Elachistosuchus
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''Elachistosuchus'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of neodiapsid
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
, most likely basal archosauromorph, 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 a ...
Arnstadt Formation of
Saxony-Anhalt Saxony-Anhalt ( ; ) is a States of Germany, state of Germany, bordering the states of Brandenburg, Saxony, Thuringia and Lower Saxony. It covers an area of and has a population of 2.17 million inhabitants, making it the List of German states ...
, central
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 ...
. It contains a single species, ''Elachistosuchus huenei'', known from a single individual ''E. huenei'', originally considered a
pseudosuchia Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
n
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 ...
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 speciose g ...
n lepidosaur, 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 commonly defined, are a group of tetrapods with an ectothermic metabolism and amniotic development. Living traditional reptiles comprise four orders: Testudines, Crocodilia, Squamata, and Rhynchocephalia. About 12,000 living spe ...
, and found it to represent a more basal reptile, potentially closely related to several early archosauromorph
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s.


Discovery

Fossils of ''Elachistosuchus'' were first described and named by the German
paleontologist 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 ...
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 ...
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 2025 * January 2 – Luis ...
, with the
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
being ''Elachistosuchus huenei''. The generic name is derived from
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 ...
''ἐλάχιστος'' / ''elachistos'', meaning "diminutive", and ''suchus'',
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
ized Greek ''souchos'', an Egyptian crocodile god, in reference to the size of the holotype and its identification by Janensch as a
pseudosuchia Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
n archosaur. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
''huenei'' honors the German paleontologist Friedrich von Huene who greatly contributed to the study of various
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
groups, especially extinct reptiles. Janensch described ''Elachistosuchus'' based on an associated partial
skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...
of a single individual, 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 ...
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, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
, while the right
humerus The humerus (; : humeri) 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 (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
associated with articulated
vertebrae Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
including the back portion of the neck and the front portion of the back with
rib In vertebrate anatomy, ribs () are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the thoracic cavity, enabling the lungs to expand and thus facilitate breathing by expanding the ...
fragments are preserved in block II, along with shoulder girdle elements including the interclavicle,
clavicle The clavicle, collarbone, or keybone is a slender, S-shaped long bone approximately long that serves as a strut between the scapula, shoulder blade and the sternum (breastbone). There are two clavicles, one on each side of the body. The clavic ...
s,
coracoid A coracoid 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 present as part of the scapula, but this is n ...
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 Eu ...
'' around 1928. They were collected from a locality which lies along the present-day highway (Bundesstrasse) B79 between
Halberstadt Halberstadt (; Eastphalian dialect, Eastphalian: ''Halverstidde'') is a town in the state of Saxony-Anhalt in central Germany, the capital of Harz (district), Harz district. Located north of the Harz mountain range, it is known for its old town ...
and
Quedlinburg Quedlinburg () is a town situated just north of the Harz mountains, in the Harz (district), district of Harz in the west of Saxony-Anhalt, Germany. As an influential and prosperous trading centre during the early Middle Ages, Quedlinburg becam ...
, on the southeastern edge of
Halberstadt Halberstadt (; Eastphalian dialect, Eastphalian: ''Halverstidde'') is a town in the state of Saxony-Anhalt in central Germany, the capital of Harz (district), Harz district. Located north of the Harz mountain range, it is known for its old town ...
in
Saxony-Anhalt Saxony-Anhalt ( ; ) is a States of Germany, state of Germany, bordering the states of Brandenburg, Saxony, Thuringia and Lower Saxony. It covers an area of and has a population of 2.17 million inhabitants, making it the List of German states ...
, from a brick-
clay Clay is a type of fine-grained natural soil material containing clay minerals (hydrous aluminium phyllosilicates, e.g. kaolinite, ). Most pure clay minerals are white or light-coloured, but natural clays show a variety of colours from impuriti ...
pit unit of the Arnstadt Formation, dating to the middle to late
Norian The Norian is a division of the Triassic geological period, Period. It has the rank of an age (geology), age (geochronology) or stage (stratigraphy), stage (chronostratigraphy). It lasted from ~227.3 to Mya (unit), million years ago. It was prec ...
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 a ...
period. 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 Nowa Sól, Neusalz (Nowa Sól), Prussian Silesia, the son of a builder and the youngest of seven children. He st ...
between 1909 and 1912, and later Janensch conducted additional excavations there between 1923 and 1928. 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 () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...
s,
crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
s, chondrichthyans, dipnoans,
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
s, stem-
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s,
phytosaur Phytosaurs (Φυτόσαυροι in Greek, meaning 'plant lizard') are an extinct group of large, mostly semiaquatic Late Triassic archosauriform or basal archosaurian reptiles. Phytosaurs belong to the order Phytosauria and are sometimes ref ...
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 lo ...
''Plateosaurus'', which is represented by some 50 specimens including at least two complete skeletons.


Description

''Elachistosuchus'' was originally thought to be a
pseudosuchia Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
n archosaur by Janensch (1949). Janensch diagnosed it among other pseudosuchians by its small body size, the lack of a specialized body plan, the presence of a large post-
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 ...
, and what he believed to be an ant
orbital fenestra The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
. 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 speciose g ...
n closely related to the only living genus '' Sphenodon'' (the
tuatara The tuatara (''Sphenodon punctatus'') is a species of reptile endemic to New Zealand. Despite its close resemblance to lizards, it is actually the only extant member of a distinct lineage, the previously highly diverse order Rhynchocephal ...
) 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 bone, zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by spe ...
, the allegedly acrodont dentition, the large posttemporal fenestra, the absence of an external
lower jaw In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
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, ...
, and a strongly twisted end of the
humerus The humerus (; : humeri) 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 (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
. Walker (1966) also argued that the antorbital fenestra identified by Janensch actually represented a damaged opening for the lacrimal canal. 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, brain-pan, or brainbox, 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 calv ...
,
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 sep ...
, 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 (based on the results of several
phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
), with a distinct back-side portion of the
frontal bone In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
,
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 ...
ry tooth row that extends behind the back margin of
eye socket In anatomy, the orbit is the cavity or socket/hole 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 ...
, 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 bone, zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by spe ...
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 (; derived from the Latin ''palatum'') are two irregular bones of the facial skeleton in many animal specie ...
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 ' and is related to Italian and Venetian ''sagrin'', deriv ...
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 ...
exposed along about one third of the side surface of
mandibula In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...
r ramus, dichocephalous chest ribs, a notched front margin of the interclavicle, and a spoon-shaped back projection of the interclavicle.


Phylogeny

The
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 ...
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 Miocene (168 to 20 or p ...
, or in a polytomy with it. The two were recovered either as the basalmost archosauromorphs, or as a lepidosauromorphs which is more advanced than sauropterygians and their relatives. Using the Ezcurra ''et al.'' (2014) data set, Sobral, Sues and Müller (2015) found ''Elachistosuchus'' with '' Coelurosauravus'' just outside Sauria 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, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
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 speciose g ...
n affinities for ''Elachistosuchus'' suggested by earlier studies. The two
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
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 In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s are not shown).


References

{{Taxonbar, from=Q5353038 Archosauromorpha 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