Batrachotomus
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''Batrachotomus'' is a genus of prehistoric
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
.
Fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
of this animal have been found in southern
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
and dated from the Ladinian stage of the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma and ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
, around 242 to 237
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
. ''Batrachotomus'' was described by
palaeontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
David J. Gower David J. Gower is a palaeontologist. Before making his debut for the Strongroom CC in 2000, he was a herpetology researcher at the Museum of Natural History in London London is the capital and List of urban areas in the United Kingdom, ...
22 years after its discovery. The locality where ''Batrachotomus'' lived was a swampy region and the name comes from the
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 ...
''batrachos/βάτραχος'' (frog) and ''tome/τομή'' (cutting, slicing), which refers to its preying on the large amphibian ''
Mastodonsaurus ''Mastodonsaurus'' (meaning "teat tooth lizard") is an extinct genus of temnospondyl amphibian from the Middle Triassic of Europe. It belongs to a Triassic group of temnospondyls called Capitosauria, characterized by their large body size and pre ...
''.Gower (1999), p. 6. In contrast with sprawling reptiles, like
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...
s, this large carnivore was very agile with locomotor superiority due to its erect stance. A remarkable feature seen on its back was a row of paired, flattened bony plates. ''Batrachotomus'' was possibly an early relative of ''
Postosuchus ''Postosuchus'', meaning "Crocodile from Post", is an extinct genus of rauisuchid reptiles comprising two species, ''P. kirkpatricki'' and ''P. alisonae'', that lived in what is now North America during the Late Triassic. ''Postosuchus'' is a me ...
'',Gower (1999), p. 1. which lived during the dawn of the
dinosaurs Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
.


Description

''Batrachotomus'' was a heavily built, large quadrupedal reptile reaching in length. A trait that characterized ''Batrachotomus'', compared to other
crurotarsans Crurotarsi is a clade of archosauriform reptiles that includes crocodilians and stem-crocodilians and possibly bird-line archosaurs too if the extinct, crocodile-like phytosaurs are more distantly related to crocodiles than traditionally though ...
, was a series of paired small plates on its back which were attached to each vertebra.Gower and Schoch (2009), p. 117. These bony deposits forming scales are called
osteoderms 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 ...
. Flattened and leaf-shaped, these extended from behind the head along the column and reducing in size, ended at the tail. There is also evidence that osteoderms were present on the ventral region of the tail, as seen in '' Ticinosuchus ferox'', and even on the flank, belly and limbs.Gower and Schoch (2009), p. 104. Like
rauisuchians "Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilians t ...
, ''Batrachotomus'' walked with an erect posture, although the limbs were not located directly under the trunk. The limbs were not equal in length as the forelimbs were about 70% of the hindlimbs.Gower and Schoch (2009), p. 109. The toe bones (
phalanges The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...
) are poorly preserved and the only well known bone is a fifth
metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the med ...
(bone in hindlimbs attached to the toe bones) which was hooked in shape. However, hypotheses suggest that probably each forelimb had four toes and each hindlimb five.Gower and Schoch (2009), p. 119 (Figure 8). ''Batrachotomus'' had a tall and narrow skull estimated at in length.Gower (1999), p. 7. It had five pairs of
fenestra A fenestra (fenestration; plural fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomical st ...
e (skull openings), two pairs of which were for the eyes (called
orbits In celestial mechanics, an orbit is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an object or position in space such as a p ...
) and the
nostril A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbi ...
s. Behind the orbits were two
temporal fenestra An infratemporal fenestra, also called the lateral temporal fenestra or simply temporal fenestra, is an opening in the skull behind the orbit in some animals. It is ventrally bordered by a zygomatic arch. An opening in front of the eye sockets, ...
e. These holes probably helped to reduce the weight of the skull and enabled the jaw to open more widely.Benton (2005), p. 112. As a typical archosaur, ''Batrachotomus'' had two
antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, birds ...
e between the orbits and nostrils, and a fifth pair of small openings at the rear part of the lower jaw. The jaws contained sharp teeth which were compressed laterally and unequal in size and shape,Gower (1999), p. 37. and this variation of tooth shape is known as
heterodonty In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. In vertebrates, heterodont pertains to animals where teeth are differentiated into different forms. For example, ...
.Gower (1999), p. 38. The teeth on the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
e (bones at the very tip of the upper jaw) were slender, unlike those of 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. The t ...
e (the main tooth-bearing bones in the upper jaw) which had a straight posterior edge.Gower (1999), pp. 37–38. The upper jaw bore 30 teeth, with each premaxilla carrying about 4 teeth and each maxilla 11, while the lower jaw held 22 teeth.


