The Halisaurinae are a

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...

mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on th ...

s, a group of Late

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...

marine lizards. They were small to medium-sized, ranging from just under 3 meters in ''Eonatator sternbergi'' to as much as 8 or 9 meters in ''Pluridens serpentis''. They tended to have relatively slender jaws and small, numerous teeth, suggesting a diet of small fish and other prey. Although the skeleton is primitive compared to other Mosasauridae in many respects, halisaurines had the distinctive hypocercal tail of other mosasaurids suggesting good swimming ability, and they persisted alongside other mosasaurs until the end of the Cretaceous. The earliest known remains of halisaurines occur in rocks of

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 mya. The ...

age and the subfamily persists until the latest

Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...

. Halisaurines are known from

North North is one of the four compass points or cardinal directions. It is the opposite of south and is perpendicular to east and west. ''North'' is a noun, adjective, or adverb indicating Direction (geometry), direction or geography. Etymology T ...

and

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...

Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...

and

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

, indicating a more or less global distribution in the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger 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 ''creta'', the ...

. Four genera are currently recognized: ''

Eonatator ''Eonatator'' is an extinct genus of marine lizard belonging to the mosasaur family. It is a close relative of ''Halisaurus'', and part of the same subfamily, the Halisaurinae. It is known from the Late Cretaceous of North America, Colombia and S ...

'', ''

Halisaurus ''Halisaurus'' is an extinct genus of marine reptile belonging to the mosasaur family. The holotype, consisting of an angular and a basicranium fragment discovered near Hornerstown, New Jersey, already revealed a relatively unique combination ...

'', '' Phosphorosaurus'' and '' Pluridens''.

Description

The Halisaurinae exhibit several characteristics that suggest they are more primitive than mosasaurs of more derived subfamilies. In particular, their flippers are poorly differentiated, meaning that while better adapted to life in oceans than plesiopedal mosasaur genera like ''

Dallasaurus ''Dallasaurus'' ("Dallas lizard") is a basal mosasauroid from the Upper Cretaceous of North America. Along with ''Russellosaurus'', ''Dallasaurus'' is one of the two oldest mosasauroid taxa currently known from North America. It is also the small ...

'' and ''

Tethysaurus ''Tethysaurus'' is an extinct genus of tethysaurine mosasauroid from the Early Turonian (Late Cretaceous) period. The only species is ''Tethysaurus nopcsai''. Discovery The name means "Tethys' lizard of Nopcsa", a reference to the Greek godd ...

'', they were poor swimmers compared to many of their contemporaries. Lacking swimming capabilities are quite surprising considering that most other small mosasaurs were adapted to be fast and nimble. The description of '' Phosphorosaurus ponpetelegans'' revealed that '' Phosphorosaurus'' was highly specialized to compensate for its lack of hydrophalangy.Takuya Konishi, Michael W. Caldwell, Tomohiro Nishimura, Kazuhiko Sakurai & Kyo Tanoue (2015) A new halisaurine mosasaur (Squamata: Halisaurinae) from Japan: the first record in the western Pacific realm and the first documented insights into binocular vision in mosasaurs. Journal of Systematic Palaeontology (advance online publication) DOI:10.1080/14772019.2015.1113447 http://www.tandfonline.com/doi/full/10.1080/14772019.2015.1113447#abstract Despite their shortcomings, the Halisaurinae had diversified and reached a global distribution by the end of the Maastrichtian whilst other subfamilies, particularly the

Plioplatecarpinae Plioplatecarpinae is a subfamily of mosasaurs, a diverse group of Late Cretaceous marine squamates. Members of the subfamily are informally and collectively known as "plioplatecarpines" and have been recovered from all continents, though the occu ...

and

Tylosaurinae The Tylosaurinae are a subfamily of mosasaurs,Williston, S. W. 1897. Range and distribution of the mosasaurs with remarks on synonymy. ''Kansas University Quarterly'' 4(4):177-185. a diverse group of Late Cretaceous marine squamates. Members of t ...

