Batrachotomus Calcaneum Labelled
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''Batrachotomus'' is a genus of prehistoric
archosaur Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
.
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 lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
and dated from the Ladinian stage of the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epoch (geology), epochs of the Triassic period (geology), period or the middle of three series (stratigraphy), series in which the Triassic system (stratigraphy), system is di ...
period Period may refer to: Common uses * Period (punctuation) * Era, a length or span of time *Menstruation, commonly referred to as a "period" Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (o ...
, around 242 to 237
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
. ''Batrachotomus'' was described by
palaeontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
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: 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 of all kno ...
''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, large ...
''.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 more loosely to include ...
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 ...
'',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 Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
.


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 thought ...
, 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 taxon, extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, Temnospondyli, ...
. 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 ...
, ''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 (: phalanx ) are digit (anatomy), digital bones in the hands and foot, feet of most vertebrates. In primates, the Thumb, thumbs and Hallux, big toes have two phalanges while the other Digit (anatomy), digits have three phalanges. ...
) are poorly preserved and the only well known bone is a fifth
metatarsal The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
(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; : fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biology, biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomy, ...
e (skull openings), two pairs of which were for the eyes (called
orbits In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an physical body, object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an satellite, artificia ...
) and the
nostril A nostril (or naris , : 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 turbinates ...
s. Behind the orbits were two
temporal fenestra Temporal fenestrae are openings in the temporal region of the skull of some amniotes, behind the orbit (eye socket). These openings have historically been used to track the evolution and affinities of reptiles. Temporal fenestrae are commonly (al ...
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 Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...
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. Human dentition is heterodont and diphyodont as an example. In vertebrates, heterodont pertains to animals where ...
.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 mammals h ...
e (bones at the very tip of the upper jaw) were slender, unlike those of the
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
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 states of Germany, 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 i ...
. 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 ...
, 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 N ...
, and in
Crailsheim Crailsheim () is a town in the States of Germany, German state of Baden-Württemberg. Incorporated in 1338, it lies east of Schwäbisch Hall and southwest of Ansbach in the Schwäbisch Hall (district), Schwäbisch Hall district. The city's mai ...
. 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), Hohenlohe district, in Baden-Württemberg, Germany. It is situated on the river Kocher, 4 km northwest of Künzelsau, and 36 km northeast of Heilbronn. History Numerous archeologica ...
,
Stuttgart Stuttgart (; ; Swabian German, Swabian: ; Alemannic German, Alemannic: ; Italian language, Italian: ; ) is the capital city, capital and List of cities in Baden-Württemberg by population, largest city of the States of Germany, German state of ...
. 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 (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 ...
(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 the ...
, 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 crocodilians ...
. 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, United States. It was founded as "Northern Illinois State Normal School" in 1895 by Illinois Governor John P. Altgeld, initially to provide the state with c ...
. Parrish hypothesized that ''Batrachotomus'' (then "Kupferzellia") belonged to the family of
Rauisuchidae Rauisuchidae is a group of large (up to ) predatory Triassic archosaurs. Some disagreement exists over which genera should be included in the Rauisuchidae and which should be in the related Prestosuchidae and Poposauridae, and indeed whether the ...
, 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 Triassic, Late Triassic period (235–228 million years ago). It ...
''.Parrish (1993), p. 301. However, the description of the braincase and a revisited
cladistic analysis Cladistics ( ; from Ancient Greek 'branch') 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 ...
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 the activity of assigning objects to some pre-existing classes or categories. This is distinct from the task of establishing the classes themselves (for example through cluster analysis). Examples include diagnostic tests, identif ...
of basal archosaurs, and using the most comprehensive
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analysis for this group (to date) found Prestosuchidae to be non-
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
. 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. Extinct crocodylomorphs were considerably mor ...
and
Rauisuchidae Rauisuchidae is a group of large (up to ) predatory Triassic archosaurs. Some disagreement exists over which genera should be included in the Rauisuchidae and which should be in the related Prestosuchidae and Poposauridae, and indeed whether the ...
. 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 Triassic, Middle or Late Triassic upper Chugwater Group of Wyoming, United States. It contains a single species, ''Heptasuchus clarki'', the first formally recog ...
'', a medium/large-sized (~6.5 m long) "rauisuchian" from the upper Chugwater Group of
Wyoming Wyoming ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States, Western United States. It borders Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho 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 s ...
'' and ''Mastodonsaurus'', and even animals like
nothosaur Nothosaurs (superfamily Nothosauroidea) were Triassic marine sauropterygian reptiles. They averaged about in length, with a long body and tail. The feet were paddle-like, and are known to have been webbed in life, to help power the animal when sw ...
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. Only about 100 of the 12,000 extant reptile species and subspecies are classed as marine reptiles, including mari ...
''
Tanystropheus ''Tanystropheus'' (~ 'long' + 'hinged') is an extinct genus of archosauromorph reptile which lived during the Triassic period, Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than t ...
''. The flora of the locality consisted of
horsetail ''Equisetum'' (; horsetail) is the only living genus in Equisetaceae, a family of vascular plants that reproduce by spores rather than seeds. ''Equisetum'' is a "living fossil", the only living genus of the entire subclass Equisetidae, which ...
s,
fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
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 sin ...
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 Loricata Middle Triassic pseudosuchians Taxa named by David J. Gower Prehistoric pseudosuchian genera Ladinian genera