Simosuchus
Encyclopedia
Simosuchus is an extinct genus
of notosuchia
n crocodylomorph
s from the Late Cretaceous
of Madagascar
. It is named for its unusually short skull. Fully grown individuals were about 0.75 metres (2.5 ft) in length. The type species
is S. clarki, found from the Maevarano Formation
in Mahajanga Province
.
The teeth of S. clarki were shaped like clove
s (maple leaves), which coupled with its short and deep snout suggest it was not a carnivore
like most other crocodylomorphs. In fact, these features have led many palaeontologists to consider it an herbivore
.
, giving the genus its name, which means "pug-nosed crocodile" in Greek
. The shape of skulls differs considerably between specimens, with variation in ornamentation and bony projections. These differences may be indications of sexual dimorphism
. The front portion of the skull, or preorbital area, is angled downwards. Simosuchus likely held its head so that the preorbital area was angled about 45° from horizontal. The teeth line the front of the jaws and are clove-shaped. At the back of the skull, the occipital condyle
(which articulates with the neck vertebrae) is downturned. 45 autapomorphies, or features unique to Simosuchus, can be found in the skull alone.
In most respects, the postcranial skeleton of Simosuchus resembles that of other terrestrial crocodyliforms. There are several differences, however, that have been used to distinguish it from related forms. The scapula
is broad and tripartite (three-pronged). On its surface, there is a laterally directed prominence. The deltopectoral crest, a crest on the upper end of the humerus
, is small. The glenohumeral
condyle of the humerus, which connects to the pectoral girdle in the shoulder joint, has a distinctive rounded ellipsoid shape. The limbs are robust. The radius
and ulna
of the forearm fit tightly together. The front feet are small with large claws, and the back feet are also reduced in size. There is a small crest along the anterior edge of the femur
. On the pelvis
, the anterior process of the ischium is spur-like.
Most of the spinal column of Simosuchus is known. There are eight cervical vertebrae in the neck, at least fifteen dorsal vertebrae in the back, two sacral vertebrae
at the hip, and no more than twenty caudal vertebrae in the tail. The number of vertebrae in the tail is less than that of most crocodyliforms, giving Simosuchus a very short tail.
Like other crocodyliforms, Simosuchus was covered in bony plates called osteoderm
s. These form shields over the back, underside, and tail. Unusually among crocodyliforms, Simosuchus also has osteoderms covering much of the limbs. Osteoderms covering the back, tail, and limbs are light and porous, while the osteoderms covering the belly are plate-like and have an inner structure resembling spongy diploë
. Simosuchus has a tetraserial paravertebral shield over its back, meaning that there are four rows of tightly locking paramedial osteoderms (osteoderms to either side of the midline of the back). To either side of the shield, there are four rows of accessory parasagittal osteoderms. These accessory osteoderms tightly interlock with one another.
Buckley et al., 2000
Pol, 2003
Carvalho et al., 2004
Turner and Calvo, 2005
Simosuchus was first considered to be a basal
member of the clade
Notosuchia, and was often considered to be closely related to Uruguaysuchus
from the Late Cretaceous of Uruguay
and Malawisuchus
from the Early Cretaceous
of Malawi
. Later phylogenetic studies have placed it closer to the genus Libycosuchus
and in a more derived
position than some other notosuchians such as Uruguaysuchus. In its initial description by Buckley et al. (2000), Simosuchus was placed in the family
Notosuchidae
. Its sister taxon was Uruguaysuchus, and the two were allied with Malawisuchus
. These taxa were placed in Notosuchidae along with Libycosuchus and Notosuchus
. Most of the following phylogenetic analyses resulted in a similar placement of Simosuchus and other genera within Notosuchia. Turner and Calvo (2005) also found a clade including Simosuchus, Uruguaysuchus, and Malawisuchus in their study.
