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
caiman
A caiman ( (also spelled cayman) from Taíno language, Taíno ''kaiman'') is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family (biology), family, the other being alligators. ...
from modern day
Argentina
Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
that lived during the
Paleocene
The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
epoch (
Danian
The Danian is the oldest age or lowest stage of the Paleocene Epoch or Series, of the Paleogene Period or System, and of the Cenozoic Era or Erathem. The beginning of the Danian (and the end of the preceding Maastrichtian) is at the Cretac ...
or possibly
Selandian
The Selandian is a stage in the Paleocene. It spans the time between . It is preceded by the Danian and followed by the Thanetian. Sometimes the Paleocene is subdivided in subepochs, in which the Selandian forms the "middle Paleocene".
Stratig ...
age, about 60 million years ago). It inhabited the marine or fluvio-lacustrine environment of the Patagonian
Salamanca Formation
The Salamanca Formation is a geologic formation in the Golfo San Jorge Basin of central Patagonia that yields well-preserved, well-dated fossils from the early Paleocene. Studies of these fossils are providing new data on plant and animal diversit ...
.
History and naming
The fossil remains of ''Necrosuchus'' were unearthed on April 3, 1931 during the First Scarritt Expedition in the Argentinian
Salamanca Formation
The Salamanca Formation is a geologic formation in the Golfo San Jorge Basin of central Patagonia that yields well-preserved, well-dated fossils from the early Paleocene. Studies of these fossils are providing new data on plant and animal diversit ...
and presented to American paleontologist Charles C. Mook. Mook however, busy with other research, was unable to describe the specimen himself. As an initial description was deemed vital to the works of several student researches at the time, Mook handed the specimen back to George G. Simpson for study. He eventually published his preliminary description in 1937, writing that his research "only carries this study as far as necessary" in the hopes of a more detailed description being later published by Mook himself. However this would not come to be and the postcranial was not described until 2012. Holotype AMNH 3219 consists of much of the postcranial anatomy alongside a right
dentary
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 ...
and assorted cranial fragments not recognized as such in 1937. Although other material from Paleocene Patagonia shares similarities with ''Necrosuchus'', none of it can be safely referred to the genus itself.
Simpson explains the meaning of the generic name as being a compound word of "nekros" meaning "dead," and "suchus" meaning crocodile. Simpson described how "a well-meaning lady asked us if it were dead", which inspired his choice of name. The specific name refers to the Las Violetas site: ''ion'' is "violet" in
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 ...
.
Description
The dentary teeth of ''Necrosuchus'' are more or less evenly spaced throughout the holotype, only the 12th and 13th teeth lying closer together than the others (however even then without being in direct contact). The upper margin of the jaw is notably concave between the 1st and 4th tooth and between the 4th and 13th, after which it remains straight. The teeth show very little differentiation from one another in morphology, however the difference in tooth size is striking and almost equal in its degree to that of extant genus ''
Caiman
A caiman ( (also spelled cayman) from Taíno language, Taíno ''kaiman'') is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family (biology), family, the other being alligators. ...
''. Although the degree of enlargement is similar, the distribution is not the same and more similar to that seen in ''Leidyosuchus'' (many of the species used for comparison being now considered '' Borealosuchus'') according to Simpson. The first two teeth are of the same size, followed by a decrease in size for tooth three. The 4th dentary tooth is the largest of the lower jaw and followed by a series of noticeably smaller teeth (a similar condition is seen in extant caimans). Unlike in extant taxa however, the 13th tooth is enlarged to the point that it almost rivals the 4th dentary tooth in size, and the transition in size before and after it is much less abrupt. The 13th tooth's status of second largerst tooth in the dentary differentiates ''Necrosuchus'' from all other caimans other than '' Purussaurus brasiliensis'' and some species of '' Paleosuchus''. ''Necrosuchus'' differs from ''Purussaurus'' through its slender mandibular ramus as well as the size of the first four dentary and from ''Paleosuchus'' through the anatomy of the atlantal ribs and the lack of mediolaterall compression on the posterior dentary teeth.
