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''Elachistosuchus'' 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
neodiapsid Neodiapsida is a clade, or major branch, of the reptilian family tree, typically defined as including all diapsids apart from some early primitive types known as the araeoscelidians. Modern reptiles and birds belong to the neodiapsid subclade ...
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
, most likely basal
archosauromorph Archosauromorpha (Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, liza ...
, known from the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...
Arnstadt Formation of
Saxony-Anhalt Saxony-Anhalt (german: Sachsen-Anhalt ; nds, Sassen-Anholt) is a state of Germany, bordering the states of Brandenburg, Saxony, Thuringia and Lower Saxony. It covers an area of and has a population of 2.18 million inhabitants, making it the ...
, central
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 ...
. It contains a single species, ''Elachistosuchus huenei'', known from a single individual. ''E. huenei'', originally considered a
pseudosuchia Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
n
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 ...
and then a
rhynchocephalia Rhynchocephalia (; ) is an order of lizard-like reptiles that includes only one living species, the tuatara (''Sphenodon punctatus'') of New Zealand. Despite its current lack of diversity, during the Mesozoic rhynchocephalians were a diverse g ...
n
lepidosaur The Lepidosauria (, from Greek meaning ''scaled lizards'') is a subclass or superorder of reptiles, containing the orders Squamata and Rhynchocephalia. Squamata includes snakes, lizards, and amphisbaenians. Squamata contains over 9,000 species ...
, was largely ignored in the scientific literature, as its small size and fragility did not permit further mechanical preparation and examination. More recently however a non-invasive μCT scanning was performed to resolve its placement within
Reptilia Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
, and found it to represent a more basal reptile, potentially closely related to several early archosauromorph
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
s.


Discovery

Fossils of ''Elachistosuchus'' were first described and named by the German
paleontologist 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 ...
Werner Janensch Werner Ernst Martin Janensch (11 November 1878 – 20 October 1969) was a German paleontologist and geologist. Biography Janensch was born at Herzberg (Elster). In addition to Friedrich von Huene, Janensch was probably Germany's most imp ...
in
1949 Events January * January 1 – A United Nations-sponsored ceasefire brings an end to the Indo-Pakistani War of 1947. The war results in a stalemate and the division of Kashmir, which still continues as of 2022. * January 2 – Luis ...
, with 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 ...
being ''Elachistosuchus huenei''. The generic name is derived from
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 ...
''ἐλάχιστος'' / ''elachistos'', meaning "diminutive", and ''suchus'',
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
ized Greek ''souchos'', an Egyptian crocodile god, in reference to the size of the holotype and its identification by Janensch as a
pseudosuchia Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
n archosaur. The specific name ''huenei'' honors the German paleontologist
Friedrich von Huene Friedrich von Huene, born Friedrich Richard von Hoinigen, (March 22, 1875 – April 4, 1969) was a German paleontologist who renamed more dinosaurs in the early 20th century than anyone else in Europe. He also made key contributions about v ...
who greatly contributed to the study of various
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
groups, especially extinct reptiles. Janensch described ''Elachistosuchus'' based on an associated partial
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
of a single individual, 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 ...
MB.R. 4520, which consists of six small blocks (I–VI) of bone-bearing rock currently housed at the fossil reptile collection of the Berlin's Natural History Museum, in Germany. Block I preserves a nearly complete but crushed
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
, while the right
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
associated with articulated
vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
including the back portion of the neck and the front portion of the back with
rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...
fragments are preserved in block II, along with shoulder girdle elements including the interclavicle,
clavicle The clavicle, or collarbone, is a slender, S-shaped long bone approximately 6 inches (15 cm) long that serves as a strut between the shoulder blade and the sternum (breastbone). There are two clavicles, one on the left and one on the rig ...
s,
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
s and one scapula, first observed using μCT scanning. Blocks III to VI consist of fragments of ribs, isolated back vertebrae, and gastralia. MB.R. 4520 and additional unprepared material attributed to ''Elachistosuchus'' were found during the excavation of a skeleton of ''
Plateosaurus ''Plateosaurus'' (probably meaning "broad lizard", often mistranslated as "flat lizard") is a genus of plateosaurid dinosaur that lived during the Late Triassic period, around 214 to 204 million years ago, in what is now Central and Northern Eur ...
'' around 1928. They were collected from a locality which lies along the present-day highway (Bundesstrasse) B79 between
Halberstadt Halberstadt ( Eastphalian: ''Halverstidde'') is a town in the German state of Saxony-Anhalt, the capital of Harz district. Located north of the Harz mountain range, it is known for its old town center that was greatly destroyed by Allied bombi ...
and
Quedlinburg Quedlinburg () is a town situated just north of the Harz mountains, in the district of Harz in the west of Saxony-Anhalt, Germany. As an influential and prosperous trading centre during the early Middle Ages, Quedlinburg became a center of in ...
, on the southeastern edge of
Halberstadt Halberstadt ( Eastphalian: ''Halverstidde'') is a town in the German state of Saxony-Anhalt, the capital of Harz district. Located north of the Harz mountain range, it is known for its old town center that was greatly destroyed by Allied bombi ...
, in
Saxony-Anhalt Saxony-Anhalt (german: Sachsen-Anhalt ; nds, Sassen-Anholt) is a state of Germany, bordering the states of Brandenburg, Saxony, Thuringia and Lower Saxony. It covers an area of and has a population of 2.18 million inhabitants, making it the ...
, from a brick-
clay Clay is a type of fine-grained natural soil material containing clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4). Clays develop plasticity when wet, due to a molecular film of water surrounding the clay par ...
pit unit of the Arnstadt Formation, dating to the middle to late
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
stage of the middle
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...
. The site was initially excavated by
Otto Jaekel Otto Max Johannes Jaekel (21 February 1863 – 6 March 1929) was a German paleontologist and geologist. Biography Jaekel was born in Neusalz (Nowa Sól), Prussian Silesia, the son of a builder and the youngest of seven children. He studied at ...
between 1909 and 1912 and later Janensch conducted additional excavations there between 1923 and 1928, but except for smaller digs by A. Hemprich in 1937 and 1938 no further work has taken place since. The diverse Late Triassic biota from Halberstadt includes
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
s,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
chondrichthyan Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. C ...
s, dipnoans,
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 ...
s, stem-
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked tu ...
s,
phytosaur Phytosaurs (Φυτόσαυροι in greek) are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. Phytosauria and Phytosauridae are often considered to be equivalent ...
s, and a haramiyid mammal, as well as the
sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lon ...
''Plateosaurus'', which is represented by some 50 specimens including at least two complete skeletons.


