Selmasaurus
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''Selmasaurus'' 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 ...
''. Two species are known, ''S. russelli'' and ''S. johnsoni''; both are exclusively known from
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 ...
deposits in the
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
. ''Selmasaurus'' is unique among the mosasaurs in that its skull is unusually akinetic, meaning that it is incapable of widening to swallow larger prey. Most mosasaurs have skulls which possess "coupled kinesis" (mesokinesis and streptostyly), that is, parts of the jaw can open widely to accommodate large prey.


History of discovery

First recognized by
geologist A geologist is a scientist who studies the structure, composition, and History of Earth, history of Earth. Geologists incorporate techniques from physics, chemistry, biology, mathematics, and geography to perform research in the Field research, ...
Samuel Wayne Shannon in his 1975 Master's thesis, "Selected Alabama Mosasaurs", the taxon remained a ''
nomen nudum In Taxonomy (biology), taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published ...
'' until it was officially described in 1988 in an article coauthored by Wright. The type specimen, formerly reposited at the Geological Survey of Alabama and cataloged as GSATC 221, was transferred in 2005 to the
Alabama Museum of Natural History 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 ...
(Tuscaloosa). The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
of this genus consists of a very well preserved but incomplete and disarticulated
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
, the left atlantal neural arch, atlas centrum, and a single neural arch from a cervical vertebra. Preserved skull elements include the frontal, parietal, left ectopterygoid, left jugal, supratemporals, basioccipital and basisphenoid, and quadrates. The species was named in honor of paleontologist Dale A. Russell, for his extensive work on mosasaurs. The holotype and only known specimen of ''S. russelli'' was collected from an unknown location in western
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 for decades, uncertainty surrounded the precise
stratigraphic Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithost ...
horizon from which the specimen had been recovered. Then in 1998, Caitlín R. Kiernan extracted chalky matrix from the basilar canal of the basioccipital and identified calcareous nanoplankton that indicated GSATC 221 had originated from basal
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 ...
beds within the lower unnamed member of the
Mooreville Chalk Formation The Mooreville Chalk is a geological formation in North America, within the U.S. states of Alabama and Mississippi, which were part of the subcontinent of Appalachia. The strata date back to the early Santonian to the early Campanian stage of th ...
(
Selma Group The Selma Group is a geological formation in North America, within the U.S. states of Alabama, Mississippi, and Tennessee. The strata date from the Santonian to the Maastrichtian stages of the Late Cretaceous. The group is composed of, in ascend ...
). In her study of Alabama mosasaur
biostratigraphy Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. "Biostratigraphy." ''Oxford Reference: Dictionary of Biology ...
, Kiernan placed ''S. russelli'' within the ''
Clidastes ''Clidastes'' 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 ''Prognathodon''. ''Clidastes'' is known from deposits ranging i ...
'' Acme Zone, though it was the rarest element in the fauna, accounting for only 0.3% of the
biozone In biostratigraphy, biostratigraphic units or biozones are intervals of geological strata that are defined on the basis of their characteristic fossil taxon, taxa, as opposed to a Lithostratigraphy, lithostratigraphic unit which is defined by the ...
's assemblage (one specimen). A remarkably well preserved and nearly complete ''Selmasaurus'' skull and partial postcranial skeleton was discovered by Steve Johnson and family in 1996, 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 ...
or Campanian marine horizon in the
Niobrara Formation The Niobrara Formation , also called the Niobrara Chalk, is a geologic formation in North America that was deposited between 87 and 82 million years ago during the Coniacian, Santonian, and Campanian stages of the Late Cretaceous. It is com ...
of
Niobrara Chalk The Niobrara Formation , also called the Niobrara Chalk, is a Formation (stratigraphy), geologic formation in North America that was deposited between 87 and 82 million years ago during the Coniacian, Santonian, and Campanian stages of the La ...
, western
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 ...
. Recovered in 1997 and donated to the Sternberg Museum of Natural History in Hays, Kansas, in 2001, the remains were determined to be a new species of ''Selmasaurus'' in 2008 after over a decade of study by Polcyn and Everhart. Named ''S. johnsoni'' after its discoverer, the skull is one of the most complete mosasaur skulls recovered and thus provides new anatomical information for ''Selmasaurus'', a better understanding of plioplatecarpine ingroup relationships, extends the geographic and temporal range of the genus, and documents further diversity within
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 ...
. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
and the only known specimen is housed at the Sternberg Museum of Natural History under catalog number FHSM-VP-13910.


Description

''Selmasaurus'' was a small predatory mosasaur at approximately 3–5 meters in length. It possesses a relatively low number of teeth for a mosasaur, the lowest of any known species at the time of its discovery. Originally classified as a plioplatecarpine mosasaur, it differs from all other plioplatecarpine mosasaurs in several respects, listed below as stated by Polcyn and Everhart (
2008 2008 was designated as: *International Year of Languages *International Year of Planet Earth *International Year of the Potato *International Year of Sanitation The Great Recession, a worldwide recession which began in 2007, continued throu ...
):Polcyn, M. J., and Everhart, M. J., 2008, Description and phylogenetic analysis of a new species of ''Selmasaurus'' (Mosasauridae: Plioplatecarpinae) from the Niobrara Chalk of western Kansas: In: Proceedings of the Second Mosasaur Meeting, edited by Everhart, M. J, Fort Hays Studies, Special Issue number 3, p. 13-28.


Classification

Wright and Shannon classified ''Selmasaurus'' as a member of the mosasaur subfamily
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 ...
, which also includes the genera ''Platecarpus'', ''Plioplatecarpus'', and ''Ectenosaurus'', largely on the "basis of the mode of circulation through the basicarnium." The genus may be most closely related to ''Ectenosaurus'', though it possesses a much shorter, stouter skull. Additional specimens would greatly expand our understanding of ''Selmasaurus russelli''. 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 the most resolved topology from a 2011 phylogenetic analysis 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 ...
by paleontologists Takuya Konishi and Michael W. Caldwell. In their 2024 description of the Moroccan plioplatecarpine ''
Khinjaria ''Khinjaria'' (meaning "dagger") is an extinct genus of plioplatecarpine mosasaurid from the Late Cretaceous Ouled Abdoun Basin of Morocco. The genus contains a single species, ''K. acuta'', known from a partial skull and vertebra. ''Khinjaria ...
'', Longrich ''et al.'' used
phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
to recover it within a monophyletic clade of non-plioplatecarpin plioplatecarpine mosasaurids. They named this clade—also containing '' Gavialimimus'', ''
Goronyosaurus ''Goronyosaurus'' is an extinct genus of marine lizard belonging to the mosasaur family. Fossils of ''Goronyosaurus'' are exclusively known from the Late Maastrichtian of the Iullemmeden Basin in West Africa, specifically the Dukamaje Formation o ...
'', and ''Selmasaurus''—the Selmasaurini. ''Selmasaurus johnsoni'', which is older and may be more primitive, was found in a more basal position than ''S. russelli''. The results of their analyses are shown in 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:


Palaeoecology

According to stable isotope analysis, ''S. johnsoni'' was one of the few plioplatecarpines that was not restricted to nearshore marine environments.


References

{{Taxonbar, from=Q7211269 Fossil taxa described in 1988 Mosasaurs of North America Mooreville Chalk Mosasaurids