The postcranial remains include 4 dorsal vertebrae that were preserved in articulation. They are procoelous with rectangular neural spines and lacking their transverse processes due to preservation. At some point in the past, the vertebrae had been identified as dorsals 5 to 8, but the basis for this assignment is not known. However, the study by Brochu confirms that this assumption was correct based on the anatomy of the hypapophyseal keel. The last two dorsals are also preserved in articulation with the sacral vertebrae. The sutures of the dorsal and sacral neurocentrals are unfused and still open, suggesting that the animal was not yet fully grown by the time of its death. The
pectoral girdle
The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans, it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists o ...
is largely preserved, with parts of both
scapula
The scapula (: 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 either side ...
humeri
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 and ulna, and consists of three sections. The humeral upper extremity consists of ...
,
ulna
The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
e and
radii
In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is the line segment or ...
as well as one radiale. They largely resemble forelimbs of modern crocodilians. Both pelvic girdles are preserved, although one of the ilia is distorted, but the corresponding bone of the other side preserves its natural state. The blades of the pubis are less symmetrical than in modern crocs. The hindlimbs preserve the
femur
The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg.
The Femo ...
,
tibia
The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
, parts of the
fibula
The fibula (: fibulae or fibulas) 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. ...
and various foot bones. Osteoderms were preserved but not in articulation.
Phylogeny
Early analysis of ''Necrosuchus'' believed it to be a close relative of ''
Leidyosuchus
''Leidyosuchus'' (meaning " Leidy's crocodile") is an extinct genus of eusuchian, either an alligatoroid crocodilian or a stem-group crocodilian, from the Late Cretaceous of Alberta. It was named in 1907 by Lawrence Lambe, and the type species ...
'', which at the time was believed to be a crocodylid. Later research showed that ''Necrosuchus'' shows derived features of alligatoroids and more precisely caimanines. By the time Brochu described the postcrania of ''Necrosuchus'', many species of ''Leidyosuchus'' originally used for comparison were reclassified as species of '' Borealosuchus''. The postcranial material however differs significantly from ''Borealosuchus'' and shows further affinities with caimans. Following a taxonomic revision and amended diagnosis published in 2020, Cicade and colleagues recovered ''Necrosuchus'' as a derived member of
Caimaninae
A caiman ( (also spelled cayman) from Taíno ''kaiman'') is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family, the other being alligators. Caimans are native to Central and ...
nested within a large
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 tree ...
with most extant non-'' paleosuchus'' caimans and ''
Purussaurus
''Purussaurus'' is an extinct genus of giant caiman that lived in South America during the Miocene epoch, from the Friasian to the Huayquerian in the SALMA classification. It is known from skull material found in the Brazilian and Peruvian Am ...
''. This polytomy was a direct result of the inclusion of ''Necrosuchus'' and '' Centenariosuchus'', and their removal resulted in a better resolved
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 ...
tree. Following a Pcr Prune analysis, two alternative placements of ''Necrosuchus'' recover it as either basal to the clade
Jacarea
Jacarea is a clade of caimans within the subfamily Caimaninae. Jacarea was first named by Norell in 1988 to include the extant species within the genera ''Caiman'' and ''Melanosuchus'', while excluding the dwarf caiman genus '' Paleosuchus''. I ...
or as basal to a clade comprising Jacarea and '' Acresuchus'' + ''
Purussaurus
''Purussaurus'' is an extinct genus of giant caiman that lived in South America during the Miocene epoch, from the Friasian to the Huayquerian in the SALMA classification. It is known from skull material found in the Brazilian and Peruvian Am ...
''. The below
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 ...
shows the results of the strict consensus phylogenetic analysis.
The following morphological
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 ...
cladogram published by Rio and colleagues recovered a more resolved tree, with ''Necrosuchus'' clading alongside '' Bottosaurus'' from
Cretaceous
The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
-
Paleocene
The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
North America and ''
Protocaiman
''Protocaiman'' is a caimanine genus of crocodylian first scientifically described, described in 2018. The type species ''Protocaiman peligrensis'' was discovered in Argentina's Salamanca Formation, and lived in Patagonia during the Paleocene epo ...