Ectenosaurus
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''Ectenosaurus'' is an extinct
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 marine lizard belonging to 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 ...
family. It is classified as part of the
Plioplatecarpinae Plioplatecarpinae is a subfamily of mosasaurs, a diverse group of Late Cretaceous marine Squamata, squamates. Members of the subfamily are informally and collectively known as "plioplatecarpines" and have been recovered from all continents, thoug ...
subfamily alongside genera like ''
Angolasaurus ''Angolasaurus'' ("Angola lizard") is an extinct genus of mosasaur. Definite remains from this genus have been recovered from the Turonian and Coniacian of Angola, and possibly the Coniacian of the United States, the Turonian of Brazil, and the M ...
'' and ''
Platecarpus ''Platecarpus'' ("oar wrist") is an extinct genus of aquatic lizards belonging to the mosasaur family, living around 84–81 million years ago during the middle Santonian to early Campanian, of the Late Cretaceous period. Fossils have been found ...
''. ''Ectenosaurus'' is known from the
Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya ( million years ago) and 83.6 ± 0.7 m ...
and
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
of
Kansas Kansas ( ) is a landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the west. Kansas is named a ...
,
Alabama Alabama ( ) is a U.S. state, state in the Southeastern United States, Southeastern and Deep South, Deep Southern regions of the United States. It borders Tennessee to the north, Georgia (U.S. state), Georgia to the east, Florida and the Gu ...
, and
Texas Texas ( , ; or ) is the most populous U.S. state, state in the South Central United States, South Central region of the United States. It borders Louisiana to the east, Arkansas to the northeast, Oklahoma to the north, New Mexico to the we ...
. The generic name means "drawn-out lizard", 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 ...
''ectenes'' ("drawn-out") and Greek ''sauros'' ("lizard") referencing the elongated muzzle.


Description

With the preserved skull about long, ''Ectenosaurus'' is estimated to have reached in length. It was a rare genus of
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 ...
with several unique characteristics that clearly separate it from other mosasaur genera. The most prominent of these features is its elongated jaws, elongated in a similar vein to other mosasaurs with elongated jaws, such as ''
Plotosaurus ''Plotosaurus'' ("swimmer lizard") is an extinct genus of large mosasaurs which lived during the Upper Cretaceous (Maastrichtian) in what is now North America. The taxon was initially described by University of California, Berkeley, Berkeley pale ...
'' and ''
Pluridens ''Pluridens'' ("many teeth") is an extinct genus of marine lizard belonging to the Mosasauridae. ''Pluridens'' is placed in the subfamily Halisaurinae with the genera ''Phosphorosaurus'', ''Eonatator'' and ''Halisaurus''.Konishi, Takuya; Caldwell ...
''. Dale Allan Russell (1967) considered the form of the teeth, the shape of the frontal and the large suprastapedial process of the quadrate as evidence of a close relation between ''Ectenosaurus'' and ''
Platecarpus ''Platecarpus'' ("oar wrist") is an extinct genus of aquatic lizards belonging to the mosasaur family, living around 84–81 million years ago during the middle Santonian to early Campanian, of the Late Cretaceous period. Fossils have been found ...
''. He separated ''Ectenosaurus'' from ''Platecarpus'' based on the elongated snout, the exclusion of the prefrontals from the narial borders and the fusion of the supra- and infrastepedial processes.


Scales and locomotion

The specimen FHSM VP-401 preserve significantly comprehensive skin impressions from ''Ectenosaurus'', which makes it possible to draw conclusions not only about mosasaur integument at large but also about mosasaur movement and propulsion. The scales are considerably smaller in size (2.7×2.0 mm) than those found in the famed LACM 128319 specimen of ''
Platecarpus ''Platecarpus'' ("oar wrist") is an extinct genus of aquatic lizards belonging to the mosasaur family, living around 84–81 million years ago during the middle Santonian to early Campanian, of the Late Cretaceous period. Fossils have been found ...
'' (3.8×4.4 mm), despite the animals being of similar sizes. The combination of small and firmly anchored body scales as well as a complex meshwork of alternating crossed-helical and longitudinal fiber bundles suggest that the anterior torso of ''Ectenosaurus'' was reasonably stiff. This also suggests that this section of the body was quite rigid during locomotion, and that the main form of propulsion would have to have been done by the tail (likely possessing a tail fin like other mosasaur species), and that it could not move by undulating its entire body like
snake Snakes are elongated limbless reptiles of the suborder Serpentes (). Cladistically squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales much like other members of the group. Many species of snakes have s ...
s do, a previously popular view of mosasaur locomotion.