Description

''Elachistosuchus'' was originally thought to be a
pseudosuchia Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
n archosaur by Janensch (1949). Janensch diagnosed it among other pseudosuchians by its small body size, the lack of a specialized body plan, and the presence of a large post
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, ...
and what he believed to be an ant
orbital fenestra The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
. Walker (1966) reinterpreted ''Elachistosuchus'' as a
rhynchocephalia Rhynchocephalia (; ) is an order of lizard-like reptiles that includes only one living species, the tuatara (''Sphenodon punctatus'') of New Zealand. Despite its current lack of diversity, during the Mesozoic rhynchocephalians were a diverse g ...
n closely related to the only living genus ''
Sphenodon Tuatara (''Sphenodon punctatus'') are reptiles endemic to New Zealand. Despite their close resemblance to lizards, they are part of a distinct lineage, the order Rhynchocephalia. The name ''tuatara'' is derived from the Māori language a ...
'' (the
tuatara Tuatara (''Sphenodon punctatus'') are reptiles endemic to New Zealand. Despite their close resemblance to lizards, they are part of a distinct lineage, the order Rhynchocephalia. The name ''tuatara'' is derived from the Māori language and m ...
), based on the long front portion of the
jugal bone The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
, the allegedly
acrodont Acrodonty (from Greek ''akros'' 'highest' + ''dont'' 'tooth') is an anatomical placement of the teeth at the summit of the alveolar ridge of the jaw, without sockets, characteristic of bony fish. Functionally, acrodont tooth implantation may be rela ...
dentition, the large posttemporal fenestra, the absence of an external
lower jaw In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
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 ...
, and a strongly twisted end of the
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
. Walker (1966) also argued that the antorbital fenestra identified by Janensch actually represented a damaged opening for the
lacrimal canal The term Lacrimal or lachrymal, may refer to: Anatomy * Lacrimal apparatus * Lacrimal artery * Lacrimal bone * Lacrimal canaliculi (singular: canaliculus), also known as ''Lacrimal ducts'' * Lacrimal fossa (disambiguation) * Lacrimal fluid, see Te ...
. Due to the small size and fragility of the holotype, further mechanical preparation is not possible, which caused ''Elachistosuchus'' to be largely ignored in the scientific literature. More recently however a non-invasive μCT scanning allowed for the examination of MB.R. 4520 and assessment of much of its internal cranial structure. Additionally, the scan revealed previously unknown parts of the skeleton concealed in the matrix, specifically the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
,
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 ...
, and parts of the shoulder girdle. Sobral, Sues and Müller (2015) provided a revised diagnosis for the species, based on these new observations. ''Elachistosuchus'' was considered by them to represent a small diapsid reptile, probably a basal
archosauromorph Archosauromorpha (Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, liza ...
(based on the results of several
phylogenetic analyses 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 ...
), with a distinct back-side portion of the
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.''Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, par ...
,
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 ...
ry tooth row that extends behind the back margin of
eye socket In anatomy, the orbit is the cavity or socket of the skull in which the eye and its appendages are situated. "Orbit" can refer to the bony socket, or it can also be used to imply the contents. In the adult human, the volume of the orbit is , of ...
, a
jugal bone The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
with free back process, a palatine ramus of the
pterygoid bone The pterygoid is a paired bone forming part of the palate of many vertebrates, behind 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 th ...
with
shagreen Shagreen is a type of rawhide consisting of rough untanned skin, historically from a horse's or onager's back, or from shark or ray. Etymology The word derives from the French ''chagrin'' and is related to Italian ''zigrino'' and Venetian '' ...
of teeth, an
angular bone The angular is a large bone in the lower jaw (mandible) of amphibians and reptiles (birds included), which is connected to all other lower jaw bones: the dentary (which is the entire lower jaw in mammals), the splenial, the suprangular, and the art ...
exposed along about one third of the side surface of
mandibula In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
r ramus, dichocephalous chest ribs, a notched front margin of 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 th ...
, and a spoon-shaped back projection of the interclavicle.