The phylogenetic analysis of Carvalho et al. (2004), based on different character values than previous studies, produced a very different relationship among Simosuchus and other notosuchians. Simosuchus, along with Uruguaysuchus and Comahuesuchus, were placed outside Notosuchia. Simosuchus was found to be the sister taxon of the Chinese
genus Chimaerasuchus in the family Chimaerasuchidae
. Like Simosuchus, Chimaerasuchus has a short snout and was probably herbivorous. Both genera were placed outside Notosuchia in the larger clade Gondwanasuchia. Uruguaysuchus, previously considered to be a basal notosuchian and a close relative of Simosuchus, was placed in its own family, Uruguaysuchidae, also outside Notosuchia. Malawisuchus was found to be a member of Peirosauroidea, specifically a member of the family Itasuchidae.
A fossorial
, or burrowing lifestyle, for Simosuchus has recently been proposed but is not widely agreed upon. Evidence for burrowing includes the robust limbs and short snout, which appears shovel-like. There are also areas on the skull that may have attached to strong neck muscles that would have been well suited for burrowing.
, the southern supercontinent
that existed throughout much of the Mesozoic and encompassed South America
, Africa
, India
, Australia
, and Antarctica. Libycosuchus, regarded as one of the closest relatives of Simosuchus, lived in Egypt
. Unresolved relationships among notosuchians along with an incomplete fossil record have made it difficult to determine the biogeographic origins of Simosuchus.
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of notosuchia
Notosuchia
Notosuchia is a suborder of primarily Gondwanan mesoeucrocodylian crocodylomorphs that lived during the Cretaceous. Fossils have been found from South America, Africa, and Asia...
n crocodylomorph
Crocodylomorpha
The Crocodylomorpha are an important group of archosaurs that include the crocodilians and their extinct relatives.During Mesozoic and early Tertiary times the Crocodylomorpha were far more diverse than they are now. Triassic forms were small, lightly built, active terrestrial animals. These were...
s from the Late Cretaceous
Late Cretaceous
The Late Cretaceous is the younger of two epochs into which the Cretaceous period is divided in the geologic timescale. Rock strata from this epoch form the Upper Cretaceous series...
of Madagascar
Madagascar
The Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...
. It is named for its unusually short skull. Fully grown individuals were about 0.75 metres (2.5 ft) in length. The type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...
is S. clarki, found from the Maevarano Formation
Maevarano Formation
The Maevarano Formation is an Upper Cretaceous sedimentary rock formation found in the Mahajanga Province of northwestern Madagascar. It is most likely Maastrichtian in age, and records a seasonal, semiarid environment with rivers that had greatly varying discharges...
in Mahajanga Province
Mahajanga Province
Mahajanga is a former province of Madagascar with an area of 150,023 km². It had a population of 1,896,000 . Its capital was Mahajanga.Except for Fianarantsoa, Mahajanga Province bordered all of the country's other provinces:...
.
The teeth of S. clarki were shaped like clove
Clove
Cloves are the aromatic dried flower buds of a tree in the family Myrtaceae. Cloves are native to the Maluku islands in Indonesia and used as a spice in cuisines all over the world...
s (maple leaves), which coupled with its short and deep snout suggest it was not a carnivore
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...
like most other crocodylomorphs. In fact, these features have led many palaeontologists to consider it an herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...
.
Description
Simosuchus was small, about 0.75 metres (2.5 ft) long based on the skeletons of mature individuals. In contrast to most other crocodyliforms, which have long, low skulls, Simosuchus has a distinctively short snout. The snout resembles that of a pugPug
The pug is a "toy" breed of dog with a wrinkly, short-muzzled face, and curled tail. The breed has a fine, glossy coat that comes in a variety of colors, and a compact square body with well-developed muscle. They have been described as multum in parvo , referring to the pug's personality and...
, giving the genus its name, which means "pug-nosed crocodile" in Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
. The shape of skulls differs considerably between specimens, with variation in ornamentation and bony projections. These differences may be indications of sexual dimorphism
Sexual dimorphism
Sexual dimorphism is a phenotypic difference between males and females of the same species. Examples of such differences include differences in morphology, ornamentation, and behavior.-Examples:-Ornamentation / coloration:...