History of discovery

''Ectenosaurus'' was originally described as a species of ''
Platecarpus ''Platecarpus'' ("oar wrist") is an extinct genus of aquatic lizards belonging to the mosasaur family, living around 84–81 million years ago during the middle Santonian to early Campanian, of the Late Cretaceous period. Fossils have been found ...
'', ''P. clidastoides'', in 1894 by
John Campbell Merriam John Campbell Merriam (October 20, 1869 – October 30, 1945) was an American paleontologist, educator, and conservationist. The first vertebrate paleontologist on the West Coast of the United States, he is best known for his taxonomy of ve ...
. The type specimen was collected by
Charles Hazelius Sternberg Charles Hazelius Sternberg (June 15, 1850 – July 20, 1943) was an American fossil collector and paleontology, paleontologist. He was active in both fields from 1876 to 1928, and collected fossils for Edward Drinker Cope and Othniel C. Marsh, ...
or Georg Bauer from
Logan County Logan County is the name of ten current counties and one former county in the United States: * Logan County, Arkansas * Logan County, Colorado * Logan County, Idaho (1889–1895) * Logan County, Illinois * Logan County, Kansas * Logan County ...
in
Kansas Kansas ( ) is a landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the west. Kansas is named a ...
and was housed in the Bayerische Staatssammlung für Paläontologie in Munich, where it was likely destroyed in 1944 during the
Second World War World War II or the Second World War (1 September 1939 – 2 September 1945) was a World war, global conflict between two coalitions: the Allies of World War II, Allies and the Axis powers. World War II by country, Nearly all of the wo ...
. A second specimen (which was also much better preserved) was discovered by
George Fryer Sternberg George Fryer Sternberg (1883 – 23 October 1969) was an American paleontologist best known for his discovery in Gove County, Kansas of the "fish-within-a-fish" of ''Xiphactinus audax'' with a recently eaten ''Gillicus arcuatus'' within its stom ...
in 1953, which he initially identified as a '' Clidastes velox''. The specimen, formerly catalogued as GFS 109-53, was about in length and largely articulated, though the tail and rear limbs were missing due to erosion. This specimen was then exhibited at the Sternberg Memorial Museum on the campus of
Fort Hays State University Fort Hays State University (FHSU) is a public university in Hays, Kansas, United States. It is the largest university in western Kansas, and the fourth largest of the six state universities governed by the Kansas Board of Regents, with a total ...
from its discovery until 1999 when the museum was closed and moved. Since the museum moved, the specimen remains in storage, where it is catalogued as FHSM VP-401. It was examined in 1963 by Dale A. Russell who determined that it represented a ''Platecarpus clidastoides'' and not a ''Clidastes velox''. In his ''Systematics and Morphology of American Mosasaurs'' (1967) he re-described the species as part of a new genus, ''Ectenosaurus''. Russell also assigned several additional specimens to ''Ectenosaurus clidastoides'', namely YPM 4671, 4672, 4673, and 4674 in the
Yale Peabody Museum The Peabody Museum of Natural History at Yale University (also known as the Yale Peabody Museum of Natural History or the Yale Peabody Museum) is one of the oldest, largest, and most prolific university natural history museums in the world. It ...
. In 2021, Alexander Willman, Takuya Konishi, and Michael Caldwell designated FHSM VP-401 as the
neotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to anchor or centralizes ...
of ''E. clidastoides'', as a replacement for the type specimen. They also named another mosasaur specimen discovered in Logan County during the 1970s, catalogued as FHSM VP-5515, as a new species of ''Ectenosaurus'', ''Ectenosaurus everhartorum''. This species was named after marine reptile researchers Mike Everhart and Pamela Everhart. Since they could not be differentiated, the YPM specimens were removed by Willman and colleagues from ''E. clidastoides'' and considered as ''Ectenosaurus'' sp. In 2023, Caitlín R. Kiernan and Jun A. Ebersole named two new species of the genus: ''E. tlemonectes'' from the Niobrara of Kansas and ''E. shannoni'' from the Mooreville Chalk of Alabama.


Classification

''Ectenosaurus'' has been seen as a plioplatecarpine for most of the time since its discovery, partly due to long having been classified as a species of ''
Platecarpus ''Platecarpus'' ("oar wrist") is an extinct genus of aquatic lizards belonging to the mosasaur family, living around 84–81 million years ago during the middle Santonian to early Campanian, of the Late Cretaceous period. Fossils have been found ...
''. Some analyses recover it as a mosasaurine however, sharing close relations with ''
Prognathodon ''Prognathodon'' is an extinct genus of marine lizard belonging to the mosasaur family. It is classified as part of the Mosasaurinae subfamily, alongside genera like ''Mosasaurus'' and ''Clidastes''. ''Prognathodon'' has been recovered from depos ...
''. The
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 ...
below follows Simões et al. (2017), collapsed to only display the Plioplatecarpinae, and shows ''Ectenosaurus'' in relation to other plioplatecarpines.


Palaeoecology

Stable isotope analysis indicates that ''Ectenosaurus'' was one of the few plioplatecarpines that was not restricted to nearshore marine habitats.


References

{{Taxonbar, from=Q3335497 Mosasaurs of North America Fossil taxa described in 1967 Taxa named by Dale Russell Mosasaurids Santonian genus first appearances Campanian genus extinctions