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''Trimerorhachis'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
dvinosauria Dvinosaurs are one of several new clades of Temnospondyl amphibians named in the phylogenetic review of the group by Yates and Warren 2000. They represent a group of primitive semi-aquatic to completely aquatic amphibians, and are known from t ...
n
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order (biology), order of small to giant tetrapods—often considered Labyrinthodontia, primitive amphi ...
within the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Trimerorhachidae Trimerorhachidae is a family of dvinosaurian temnospondyls, including Trimerorhachis and Neldasaurus. They are vertebrates and carnivores. Gallery Trimerorhachis insignis life restoration.jpg, '' Trimerorhachis insignis'', of the early Permia ...
. It is known from the
Early Permian 01 or '01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * '01 (Richard Müller album), 01'' (Richard Müller album), 2001 * 01 (Son of Dave album), ''01'' (Son of Dave album), 2000 * 01 (Urban ...
of the southwestern United States, with most fossil specimens having been found in the Texas Red Beds. 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 specimen ...
of ''Trimerorhachis'', ''T. insignis'', was named by American paleontologist
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy interested ...
in 1878. Cope named a second species from Texas, ''T. mesops'', in 1896. The species ''T. rogersi'' (named in 1955) and ''T. greggi'' (named in 2013) are also from Texas, and the species ''T. sandovalensis'' (named in 1980) is from New Mexico.


Description

The length of the largest specimens of ''Trimerorhachis'' is estimated to have been almost a metre (3.3 feet) in length. ''Trimerorhachis'' has a large triangular head with upward-facing eyes positioned near the front of the skull. The trunk is long and the limbs are relatively short. The presence of a
branchial apparatus The pharyngeal apparatus is an embryological structure. It consists of: * pharyngeal grooves (from ectoderm) * pharyngeal arch The pharyngeal arches, also known as visceral arches'','' are structures seen in the embryonic development of vert ...
indicates that ''Trimerorhachis'' had external gills in life, much like the modern
axolotl The axolotl (; from nci, āxōlōtl ), ''Ambystoma mexicanum'', is a paedomorphic salamander closely related to the tiger salamander. Axolotls are unusual among amphibians in that they reach adulthood without undergoing metamorphosis. I ...
. The body of ''Trimerorhachis'' is also completely covered by small and very thin
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amp ...
s, which overlap and can be up to 20 layers thick. These osteoderms act as an armor-like covering, especially around the tail. Their weight may have helped ''Trimerorhachis'' sink to the bottom of lakes and rivers where it would feed.


History

''Trimerorhachis'' was first described by Edward Drinker Cope in 1878. Specimens are often preserved as masses of bones that are mixed together and densely packed in slabs of rock. Fossils are rarely found in articulation, although a slab of rock has been found with sixteen skulls and their associated vertebrae in an intact position. Most of these fossils preserve skulls and dorsal vertebrae, but rarely any other bones. Paleontologist S.W. Williston of the
University of Chicago The University of Chicago (UChicago, Chicago, U of C, or UChi) is a private research university in Chicago, Illinois. Its main campus is located in Chicago's Hyde Park neighborhood. The University of Chicago is consistently ranked among the b ...
commented in 1915 that "it will only be by the fortunate discovery of a connected skeleton that the tail, ribs, and feet will be made known." A nearly complete specimen was discovered the following year near
Seymour, Texas Seymour is a city in and the county seat of Baylor County, Texas, United States. Its population was 2,575 as of the 2020 Census. Geography Seymour is located on the Brazos River. It is southwest of Wichita Falls and north-northeast of Abilen ...
, and Williston was able to describe the entire postcranial skeleton of ''Trimerorhachis''. In 1955, paleontologist
Edwin Harris Colbert Edwin Harris "Ned" Colbert (September 28, 1905 – November 15, 2001)O'Connor, Anahad ''The New York Times'', November 25, 2001. was a distinguished American vertebrate paleontologist and prolific researcher and author. Born in Clarinda, Iowa, h ...
described the scales of ''Trimerorhachis''. He noted that they were oval-shaped and overlapping and that each had a base layer of longitudinal striations covered by another layer ring-like ridges, the growth rings of the scales. The scales were more similar to fish scales than they were to reptile scales. In 1979, paleontologist Everett C. Olson claimed that there were no such scales in ''Trimerorhachis'', and that Colbert was incorrect in his interpretation of the body covering of ''Trimerorhachis''. A second species called ''T. sandovalensis'' was named from New Mexico in 1980. A nearly complete skeleton from the
Abo Formation The Abo Formation is a geologic formation in New Mexico. It contains fossils characteristic of the Cisuralian epoch of the Permian period. Description The Abo Formation consists of fluvial redbed mudstones and sandstones, including river channel ...
near Jemez Springs has been designated the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
, but other fossils of the species are found throughout the state, giving it a wide distribution.