. The front portion of the skull, or preorbital area, is angled downwards. Simosuchus likely held its head so that the preorbital area was angled about 45° from horizontal. The teeth line the front of the jaws and are clove-shaped. At the back of the skull, the occipital condyle
Occipital condyle
The occipital condyles are undersurface facets of the occipital bone in vertebrates, which function in articulation with the superior facets of the atlas vertebra....
(which articulates with the neck vertebrae) is downturned. 45 autapomorphies, or features unique to Simosuchus, can be found in the skull alone.
In most respects, the postcranial skeleton of Simosuchus resembles that of other terrestrial crocodyliforms. There are several differences, however, that have been used to distinguish it from related forms. The scapula
Scapula
In anatomy, the scapula , omo, or shoulder blade, is the bone that connects the humerus with the clavicle ....
is broad and tripartite (three-pronged). On its surface, there is a laterally directed prominence. The deltopectoral crest, a crest on the upper end of the humerus
Humerus
The humerus is a long bone in the arm or forelimb that runs from the shoulder to the elbow....
, is small. The glenohumeral
Glenohumeral joint
The glenohumeral joint, or shoulder joint, is a multiaxial synovial ball and socket joint and involves articulation between the glenoid fossa of the scapula and the head of the humerus...
condyle of the humerus, which connects to the pectoral girdle in the shoulder joint, has a distinctive rounded ellipsoid shape. The limbs are robust. The radius
Radius
In classical geometry, a radius of a circle or sphere is any line segment from its center to its perimeter. By extension, the radius of a circle or sphere is the length of any such segment, which is half the diameter. If the object does not have an obvious center, the term may refer to its...
and ulna
Ulna
The ulna is one of the two long bones in the forearm, the other being the radius. It is prismatic in form and runs parallel to the radius, which is shorter and smaller. In anatomical position The ulna is one of the two long bones in the forearm, the other being the radius. It is prismatic in form...
of the forearm fit tightly together. The front feet are small with large claws, and the back feet are also reduced in size. There is a small crest along the anterior edge of the femur
Femur
The femur , or thigh bone, is the most proximal bone of the leg in tetrapod vertebrates capable of walking or jumping, such as most land mammals, birds, many reptiles such as lizards, and amphibians such as frogs. In vertebrates with four legs such as dogs and horses, the femur is found only in...
. On the pelvis
Pelvis
In human anatomy, the pelvis is the lower part of the trunk, between the abdomen and the lower limbs .The pelvis includes several structures:...
, the anterior process of the ischium is spur-like.
Most of the spinal column of Simosuchus is known. There are eight cervical vertebrae in the neck, at least fifteen dorsal vertebrae in the back, two sacral vertebrae
Sacrum
In vertebrate anatomy the sacrum is a large, triangular bone at the base of the spine and at the upper and back part of the pelvic cavity, where it is inserted like a wedge between the two hip bones. Its upper part connects with the last lumbar vertebra, and bottom part with the coccyx...
at the hip, and no more than twenty caudal vertebrae in the tail. The number of vertebrae in the tail is less than that of most crocodyliforms, giving Simosuchus a very short tail.
Like other crocodyliforms, Simosuchus was covered in bony plates called osteoderm
Osteoderm
Osteoderms are bony deposits forming scales, plates or other structures in the dermal layers of the skin. Osteoderms are found in many groups of extant and extinct reptiles, including lizards, various groups of dinosaurs , crocodilians, phytosaurs, aetosaurs, placodonts, and hupehsuchians...
s. These form shields over the back, underside, and tail. Unusually among crocodyliforms, Simosuchus also has osteoderms covering much of the limbs. Osteoderms covering the back, tail, and limbs are light and porous, while the osteoderms covering the belly are plate-like and have an inner structure resembling spongy diploë
Diploë
Diploë refers to the spongy bone structure of the internal part of short, irregular, and flat bones.In the cranial bones, the layers of compact tissue are familiarly known as the tables of the skull; the outer one is thick and tough; the inner is thin, dense, and brittle, and hence is termed the...
. Simosuchus has a tetraserial paravertebral shield over its back, meaning that there are four rows of tightly locking paramedial osteoderms (osteoderms to either side of the midline of the back). To either side of the shield, there are four rows of accessory parasagittal osteoderms. These accessory osteoderms tightly interlock with one another.
