Lagosuchus
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''Lagosuchus'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
avemetatarsalia Avemetatarsalia (meaning "bird metatarsals") is a clade of diapsid reptiles containing all archosaurs more closely related to birds than to crocodilians. The two most successful groups of avemetatarsalians were the dinosaurs and pterosaurs. ...
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-avia ...
from the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch ...
of
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the List of South American countries by area, second-largest ...
. 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 specim ...
of ''Lagosuchus'', ''Lagosuchus talampayensis,'' is based on a small partial skeleton recovered from the early
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227 m ...
-age Chañares Formation. The holotype skeleton of ''L. talampayensis'' is fairly fragmentary, but it does possess some traits suggesting that ''Lagosuchus'' was a probable
dinosauriform Dinosauromorpha is a clade of avemetatarsalian archosaurs (reptiles closer to birds than to crocodilians) that includes the Dinosauria (dinosaurs) and some of their close relatives. It was originally defined to include dinosauriforms and lage ...
, closely related to
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 ...
. A second potential species of ''Lagosuchus'', ''L. lilloensis'', is based on an assortment of slightly larger and more well-preserved fossils. These larger specimens have been considered much more diagnostic and informative than the original small ''L. talampayensis'' skeleton. As a result, some paleontologists have placed the larger specimens into a new genus, ''
Marasuchus ''Marasuchus'' (meaning "Mara crocodile") is a genus of basal dinosauriform archosaur which is possibly synonymous with '' Lagosuchus''. Both genera lived during the Late Triassic in what is now La Rioja Province, Argentina. ''Marasuchus'' conta ...
''. ''Marasuchus'' is generally considered one of the more complete early dinosauriforms, useful for estimating ancestral traits for the origin of dinosaurs. This would also render ''Lagosuchus'' a
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium'' it may be impossible to determine whether a s ...
, simply a name referring to a fossil which is too fragmentary to have a formal genus. However, other paleontologists support the argument that ''Lagosuchus'' is a valid genus, and that ''Marasuchus'' is a
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...
of it.


History

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 specim ...
''Lagosuchus talampayensis'' was first described by Alfred S. Romer in 1971, who considered it a "
pseudosuchian 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 ...
" (then a collection of various non-dinosaurian "
thecodont Thecodontia (meaning 'socket-teeth'), now considered an obsolete taxonomic grouping, was formerly used to describe a diverse "order" of early archosaurian reptiles that first appeared in the latest Permian period and flourished until the end of t ...
s"). In 1972 he named a second species, '' Lagosuchus lilloensis'', known from larger and more well-preserved specimens. A later review by
Jose Bonaparte Jose is the English transliteration of the Hebrew and Aramaic name ''Yose'', which is etymologically linked to ''Yosef'' or Joseph. The name was popular during the Mishnaic and Talmudic periods. * Jose ben Abin * Jose ben Akabya *Jose the Galil ...
in 1975 synonymized the two species and considered ''Lagosuchus'' intermediate between "pseudosuchians" and
saurischian Saurischia ( , meaning "reptile-hipped" from the Greek ' () meaning 'lizard' and ' () meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia), classified by their hip structure. Saurischia and Ornithis ...
dinosaurs. Modern authors now consider at least ''L. lilloensis'' to be firmly on the lineage of archosaurs leading to dinosaurs. However, the genus ''Lagosuchus'' is regarded by some to be
dubious Doubt is a mental state in which the mind remains suspended between two or more contradictory propositions, unable to be certain of any of them. Doubt on an emotional level is indecision between belief and disbelief. It may involve uncertainty ...
.
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at si ...
and
Andrea Arcucci Andrea is a given name which is common worldwide for both males and females, cognate to Andreas, Andrej and Andrew. Origin of the name The name derives from the Greek word ἀνήρ (''anēr''), genitive ἀνδρός (''andrós''), that re ...
considered ''L. talampayensis'' to be undiagnosable in a 1994 study, and reclassified ''L. lilloensis'' as a new genus, ''
Marasuchus ''Marasuchus'' (meaning "Mara crocodile") is a genus of basal dinosauriform archosaur which is possibly synonymous with '' Lagosuchus''. Both genera lived during the Late Triassic in what is now La Rioja Province, Argentina. ''Marasuchus'' conta ...
.'' In 2019, PULR 09, the holotype skeleton of ''L. talampayensis'', was redescribed by Federico Agnolin and
Martin Ezcurra Martin may refer to: Places * Martin City (disambiguation) * Martin County (disambiguation) * Martin Township (disambiguation) Antarctica * Martin Peninsula, Marie Byrd Land * Port Martin, Adelie Land * Point Martin, South Orkney Islands Austra ...
. They argued that the skeleton was not only diagnostic, but indistinguishable from specimens of ''Marasuchus lilloensis''. As a result, they supported the synonymy proposed by Bonaparte, referring specimens of ''Marasuchus lilloensis'' back to ''Lagosuchus talampayensis''. The Chañares Formation's age has been through much debate. It has traditionally been considered to belong to the
Ladinian The Ladinian is a stage and age in the Middle Triassic series or epoch. It spans the time between Ma and ~237 Ma (million years ago). The Ladinian was preceded by the Anisian and succeeded by the Carnian (part of the Upper or Late Triassic) ...
stage, the last 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 ...
.
Radiometric dating Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares ...
in 2016 has dated the main fossiliferous section of the formation to the early
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227 m ...
stage, near the start of the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch ...
.