Paleobiology


Environment

''Trimerorhachis'' was probably a fully aquatic temnospondyl. Like most dvinosaurs, it had external gills. The
interclavicle An interclavicle is a bone which, in most tetrapods, is located between the clavicles. Therian mammals (marsupials and placentals) are the only tetrapods which never have an interclavicle, although some members of other groups also lack one. In t ...
and
clavicle The clavicle, or collarbone, is a slender, S-shaped long bone approximately 6 inches (15 cm) long that serves as a strut between the shoulder blade and the sternum (breastbone). There are two clavicles, one on the left and one on the rig ...
of the
pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of ...
are both very large, a feature that is shared with other aquatic temnospondyls. Many bones are poorly
ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by Cell (biology), cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes ...
, indicating that ''Trimerorhachis'' was poorly suited for movement on land. ''Trimerorhachis'' was probably an aquatic predator that fed on fish and small vertebrates. Microanatomical data also suggest a fairly aquatic lifestyle, as shown by the obvious osteosclerosis of its femur. During the Early Permian, the area of New Mexico and Texas was a broad coastal plain that stretched from an ocean in the south to highlands in the north. Other common animals that lived alongside ''Trimerorhachis'' included
lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
and
crossopterygian Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includin ...
s, the
lepospondyl Lepospondyli is a diverse taxon of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco (''Diplocaulus minumus''), lepospondyls lived from the Early Carboniferous ( Mississippian) to the Early Per ...
''
Diplocaulus ''Diplocaulus'' (meaning "double caul") is an extinct genus of lepospondyl amphibians which lived from the Late Carboniferous to the Late Permian of North America and Africa. ''Diplocaulus'' is by far the largest and best-known of the lepospondy ...
'', and the large sail-backed
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
''
Dimetrodon ''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodontid ...
''.


Brooding

Small bones that likely belong to immature ''Trimerorhachis'' individuals have been found in the pharyngeal pouches of larger ''Trimerorhachis'' specimens. At first these bones were thought to be part of the
branchial arch Branchial arches, or gill arches, are a series of bony "loops" present in fish, which support the gills. As gills are the primitive condition of vertebrates, all vertebrate embryos develop pharyngeal arches, though the eventual fate of these arc ...
es which surround the pouch, or remains of prey that had just been eaten before the animal died. If ''Trimerorhachis'' was a
mouth brooder Mouthbrooding, also known as oral incubation and buccal incubation, is the care given by some groups of animals to their offspring by holding them in the mouth of the parent for extended periods of time. Although mouthbrooding is performed by a va ...
, the closest living analogue would be
Darwin's Frog Darwin’s frog (''Rhinoderma darwinii''), also called the Southern Darwin's frog, is a species of Chilean/Argentinian frog of the family Rhinodermatidae. It was discovered by Charles Darwin during his voyage on voyage on HMS ''Beagle''. on a ...
, which broods its young in its
vocal sac The vocal sac is the flexible membrane of skin possessed by most male frogs and toads. The purpose of the vocal sac is usually as an amplification of their mating or advertisement call. The presence or development of the vocal sac is one way ...
. The bones in ''Trimerorhachis'' belong to juveniles that were much larger than those of Darwin's Frog, however. The young of the
Gastric-brooding frog The gastric-brooding frogs or platypus frogs (''Rheobatrachus'') is a genus of extinct ground-dwelling frogs native to Queensland in eastern Australia. The genus consisted of only two species, both of which became extinct in the mid-1980s. The g ...
s of Australia were comparable in size to those of ''Trimerorhachis'' but were brooded in the stomach rather than the throat. The number of brooded young in Darwin's Frog and the Gastric-brooding frogs is also much higher than that of ''Trimerorhachis'', as only a few individuals can be distinguished in the collection of bones. The only living amphibian that raises similarly sized young is the Golden coquí, although it does so through
ovovivipary Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop ins ...
rather than brooding. Another possible explanation for the small bones is that they were originally located in the throat and were pushed into the pharyngeal pouch during fossilization. If this was the case, ''Trimerorhachis'' may have eaten its young instead of brooding them. This type of cannibalism is widespread in living amphibians, and most likely occurred among some prehistoric amphibians as well.


See also

*
List of prehistoric amphibians This list of prehistoric amphibians is an attempt to create a comprehensive listing of all Genus, genera from the fossil record that have ever been considered to be amphibians, excluding purely vernacular terms. The list includes all commonly accep ...


References

{{Taxonbar, from=Q7842192 Dvinosaurs Cisuralian temnospondyls of North America Permian geology of New Mexico Permian geology of Texas Taxa named by Edward Drinker Cope Fossil taxa described in 1878