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-avia ...
.
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 ...
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 betwee ...
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 an ...
period, 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 foss ...
David J. Gower 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''.Gower (1999), p. 6. In contrast with sprawling reptiles, like
crocodile Crocodiles (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 include all extant me ...
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'',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 the ...
.


Description

''Batrachotomus'' was a heavily built, large quadrupedal reptile reaching in length. A trait that characterized ''Batrachotomus'', compared to other crurotarsans, 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. 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, ''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 me ...
(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 fenestrae (skull openings), two pairs of which were for the eyes (called orbits) 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 fenestrae. 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 fenestrae 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.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 ...
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. 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 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 Fossil collecting (sometimes, in a non-scientific sense, fossil hunting) is the collection of fossils for scientific study, hobby, or profit. Fossil collecting, as practiced by amateurs, is the predecessor of modern paleontology and many sti ...
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, ...
. 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,
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 Sw ...
. 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 seve ...
(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. The family name "Prestosuchidae" was established in 1966 by American paleontologist Alfred Romer. 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, 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 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 ...
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 gr ...
. The members of this clade were recovered as basal loricatan, of which ''Batrachotomus'' was found to be the most derived i.e. most closely related to the clade containing Crocodylomorpha and Rauisuchidae. Subsequent derivatives of this analysis further support this hypothesis. In a yet to be formally published revision of '' Heptasuchus'', a medium/large-sized (~6.5 m long) "rauisuchian" from the upper Chugwater Group of
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 t ...
, 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 ...
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 ''Gerrothorax'' ("wicker chest") is an extinct genus of temnospondyl amphibian from the Triassic period of Greenland, Germany, Poland, Sweden, and possibly Thailand. It is known from a single species, ''G. pulcherrimus'', although several other ...
'' and ''Mastodonsaurus'', and even animals like nothosaurs 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 th ...
'' Tanystropheus''. 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 ...
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 exce ...
s,
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male o ...
s and
conifer Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...
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