History
The first specimen of Simosuchus clarki, which served as the basis for its initial description in 2000, included a complete skull and lower jaw, the front of the postcranial skeleton, and parts of the posterior postcranial skeleton. Five more specimens were later described, representing the majority of the skeleton. Many isolated teeth have also been found in the Mahajanga Basin. Most remains of Simosuchus were found as part of the Mahajanga Basin Project, directed by the Université d'Antananarivo and Stony Brook University. Material was usually found in clays that were part of flow deposits in the Anembalemba Member of the Maevarano Formation.Classification
Simosuchus was first considered to be a basal
Basal (phylogenetics)
In phylogenetics, a basal clade is the earliest clade to branch in a larger clade; it appears at the base of a cladogram.A basal group forms an outgroup to the rest of the clade, such as in the following example:...
member of the clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
Notosuchia, and was often considered to be closely related to Uruguaysuchus
Uruguaysuchus
Uruguaysuchus is an extinct genus of crocodylomorph from the Late Cretaceous of Uruguay. It was related to Simosuchus and Malawisuchus.-Sources:* A Pictorial Guide to Fossils by Gerard Ramon Case...
from the Late Cretaceous of Uruguay
Uruguay
Uruguay ,officially the Oriental Republic of Uruguay,sometimes the Eastern Republic of Uruguay; ) is a country in the southeastern part of South America. It is home to some 3.5 million people, of whom 1.8 million live in the capital Montevideo and its metropolitan area...
and Malawisuchus
Malawisuchus
Malawisuchus is an extinct genus of notosuchian mesoeucrocodylian from the Early Cretaceous of Malawi. It was described in 1997 by Elizabeth Gomani as a member of the family Notosuchidae. The type species is M. mwakasyungutiensis, referring to Mwakasyunguti, the area of northern Malawi where it...
from the Early Cretaceous
Early Cretaceous
The Early Cretaceous or the Lower Cretaceous , is the earlier or lower of the two major divisions of the Cretaceous...
of Malawi
Malawi
The Republic of Malawi is a landlocked country in southeast Africa that was formerly known as Nyasaland. It is bordered by Zambia to the northwest, Tanzania to the northeast, and Mozambique on the east, south and west. The country is separated from Tanzania and Mozambique by Lake Malawi. Its size...
. Later phylogenetic studies have placed it closer to the genus Libycosuchus
Libycosuchus
Libycosuchus is an extinct genus of North African crocodylomorph possibly related to Notosuchus. It was terrestrial, living approximately 95 million years ago in the Cenomanian stage of the Late Cretaceous...
and in a more derived
Derived
In phylogenetics, a derived trait is a trait that is present in an organism, but was absent in the last common ancestor of the group being considered. This may also refer to structures that are not present in an organism, but were present in its ancestors, i.e. traits that have undergone secondary...
position than some other notosuchians such as Uruguaysuchus. In its initial description by Buckley et al. (2000), Simosuchus was placed in the family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
Notosuchidae
Notosuchidae
Notosuchidae is a Gondwanan family of notosuchians. They were small-bodied terrestrial crocodyliforms that lived during the Late Cretaceous.-Genera:*Notosuchus: genus from the Coniacian - Santonian of Argentina....
. Its sister taxon was Uruguaysuchus, and the two were allied with Malawisuchus
Malawisuchus
Malawisuchus is an extinct genus of notosuchian mesoeucrocodylian from the Early Cretaceous of Malawi. It was described in 1997 by Elizabeth Gomani as a member of the family Notosuchidae. The type species is M. mwakasyungutiensis, referring to Mwakasyunguti, the area of northern Malawi where it...
. These taxa were placed in Notosuchidae along with Libycosuchus and Notosuchus
Notosuchus
Notosuchus is an extinct genus of South American notosuchian crocodylomorph. It was terrestrial, living approximately 85 million years ago in the Coniacian or Santonian stages of the Late Cretaceous. Remains have been found in the Bajo de la Carpa Formation in Patagonia, Argentina. First named in...
