''Carsosaurus'' is a genus of extinct amphibious reptiles, in the
mosasaur
Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Ancient Greek, Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains wer ...
superfamily, containing only the species ''Carsosaurus marchesetti''. It is known from a single individual from the
Komen Limestone of
Slovenia
Slovenia, officially the Republic of Slovenia, is a country in Central Europe. It borders Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the south and southeast, and a short (46.6 km) coastline within the Adriati ...
. The specimen is well-preserved, containing many different bones as well as some skin impressions and sternal
cartilage
Cartilage is a resilient and smooth type of connective tissue. Semi-transparent and non-porous, it is usually covered by a tough and fibrous membrane called perichondrium. In tetrapods, it covers and protects the ends of long bones at the joints ...
. While more remains are needed to be certain, it is generally thought to belong to the
Aigialosauridae
Aigialosauridae (from Ancient Greek, Greek, ''aigialos'', meaning "seashore", and ''sauros'', "lizard") is a family of Late Cretaceous semiaquatic Pythonomorpha, pythonomorph lizards closely related to the mosasaurs. Regarded by some paleontologi ...
. In life, it was an amphibious creature that spent most of its time on land, although its later relatives would become fully aquatic.
Discovery and naming
''Carsosaurus marchesetti'' was described from a single, mostly complete skeleton at the
Civico Museo di Storia Naturale di Trieste, uncovered from the
Karst Plateau
The Karst Plateau or the Karst region (, ), also locally called Karst, is a karst plateau region extending across the border of southwestern Slovenia and northeastern Italy.
It lies between the Vipava Valley, the low hills surrounding the val ...
near
Komen (modern-day
Slovenia
Slovenia, officially the Republic of Slovenia, is a country in Central Europe. It borders Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the south and southeast, and a short (46.6 km) coastline within the Adriati ...
) by
Andreas Kornhuber in 1893. He compared it to ''
Acteosaurus tommasinii'', as both were uncovered from the same area. There were numerous noticeable differences between the two, and Kornhuber concluded that they were not closely related at all: ''Acteosaurus tommasinii'' belonged to the family
Dolichosauridae
Dolichosauridae (from Latin, ''dolichos'' = "long" and Ancient Greek, Greek ''sauros''= lizard) is a family of Cretaceous Aquatic animal, aquatic lizards. They are widely considered to be the earliest and most primitive members of Mosasauria, tho ...
, while ''Carsosaurus'' more closely resembled a
monitor lizard
Monitor lizards are lizards in the genus ''Varanus,'' the only extant genus in the family Varanidae. They are native to Africa, Asia, and Oceania, and West African Nile monitor, one species is also found in south America as an invasive species. A ...
. For the "beautiful and memorable lizard of the Karst", in his words, he chose the genus name ''Carsosaurus''. The
specific epithet
In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
''marchesetti'' was in honour of the museum's director, Dr. Carlo de Marchesetti.
Description

''Carsosaurus'' is known only from a single specimen, currently housed at the Civico Museo di Storia Naturale di Trieste. It is missing the skull, anterior
cervical vertebrae
In tetrapods, cervical vertebrae (: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In saurop ...
, and much of the tail, but is otherwise very complete.
The total length of the parts preserved is . Kornhuber hypothesised that the mostly absent tail may have been twice as long as the body — — at . At its trunk, the skeleton is wide. There are only 3 cervical vertebrae present (the 3 furthest from the head), but in life there were likely 7–9. Similarly, only 12
caudal vertebrae
Caudal vertebrae are the vertebrae of the tail in many vertebrates. In birds, the last few caudal vertebrae fuse into the pygostyle, and in apes, including humans, the caudal vertebrae are fused into the coccyx.
In many reptiles, some of the caud ...
(located in the tail) are present in the fossil, however there may have been over 100. There are 5 pairs of
true ribs, and a presumed 3–4 additional pairs of
false ribs. Impressions of the
epidermis
The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermal layer provides a barrier to infection from environmental pathogens and regulates the ...
are preserved as well, which show rhombus-shaped scales, thickened at the margins.