Phylogeny

The
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 ...
position of ''Elachistosuchus'' was explored by Sobral, Sues and Müller (2015) two recently constructed data matrix. Using Chen ''et al.'' (2014) data set, ''Elachistosuchus'' was recovered as the sister taxon of
Choristodera Choristodera (from the Greek χωριστός ''chōristos'' + δέρη ''dérē'', 'separated neck') is an extinct order of semiaquatic diapsid reptiles that ranged from the Middle Jurassic, or possibly Triassic, to the late Miocene (168 to 1 ...
, or in a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with it. The two were recovered either as the
basalmost In phylogenetics, basal is the direction of the ''base'' (or root) of a rooted phylogenetic tree or cladogram. The term may be more strictly applied only to nodes adjacent to the root, or more loosely applied to nodes regarded as being close to th ...
archosauromorph Archosauromorpha (Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, liza ...
s, or alternatively as
lepidosauromorph Lepidosauromorpha (in PhyloCode known as ''Pan-Lepidosauria'') is a group of reptiles comprising all diapsids closer to lizards than to archosaurs (which include crocodiles and birds). The only living sub-group is the Lepidosauria, which cont ...
s more advanced than
sauropterygian Sauropterygia (" lizard flippers") is an extinct taxon of diverse, aquatic reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosauria bec ...
s and their relatives. Using the Ezcurra ''et al.'' (2014) data set, Sobral, Sues and Müller (2015) found ''Elachistosuchus'' with ''
Coelurosauravus ''Coelurosauravus'' (meaning "hollow lizard grandfather") is a genus of basal diapsid reptiles, known from the Late Permian of Madagascar. Like other members of the family Weigeltisauridae, members of this genus possessed long, rod-like ossifica ...
'' just outside
Sauria Sauria is the clade containing the most recent common ancestor of archosaurs (such as crocodilians, dinosaurs, etc.) and lepidosaurs ( lizards and kin), and all its descendants. Since most molecular phylogenies recover turtles as more closely re ...
using parsimony, or alternatively as the basalmost archosauromorph (out of the included taxa), using the Bayesian approach. Although still uncertain, the phylogenetic position as broadly recovered by these analyses, rejects
pseudosuchia Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
n and
rhynchocephalia Rhynchocephalia (; ) is an order of lizard-like reptiles that includes only one living species, the tuatara (''Sphenodon punctatus'') of New Zealand. Despite its current lack of diversity, during the Mesozoic rhynchocephalians were a diverse g ...
n affinities for ''Elachistosuchus'', suggested by earlier studies. The two
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 ...
s below shows simplified topologies recovered by Sobral, Sues and Müller (2015) when using the Bayesian approach for Chen ''et al.'' (2014) and Ezcurra ''et al.'' (2014), respectively (relations within bold terminal
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
s are not shown).


References

{{Taxonbar, from=Q5353038 Prehistoric archosauromorphs Prehistoric reptile genera Norian genera Late Triassic reptiles of Europe Triassic Germany Fossils of Germany Fossil taxa described in 1949 Taxa named by Werner Janensch