Gerobatrachus hottoni
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''Gerobatrachus'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
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
amphibamid The Amphibamidae are an extinct Family (biology), family of Dissorophoidea, dissorophoid temnospondyls known from Late Carboniferous-Early Permian strata in the United States. Classification Amphibamidae has traditionally included small-bodied ...
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 ...
(represented by 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 ...
''Gerobatrachus hottoni'') that lived in 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 ...
, approximately 290 million years ago (Ma), in the area that is now
Baylor County, Texas Baylor County is a county located in the U.S. state of Texas. As of the 2020 census, its population was 3,465. Its county seat is Seymour. History In 1858, the Texas Legislature established Baylor County, naming it for Henry Weidner Baylor, ...
. When it was first described in 2008, ''Gerobatrachus'' was announced to be the closest relative of
Batrachia The Batrachia are a clade of amphibians that includes frogs and salamanders, but not caecilians nor the extinct allocaudates. The name Batrachia was first used by French zoologist Pierre André Latreille in 1800 to refer to frogs, but has more ...
, the group that includes modern frogs and salamanders. It possesses a mixture of characteristics from both groups, including a large frog-like head and a salamander-like tail. These features have led to it being dubbed a frogamander by the press. Some more recent studies place ''Gerobatrachus'' as the closest relative of
Lissamphibia The Lissamphibia is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia (frogs, toads, and their extinct relatives), the Caudata (salamanders, newts, and their extinct relatives), ...
, the group that contains all modern amphibians including frogs, salamanders, and
caecilian Caecilians (; ) are a group of limbless, vermiform or serpentine amphibians. They mostly live hidden in the ground and in stream substrates, making them the least familiar order of amphibians. Caecilians are mostly distributed in the tropics of ...
s, or place modern amphibians far from ''Gerobatrachus'' within a group called
Lepospondyli 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 ...
.


Description

The only known specimen of ''Gerobatrachus'' is a nearly complete skeleton (USNM 489135) about long, that is articulated, preserved in ventral view, missing only the stylopodia, zeugopodia, and ventral portions of the skull and pectoral girdle. It is preserved in red
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, p ...
with only its underside exposed. Like other amphibamid temnospondyls, ''Gerobatrachus'' has a rounded and flattened head, well-developed limbs, and a small tail. Its vertebral column is somewhat shorter than those of related amphibamids. The large, round head and shortened vertebral column are features ''Gerobatrachus'' shares in common with frogs and the early salamander ''
Karaurus ''Karaurus'' (meaning ''head-tail'') is an extinct genus of stem-group salamander (Caudata) from the Middle to Late Jurassic (Callovian–Kimmeridgian) Karabastau Formation of Kazakhstan. It is one of the oldest salamanders known. ''Karaurus'' w ...
''.''Science Daily'', "Ancient Amphibian: Debate Over Origin Of Frogs And Salamanders Settled With Discovery Of Missing Link"
Accessed 22 May 2008.
''Gerobatrachus'' also has a large embayment at the back of the skull called an
otic notch Otic notches are invaginations in the posterior margin of the skull roof, one behind each orbit. Otic notches are one of the features lost in the evolution of amniotes from their tetrapod ancestors. The notches have been interpreted as part of an ...
, which is seen other amphibamids and in frogs and supports the tympanum, an eardrum-like structure used in hearing. Many finer details of the skull link ''Gerobatrachus'' with modern amphibians. ''Gerobatrachus'' has a row of very small
pedicellate teeth Pedicellate teeth are a tooth morphology today unique to modern amphibians, but also seen in a variety of extinct labyrinthodonts. Pedicellate teeth consist of a tooth crown and a base (both composed of dentine) separated by a layer of uncalcifie ...
, a feature shared with modern amphibians. Pedicellate teeth are characterized by two layers of hardened
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
, one at the tooth base and one at the tooth tip, surrounding a middle layer of softer, uncalcified dentine. The
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
, which in other amphibians bears many distinguishing characteristics, is poorly preserved in ''Gerobatrachus''. The preserved portion lacks the large palatal teeth commonly found in other temnospondyls. The
vomer The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxill ...
bone is much shorter in length than it is in other amphibamids, but similar in proportion to living amphibians. As in most frogs, the
palatine bone In anatomy, the palatine bones () are two irregular bones of the facial skeleton in many animal species, located above the uvula in the throat. Together with the maxillae, they comprise the hard palate. (''Palate'' is derived from the Latin ''pa ...
forms a narrow strip along the side of the palate. ''Gerobatrachus'' possesses another modern amphibian characteristic at the back of the skull, a widened bone called the parasphenoid basal plate. Many of the features that link ''Gerobatrachus'' with modern amphibians are also seen in other amphibamids. For example, the amphibamids ''
Amphibamus ''Amphibamus'' is a genus of amphibamid temnospondyl amphibians from the Carboniferous (middle Pennsylvanian) of North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisp ...
'', ''
Doleserpeton ''Doleserpeton'' is an extinct, monospecific genus of dissorophoidean temnospondyl within the family Amphibamidae that lived during the Upper Permian, 285 million years ago. ''Doleserpeton'' is represented by a single species, ''Doleserpeton ann ...
'', ''
Eoscopus ''Eoscopus'' is an extinct genus of dissorophoidean euskelian temnospondyl in the family Micropholidae. It is known from Hamilton Quarry, a Late Carboniferous lagerstätte near Hamilton, Kansas. Members of Micropholidae were historically inclu ...
'', and ''
Platyrhinops ''Platyrhinops'' is an extinct genus amphibamid temnospondyl from the Late Carboniferous (late Westphalian stage) of Ohio and the Czech Republic. It is known from many partial skeletons from the Linton site in Saline Township, Ohio and at lea ...
'' also have large otic notches; ''Amphibamus'', ''Doleserpeton'', and ''
Tersomius ''Tersomius'' is an extinct genus of dissorophoid temnospondyl within the family Micropholidae. It is known from the early Permian of North America (Oklahoma and Texas). History of study ''Tersomius'' was first named in 1910 by American paleo ...
'' also have pedicellate teeth; and ''Amphibamus'' and ''Doleserpeton'' also have a wide parasphenoid plate.


