. 1996. The beginning of the mammal era in the San Juan Basin, New Mexico; biostratigraphy and evolution of Paleocene mammals of the Nacimiento Formation. Bulletin of the New Mexico Museum of Natural History and Science 8:1-141. *T. J. D. Halliday, P. Upchurch, and A. Goswami. 2017. Resolving relationships of Paleocene placental mammals. 92:521-550 *Shelley SL, Williamson TE, Brusatte SL (2018) The osteology of Periptychus carinidens: A robust, ungulate-like placental mammal (Mammalia: Periptychidae) from the Paleocene of North America. PLoS ONE 13(7): e0200132. https://doi.org/10.1371/journal.pone.0200132 {{Taxonbar, from=Q60978042 Condylarths Paleocene mammals of North America Paleogene United States Fossils of the United States Puercan Torrejonian Tiffanian Fossil taxa described in 1881 Prehistoric placental genera Taxa named by Edward Drinker Cope