Description

''Lagosuchus talampayensis'', in its most restricted form, can only be described based on the incomplete holotype skeleton. It was a lightly built archosaur, with long, slender legs and well-developed feet - features it shares with certain dinosaurs. With its short forelimbs, long shin bones, and narrow stance, it was likely an agile
biped Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
adapted for running. Gregory S. Paul inferred that ''Lagosuchus'' was one of the smallest Triassic archosaurs, with a weight of about 167 g, similar in size and ecology to a
weasel Weasels are mammals of the genus ''Mustela'' of the family Mustelidae. The genus ''Mustela'' includes the least weasels, polecats, stoats, ferrets and European mink. Members of this genus are small, active predators, with long and slend ...
or
ferret The ferret (''Mustela furo'') is a small, domesticated species belonging to the family Mustelidae. The ferret is most likely a domesticated form of the wild European polecat (''Mustela putorius''), evidenced by their interfertility. Other mus ...
.
Thomas Holtz Thomas Richard Holtz Jr. (born September 13, 1965) is an American vertebrate palaeontologist, author, and principal lecturer at the University of Maryland's Department of Geology. He has published extensively on the phylogeny, morphology, ecomor ...
estimated that ''Lagosuchus'' could have obtained a total length of 1.7 ft (51 cm) and a weight similar to that of a
pigeon Columbidae () is a bird family consisting of doves and pigeons. It is the only family in the order Columbiformes. These are stout-bodied birds with short necks and short slender bills that in some species feature fleshy ceres. They primarily ...
(50-500 g).Holtz, Thomas R. Jr. (2012) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages,'
Winter 2011 Appendix.
/ref> However, these estimates may have been based on specimens referred to ''Marasuchus,'' some of which were significantly larger than the ''Lagosuchus'' holotype.


Vertebrae and forelimbs

The
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal c ...
(trunk)
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 characteristi ...
had
centra Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the stores are all owned by individual franchisees. The chain has three different ...
about three times longer than tall, slightly more elongated than those referred to ''Marasuchus''. Yet the dorsals also had many traits in common with ''Marasuchus'', such as large excavations below the
transverse processes 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 ...
. In addition, they both have trapezoidal
neural spines 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 ...
with thickened upper edges which expand forwards and backwards to contact those of adjacent vertebrae. The contemporary silesaurid ''
Lewisuchus ''Lewisuchus'' is a genus of archosaur that lived during the Late Triassic (early Carnian). As a silesaurid dinosauriform, it was a member of the group of reptiles most commonly considered to be the closest relatives of dinosaurs (possibly true ...
'' has a similar neural spine morphology, but it possesses osteoderms on its neural spines, unlike ''Lagosuchus'' and ''Marasuchus''. The two sacral (hip) vertebrae had large and slightly tapering transverse processes. This is also the case for the first four
caudal Caudal may refer to: Anatomy * Caudal (anatomical term) (from Latin ''cauda''; tail), used to describe how close something is to the trailing end of an organism * Caudal artery, the portion of the dorsal aorta of a vertebrate that passes into the ...
(tail) vertebrae at the base of the tail. Further back, the caudals have much shorter transverse processes and more elongated centra, like those of ''Marasuchus''. The
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eith ...
(shoulder blade) was narrow, with a slightly expanded upper tip and a thick longitudinal ridge on its inner surface. 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 r ...
was also quite narrow, with a subtriangular deltopectoral crest in its upper part. The deltopectoral crest extends about 31% down the length of the humerus, making it somewhat less extensive than that of other avemetatarsalians (including ''Marasuchus''). The
radius In classical geometry, a radius (plural, : radii) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', ...
and
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
are thin, simple, and unusually short, only about 65% the length of the humerus. In contrast, ''Marasuchus'' has a larger ulna with a strong olecranon process. The forelimbs in general were much smaller than the hindlimbs.


Hip and hindlimbs

The
pelvis The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
(hip) is similar to that of ''Marasuchus'', with a thin pubis and a plate-like
ischium The ischium () form ...
which has a large ridge on its rear edge. The
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates ...
is elongated and has a slightly inturned femoral head. The femoral head is characteristically ‘globose’, with a strong projecting convex surface stretching up along its inner and upper portions. This trait is shared with ''Marasuchus'' and lagerpetids. The femur also possessed a knob-like anterior trochanter and a distinct
fourth trochanter The fourth trochanter is a shared characteristic common to archosaurs. It is a knob-like feature on the posterior-medial side of the middle of the femur shaft that serves as a muscle attachment, mainly for the '' musculus caudofemoralis longus'' ...
. The
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it conn ...
and
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity ...
were narrow and about 10% longer than the femur. Like other dinosauriforms, the tibia had a strong cnemial crest at the knee and a lateral groove near the ankle. The ankle has a small distal tarsal 3, a larger distal tarsal 4, and a rounded
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
, but the
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is the point of the hock. S ...
is missing from the fossil. The middle three
metatarsals 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 ...
are elongated, with metatarsal III as the longest. Metatarsal V is short and tapers to a sharp tip. Metatarsal I is also short, though still longer than V (unlike ''Marasuchus'').
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 slender, and an isolated pedal ungual (toe claw) is weakly curved and flattened sideways.


Palaeobiology


Metabolism

It is believed that ''Lagosuchus'' and ''Marasuchus'' were transitional between cold blooded reptiles and
warm blooded Warm-blooded is an informal term referring to animal species which can maintain a body temperature higher than their environment. In particular, homeothermic species maintain a stable body temperature by regulating metabolic processes. The onl ...
dinosaurs.


References

{{Taxonbar, from=Q132937 Prehistoric dinosauromorphs Late Triassic reptiles of South America Triassic Argentina Fossils of Argentina Chañares Formation Taxa described in 1971 Taxa named by Alfred Romer