Trimerorhachis Cope
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''Trimerorhachis'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
dvinosauria Dvinosaurs are one of several new clades of temnospondyls named in the phylogenetic review of the group by Yates and Warren 2000. They represent a group of primitive semi-aquatic to completely aquatic temnospondyls, and are known from the Late C ...
n
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
within the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Trimerorhachidae Trimerorhachidae is a family of dvinosaurian temnospondyls, including ''Lafonius'', ''Trimerorhachis'', ''Procuhy'' and ''Neldasaurus''. They were semiaquatic carnivores that lived from the Pennsylvanian (geology), Late Carboniferous to the Earl ...
. 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 (Urban Zakapa album), ''01'' (Urban Zakapa album), 2011 * ''01011 ...
of the southwestern United States, with most fossil specimens having been found in the Texas Red Beds. The
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
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, paleontology, paleontologist, comparative anatomy, comparative anatomist, herpetology, herpetologist, and ichthyology, ichthyologist. Born to a wealthy Quaker fam ...
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

Largest specimens of ''Trimerorhachis'' reached a length of about 1.5 meters (5 ft), although most have been no more than 1.2 meters (4 ft) 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 indicates that ''Trimerorhachis'' had external gills in life, much like the modern
axolotl The axolotl (; from ) (''Ambystoma mexicanum'') is a neoteny, paedomorphic salamander, one that Sexual maturity, matures without undergoing metamorphosis into the terrestrial adult form; adults remain Aquatic animal, fully aquatic with obvio ...
. 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 amph ...
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, or UChi) is a Private university, private research university in Chicago, Illinois, United States. Its main campus is in the Hyde Park, Chicago, Hyde Park neighborhood on Chicago's South Side, Chic ...
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, he ...
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 Everett Claire Olson (November 6, 1910 – November 27, 1993) was an American zoologist, paleontologist, and geologist noted for his seminal research of origin and evolution of vertebrate animals. Through his research studying terrestrial verte ...
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 near Jemez Springs has been designated the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
, 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 ...
and
clavicle The clavicle, collarbone, or keybone is a slender, S-shaped long bone approximately long that serves as a strut between the scapula, shoulder blade and the sternum (breastbone). There are two clavicles, one on each side of the body. The clavic ...
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 o ...
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 cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...
, 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 class Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, inc ...
and
crossopterygian Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...
s, the
lepospondyl Lepospondyli is a diverse clade of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco ('' Diplocaulus minimus''), lepospondyls lived from the Visean stage of the Early Carboniferous to the Earl ...
''
Diplocaulus ''Diplocaulus'' (meaning "double stalk") 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 lepospond ...
'', and the large sail-backed
synapsid Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
''
Dimetrodon ''Dimetrodon'' ( or ; ) is an extinct genus of sphenacodontid synapsid that lived during the Cisuralian (Early Permian) Epoch (geology), epoch of the Permian period, around 295–272 million years ago. With most species measuring long and ...
''.


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 paired bony/ cartilaginous "loops" behind the throat ( pharyngeal cavity) of fish, which support the fish gills. As chordates, all vertebrate embryos develop pharyngeal arches, though the eve ...
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 frog of the family Rhinodermatidae. It was discovered by Charles Darwin during his voyage on HMS ''Beagle''. on a trip to Chile. In 1841, French ...
, which broods its young in its
vocal sac The vocal sac is the flexible Biological membrane, 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 voca ...
. 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 ''Rheobatrachus'', whose members are known as the gastric-brooding frogs or platypus frogs, is a genus of extinct ground-dwelling frogs native to Queensland Queensland ( , commonly abbreviated as Qld) is a States and territories of Austra ...
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 The golden coquí (''Eleutherodactylus jasperi'') is a rare, possibly extinct species of frog endemic to Puerto Rico. General description Golden coquís are roughly in snout-vent length and are olive-gold to yellow-gold without pattern. The juv ...
, although it does so through
ovovivipary Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop inside eggs that rema ...
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 genera from the fossil record that have ever been considered to be amphibians, excluding purely vernacular terms. The list includes all commonly accepted gen ...


References

{{Taxonbar, from=Q7842192 Dvinosauria 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