Discovery and history

Remains of ''Batrachotomus'' have been found in southern Germany, mainly in the
Kupferzell Kupferzell is a small German town in the district of Hohenlohe in Baden-Württemberg, Germany named after the Kupfer river flowing through it. The largest neighbouring towns are Künzelsau (to the north) and Schwäbisch Hall (to the south). Hi ...
fossil locality in northern
Baden-Württemberg Baden-Württemberg (; ), commonly shortened to BW or BaWü, is a German state () in Southwest Germany, east of the Rhine, which forms the southern part of Germany's western border with France. With more than 11.07 million inhabitants across a ...
. Fossil collector Johann G. Wegele discovered the first specimens in a 1977 excavation at the
Erfurt Formation The Erfurt Formation, also known as the Lower Keuper (German: ''Untere Keuper'', ''Lettenkeuper'', ''Lettenkohle'' or ''Lettenkohlenkeuper''), is a stratigraphic formation of the Keuper group and the Germanic Trias supergroup. It was deposited du ...
, dated from the Longobardian (late Ladinian) age.Gower (1999), p. 5.Gower and Schoch (2009), p. 121. Other remains attributed to ''Batrachotomus'' have been collected in Vellberg-Eschenau, about 10 km east of
Schwäbisch Hall Schwäbisch Hall (; "Swabian Hall"; from 1802 until 1934 and colloquially: ''Hall'' ) is a city in the German state of Baden-Württemberg located in the valley of the Kocher river, the longest tributary (together with its headwater Lein) of the ...
, and in
Crailsheim Crailsheim is a town in the German state of Baden-Württemberg. Incorporated in 1338, it lies east of Schwäbisch Hall and southwest of Ansbach in the Schwäbisch Hall district. The city's main attractions include two Evangelical churches, a ...
. The most notable are from Vellberg-Eschenau, which are represented by well preserved ribs and vertebrae (MHI 1895), and evidence of forelimbs and hindlimbs (SMNS 90018). ''Batrachotomus'' today is displayed in the Muschelkalk Museum,
Ingelfingen Ingelfingen is a town in the Hohenlohe district, in Baden-Württemberg, Germany. It is situated on the river Kocher, 4 km northwest of Künzelsau, and 36 km northeast of Heilbronn. Twin towns Ingelfingen is twinned with: * Saint ...
,
Stuttgart Stuttgart (; Swabian: ; ) is the capital and largest city of the German state of Baden-Württemberg. It is located on the Neckar river in a fertile valley known as the ''Stuttgarter Kessel'' (Stuttgart Cauldron) and lies an hour from the ...
. The fossils recovered from a
marlstone Marl is an earthy material rich in carbonate minerals, clays, and silt. When hardened into rock, this becomes marlstone. It is formed in marine or freshwater environments, often through the activities of algae. Marl makes up the lower part ...
remained undescribed until 1999 and palaeontologists referred to the genus simply as "rauisuchid" or "Kupferzellia". In 1999, palaeontologist David J. Gower described 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 ...
(SMNS 52970) from the 1977 excavation, which is the largest specimen of the genus, comprised by incomplete skull and postcranial material. Anatomy of the braincase (SMNS 80260) was made three years later, shedding light on the evolutionary relationships of the poorly known group of Rauisuchia.Gower (2002), p. 49. In 2009, Gower and Rainer R. Schoch reported a detailed reconstruction of the postcranial skeleton for the first time.Gower and Schoch (2009), p. 103.