Discovery

''Gerobatrachus hottoni'' was described for the first time on May 22, 2008 in the journal
Nature Nature, in the broadest sense, is the physics, physical world or universe. "Nature" can refer to the phenomenon, phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. ...
. The nearly complete
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 ...
skeleton USNM 489135 was collected from a fossil locality known as Don's Dump Fish Quarry in Baylor County, Texas in 1995. It was then rediscovered in the collections of the
National Museum of Natural History The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. In 2021, with 7 ...
, Washington, D.C., in 2004. The genus name means "elder frog" in
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
, from (géros, "elder") and (batrakhos, "frog"). The species name honors
Nicholas Hotton III Nicholas Hotton III (January 28, 1921 – November 29, 1999) was an American paleontologist renowned as an expert on dinosaurs and reptiles. Early life and education Hotton was born in Sault Ste. Marie, Michigan and was educated at the Univers ...
, a paleontologist who had worked for the National Museum of Natural History.


Relationships

When ''Gerobatrachus'' was first described in 2008, it was incorporated into a
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis that found it to be the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
or closest relative of
Batrachia The Batrachia are a clade of amphibians that includes frogs and salamanders, but not caecilians nor the extinct allocaudates. The name Batrachia was first used by French zoologist Pierre André Latreille in 1800 to refer to frogs, but has more ...
, an evolutionary group that includes living frogs and salamanders but not caecilians, which are the third major lineage of modern amphibians (the three main groups of modern amphibians are collectively known as lissamphibians). As was the case in some previous analyses, caecilians were found to be the descendants of a group of small amphibious Paleozoic tetrapods called Lepospondyli while frogs and salamanders had an independent origin within Temnospondyli. The phylogeny supported by the 2008 analysis has been called the "
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...
hypothesis" and is one of three main hypotheses for lissamphibian origins. The other two hypotheses are the "lepospondyl hypothesis", which places all lissamphibians as the closest relatives of a group of lepospondyls called lysorophians, and the "temnospondyl hypothesis", which has all lissamphibians derived from amphibamid temnospondyls like ''Gerobatrachus''. Paleontologists David Marjanović and Michel Laurin published studies in 2008 and 2009 that supported the lepospondyl hypothesis. Their phylogenetic analyses confirmed that ''Gerobatrachus'' was an amphibamid temnospondyl, and since all modern amphibians nested within Lepospondyli, it was positioned far from the ancestry of frogs and salamanders. A 2012 study of the
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
-
caecilian Caecilians (; ) are a group of limbless, vermiform or serpentine amphibians. They mostly live hidden in the ground and in stream substrates, making them the least familiar order of amphibians. Caecilians are mostly distributed in the tropics of ...
''
Eocaecilia ''Eocaecilia'' is an extinct genus of gymnophionan amphibian from the early Jurassic Kayenta Formation of Arizona, United States. One species is described, ''Eocaecilia micropodia''. ''Eocaecilia'' shared some characteristics with salamanders a ...
'' found ''Gerobatrachus'' to group within Lissamphibia. In this phylogeny, ''Gerobatrachus'' is more closely related to frogs and salamanders than it is to caecilians, meaning that ''Gerobatrachus'' would have been a descendant of the
last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of modern amphibians. Below is a
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
from the original 2008 phylogenetic analysis (left) and a cladogram from the 2012 analysis (right):


References

{{Taxonbar, from=Q132803 Amphibamids Dissorophids Cisuralian temnospondyls of North America Permian geology of Texas Transitional fossils Fossil taxa described in 2008 Prehistoric amphibian genera