There is also some
mineralised sternal
cartilage
Cartilage is a resilient and smooth type of connective tissue. Semi-transparent and non-porous, it is usually covered by a tough and fibrous membrane called perichondrium. In tetrapods, it covers and protects the ends of long bones at the joints ...
, which is large and shield-shaped.
Classification
Kornhuber initially classified ''Carsosaurus'' as a member of the
Aigialosauridae
Aigialosauridae (from Ancient Greek, Greek, ''aigialos'', meaning "seashore", and ''sauros'', "lizard") is a family of Late Cretaceous semiaquatic Pythonomorpha, pythonomorph lizards closely related to the mosasaurs. Regarded by some paleontologi ...
, due to its post-cranial features. A century later, in 1995,
a quantitative analysis including fossils of other species found in the intervening years supported this phylogenetic placement. Due to the lack of a skull, it is cannot be determined whether ''C. marchesetti'' might belong to the genus ''
Aigialosaurus''.
Because ''Aigialosaurus'' is the older name, ''Carsosaurus'' can therefore be considered 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 ...
''.
Paleobiology
Kornhuber considered ''Carsosaurus'' to be amphibious, although mainly land-dwelling. It long tail may have served as an emergency rudder, as well as a tool for defence, grasping, climbing, and propulsion. Its fossil contains what Kornhuber interpreted as the remains of many small fishes, lizards, and possibly amphibians, indicating that ''Carsosaurus'' was a hunter that consumed live prey whole.
However, in 2001, Caldwell and Michael S.Y. Lee proposed that these were not gut contents, but rather the remains of embryos.
Their positioning indicates that they would have been born tail-first, to lessen the chance of drowning, as this way their nostrils would emerge last. Since one is located in the
pelvis
The pelvis (: pelves or pelvises) is the lower part of an Anatomy, anatomical Trunk (anatomy), trunk, between the human abdomen, abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also c ...
, it is possible that the specimen died while giving birth. The ability of ''Carsosaurus'' and other early amphibious aigialosaurs to give live birth would have lessened their dependency on land, allowing their evolution into massive, fully aquatic
mosasaurs
Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains were discovered in ...
,
which would exist from 98 million years ago until the end of the
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 ...
, 65.5 million years ago.
Paleoecology
The single known specimen of ''Carsosaurus'' lived sometime between the
Cenomanian
The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
and
Turonian
The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...
, in the
Upper Cretaceous
The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cret ...
.
During the Cenomanian, much of the Komen area would have been covered by water, with a
tropical
The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
or
subtropical
The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones immediately to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Ge ...
climate. The upper levels were likely well-oxygenated, due to the large number of fish recovered, while the bottom would have been
anoxic
Anoxia means a total depletion in the level of oxygen, an extreme form of hypoxia or "low oxygen". The terms anoxia and hypoxia are used in various contexts:
* Anoxic waters, sea water, fresh water or groundwater that are depleted of dissolved ox ...
or dysoxic due to the scarcity of
benthic
The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
invertebrate
Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
fossils. Exposed land would not have been far, given the partially terrestrial habits of many species there, including the aigialosaurs.
Other taxa that lived or were likely to have lived in the Komen area during the Upper Cretaceous include ''
Komensaurus'' (another genus of aigialosaur),
Myctophidae (a family of fish whose modern members can be found in deep water
throughout the world), hard-shelled
molluscs
Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
,
crustaceans
Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of Arthropod, arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquat ...
,
conifers
Conifers () are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All e ...
, and
ammonites
Ammonoids are extinct, (typically) coiled-shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living octopuses, squid, and cuttlefish (which comprise the clade Coleoidea) than they are to nautiluses (family N ...
.
See also
*
List of mosasaurs
References
{{Taxonbar, from=Q16852514
Fossil taxa described in 1893
Late Cretaceous reptiles of Europe
Aigialosauridae