. Most of the following phylogenetic analyses resulted in a similar placement of Simosuchus and other genera within Notosuchia. Turner and Calvo (2005) also found a clade including Simosuchus, Uruguaysuchus, and Malawisuchus in their study.
The phylogenetic analysis of Carvalho et al. (2004), based on different character values than previous studies, produced a very different relationship among Simosuchus and other notosuchians. Simosuchus, along with Uruguaysuchus and Comahuesuchus, were placed outside Notosuchia. Simosuchus was found to be the sister taxon of the Chinese
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...
genus Chimaerasuchus in the family Chimaerasuchidae
Chimaerasuchidae
Chimaerasuchidae is a family of mesoeucrocodylians. It was erected as a clade in 2004 by Carvalho et al and included Chimaerasuchus from the Early Cretaceous of China and Simosuchus from the Late Cretaceous of Madagascar...
. Like Simosuchus, Chimaerasuchus has a short snout and was probably herbivorous. Both genera were placed outside Notosuchia in the larger clade Gondwanasuchia. Uruguaysuchus, previously considered to be a basal notosuchian and a close relative of Simosuchus, was placed in its own family, Uruguaysuchidae, also outside Notosuchia. Malawisuchus was found to be a member of Peirosauroidea, specifically a member of the family Itasuchidae.
Paleobiology
Simosuchus, like other notosuchians, was fully terrestrial. Features of the skull such as the clove-shaped teeth strongly suggest terrestrial herbivory. The short tail would have had little use in swimming. The osteoderm shield was inflexible, restricting lateral movement in Simosuchus as a possible adaptation to an entirely terrestrial lifestyle. Robust legs are also consistent with terrestrial locomotion. The deltopectoral crest on the humerus and the anterior crest on the femur served as attachment points for strong limb muscles. The hindlimbs of Simosuchus were semierect, unlike the fully erect posture of most other notosuchians. Due to its posture, Simosuchus probably did not run.A fossorial
Fossorial
A fossorial organism is one that is adapted to digging and life underground such as the badger, the naked mole rat, and the mole salamanders Ambystomatidae...
, or burrowing lifestyle, for Simosuchus has recently been proposed but is not widely agreed upon. Evidence for burrowing includes the robust limbs and short snout, which appears shovel-like. There are also areas on the skull that may have attached to strong neck muscles that would have been well suited for burrowing.
Paleobiogeography
It is unknown how Simosuchus arrived in Madagascar. A similar crocodyliform, Araripesuchus tsangatsangana, is also known from the Maevarano Formation, but its relation to Simosuchus is unclear. It has been classified as both a notosuchian and a basal neosuchian in various phylogenetic analyses. Nearly all notosuchians are known from GondwanaGondwana
In paleogeography, Gondwana , originally Gondwanaland, was the southernmost of two supercontinents that later became parts of the Pangaea supercontinent. It existed from approximately 510 to 180 million years ago . Gondwana is believed to have sutured between ca. 570 and 510 Mya,...
, the southern supercontinent
Supercontinent
In geology, a supercontinent is a landmass comprising more than one continental core, or craton. The assembly of cratons and accreted terranes that form Eurasia qualifies as a supercontinent today.-History:...
that existed throughout much of the Mesozoic and encompassed South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...
, Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...
, India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...
, Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...
, and Antarctica. Libycosuchus, regarded as one of the closest relatives of Simosuchus, lived in Egypt
Egypt
Egypt , officially the Arab Republic of Egypt, Arabic: , is a country mainly in North Africa, with the Sinai Peninsula forming a land bridge in Southwest Asia. Egypt is thus a transcontinental country, and a major power in Africa, the Mediterranean Basin, the Middle East and the Muslim world...
. Unresolved relationships among notosuchians along with an incomplete fossil record have made it difficult to determine the biogeographic origins of Simosuchus.
External links
- Mikko's Phylogenic Archives http://www.fmnh.helsinki.fi/users/haaramo/Metazoa/Deuterostoma/Chordata/Archosauria/Crocodylia/Crocodyliformes.htm
- Digimorph Entry http://www.digimorph.org/specimens/Simosuchus_clarki/