Classification

''Batrachotomus'' was a
prestosuchid Prestosuchidae (in its widest usage) is a polyphyletic grouping of carnivorous archosaurs that lived during the Triassic. They were large active terrestrial apex predators, ranging from around in length. They succeeded the Erythrosuchidae as th ...
, a member of a family of carnivorous archosaurs within the larger group
Rauisuchia "Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilia ...
. The family name "Prestosuchidae" was established in 1966 by American paleontologist
Alfred Romer Alfred Sherwood Romer (December 28, 1894 – November 5, 1973) was an American paleontologist and biologist and a specialist in vertebrate evolution. Biography Alfred Romer was born in White Plains, New York, the son of Harry Houston Romer an ...
. Prestosuchids were quadrupedal reptiles, medium to large in size, characterized by erect posture, large and narrow skull and large antorbital openings. Attention was first brought to ''Batrachotomus'' in 1993 by Michael Parrish, a palaeontologist at
Northern Illinois University Northern Illinois University (NIU) is a public research university in DeKalb, Illinois. It was founded as Northern Illinois State Normal School on May 22, 1895, by Illinois Governor John P. Altgeld as part of an expansion of the state's system ...
. Parrish hypothesized that ''Batrachotomus'' (then "Kupferzellia") belonged to the family of
Rauisuchidae Rauisuchidae is a group of large (up to or more) predatory Triassic archosaurs. There is some disagreement over which genera should be included in Rauisuchidae and which should be in the related Prestosuchidae and Poposauridae, and indeed whethe ...
, another clade of carnivorous reptiles, and species of ''
Rauisuchus ''Rauisuchus'' (meaning "Wilhelm Rau's crocodile") is a genus of extinct archosaurs which lived in what is now the Geopark of Paleorrota (Santa María Formation), Brazil, during the Late Triassic period (235–228 million years ago). It conta ...
''.Parrish (1993), p. 301. However, the description of the braincase and a revisited
cladistic analysis Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived chara ...
by Benton and
Walker Walker or The Walker may refer to: People *Walker (given name) *Walker (surname) *Walker (Brazilian footballer) (born 1982), Brazilian footballer Places In the United States *Walker, Arizona, in Yavapai County *Walker, Mono County, California * ...
, showing the close relationships between ''Batrachotomus'' and ''Prestosuchus'', led to the transfer of ''Batrachotomus'' to the family Prestosuchidae. Sterling J. Nesbitt (2011) revised the
classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organizat ...
of basal archosaurs, and using the most comprehensive
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 for this group (to date) found Prestosuchidae to be non-
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. The members of this clade were recovered as basal
loricata Loricata is a clade of archosaur reptiles that includes crocodilians and some of their Triassic relatives, such as ''Postosuchus'' and ''Prestosuchus''. More specifically, Loricata includes Crocodylomorpha (the persistent archosaur subset which ...
n, of which ''Batrachotomus'' was found to be the most derived i.e. most closely related to the clade containing
Crocodylomorpha Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cro ...
and
Rauisuchidae Rauisuchidae is a group of large (up to or more) predatory Triassic archosaurs. There is some disagreement over which genera should be included in Rauisuchidae and which should be in the related Prestosuchidae and Poposauridae, and indeed whethe ...
. Subsequent derivatives of this analysis further support this hypothesis. In a yet to be formally published revision of ''
Heptasuchus ''Heptasuchus'' is an extinct genus of loricatan pseudosuchian known from the Middle or Late Triassic upper Chugwater Group of Wyoming, United States. It contains a single species, ''Heptasuchus clarki'', the first formally recognized "rauisuch ...
'', a medium/large-sized (~6.5 m long) "rauisuchian" from the upper
Chugwater Group Chugwater is a town in Platte County, Wyoming, United States. As of the 2020 census, the town population was 175. History The Chugwater area, with its proximity to Fort Laramie, was visited by some of the earliest Western expeditions, includin ...
of
Wyoming Wyoming () is a U.S. state, state in the Mountain states, 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 south ...
, it was recovered as 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 ...
of ''Batrachotomus'' using a derivative of Nesbitt (2011) analysis. The cladogram below follows an analysis by Sterling J. Nesbitt (2011):


Paleoecology

Since 1977, the rich vertebrate fauna found at Baden-Württemberg reflects a moist region of the Middle Triassic in Germany. Along with ''Batrachotomus'', palaeontologists recovered remains of fishes, amphibians, such as '' Gerrothorax'' and ''Mastodonsaurus'', and even animals like
nothosaur Nothosaurs (order Nothosauroidea) were Triassic marine sauropterygian reptiles that may have lived like seals of today, catching food in water but coming ashore on rocks and beaches. They averaged about in length, with a long body and tail.F. v. ...
s and the distinct
marine reptile Marine reptiles are reptiles which have become secondarily adapted for an aquatic or semiaquatic life in a marine environment. The earliest marine reptile mesosaurus (not to be confused with mosasaurus), arose in the Permian period during the ...
''
Tanystropheus ''Tanystropheus'' (Greek ~ 'long' + 'hinged') is an extinct archosauromorph reptile from the Middle Triassic, Middle and Late Triassic epochs. It is recognisable by its extremely elongated neck, which measured long—longer than its body and t ...
''. The flora of the locality consisted of
horsetail ''Equisetum'' (; horsetail, snake grass, puzzlegrass) is the only living genus in Equisetaceae, a family of ferns, which reproduce by spores rather than seeds. ''Equisetum'' is a "living fossil", the only living genus of the entire subclass Eq ...
s,
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
s,
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
s and
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
s, suggesting that there was rich vegetation.Rozynek (2008), p. 4.


References

;Notes ;Sources * * * * * * * *


External links


''Batrachotomus''
at palaeos.com.
Der "Lurchenschlächter" ''Batrachotomus''
at bestiarium.kryptozoologie.net. {{Taxonbar, from=Q133494 Middle Triassic reptiles of Europe Paracrocodylomorphs Middle Triassic pseudosuchians Taxa named by David J. Gower Prehistoric pseudosuchian genera