, appear to have been in decline. There is also evidence supporting that they were accomplished intercontinental migrators, with a known large-scale migration event involving ''

'' during the Campanian from the

Western Interior Seaway The Western Interior Seaway (also called the Cretaceous Seaway, the Niobraran Sea, the North American Inland Sea, and the Western Interior Sea) was a large inland sea that split the continent of North America into two landmasses. The ancient sea ...

North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...

to what today is southern

Sweden Sweden, formally the Kingdom of Sweden,The United Nations Group of Experts on Geographical Names states that the country's formal name is the Kingdom of SwedenUNGEGN World Geographical Names, Sweden./ref> is a Nordic country located on ...

. Bardet et al. (2005, p. 464Bardet, N., Pereda Suberbiola, X., Iarochene, M., Bouya, B. & Amaghzaz, M. 2005. A new species of Halisaurus from the Late Cretaceous phosphates of Morocco, and the phylogenetical relationships of the Halisaurinae (Squamata: Mosasauridae). Zoological Journal of the Linnean Society, 143, 447–472.) diagnosed the Halsaurinae as all mosasaurs more closely related to ''

'' than ''

Moanasaurus ''Moanasaurus'' (From Māori ''moana'' "sea" and Greek ''sauros'' "lizard"; meaning "Sea Lizard") was a genus of mosasaur from the Late Cretaceous period. Its fossil remains have been discovered in the North Island of New Zealand. ''Moanasauru ...

''. Unambiguous character states were listed as follows: "

premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...

maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...

sutural contact vertical anteriorly, oblique at midpoint and horizontal posteriorly; contact plane between the parietal and the supratemporal oblique; preaxial ridge extending on two-thirds of the length of the radius;

tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...

and

fibula The fibula 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. Its upper extremity is ...

long and slender with slightly expanded extremities. Ambiguous characters include "

dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal c ...

median ridge borne on the anterior two-thirds of the

frontal Front may refer to: Arts, entertainment, and media Films * ''The Front'' (1943 film), a 1943 Soviet drama film * ''The Front'', 1976 film Music * The Front (band), an American rock band signed to Columbia Records and active in the 1980s and e ...

; frontal with

ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Th ...

boss; parietal foramen surrounded by a ventral boss; quadrate with large infrastapedial process; coalescent infra- and suprastapedial processes of quadrate; zygosphene- zygantrum complex absent; synapophyses of the

cervical vertebrae In tetrapods, cervical vertebrae (singular: 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 ...

extending ventrally to the ventral surface of the centrum; fused haemal spines."

Phylogeny

Designation of this subfamily followed many decades of confusion surrounding the

type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal f ...

, ''Halisaurus,'' which had long been seen as (and remains) an important taxon in studies of mosasaur

phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

. The confusion was especially centered on ''H. sternbergii'', a species from 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 t ...

Alabama (We dare defend our rights) , anthem = "Alabama (state song), Alabama" , image_map = Alabama in United States.svg , seat = Montgomery, Alabama, Montgomery , LargestCity = Huntsville, Alabama, Huntsville , LargestCounty = Baldwin County, Al ...

and the Niobrara Chalk of

Kansas Kansas () is a state in the Midwestern United States. Its capital is Topeka, and its largest city is Wichita. Kansas is a landlocked state bordered by Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the ...

. Originally, the

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

had been referred to the mosasaurine ''

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 ...

'',Wiman CJ. 1920. Some reptiles from the Niobrara group in Kansas. ''Bulletin of the Geological Institute of Uppsala'' 18: 9-18 (9 figs., pls. II-IV). then to ''Halisaurus'' (Russell, 1967; p. 369 Russell DA. 1967. Systematics and morphology of American mosasaurs. ''Peabody Museum of Natural History, Yale University, Bulletin'' 23.), which was also considered a member of the

Mosasaurinae The Mosasaurinae are a subfamily of mosasaurs, a diverse group of Late Cretaceous marine squamates. Members of the subfamily are informally and collectively known as "mosasaurines" and their fossils have been recovered from every continent excep ...

at that time. Later workers Wright KR. 1988. A new specimen of ''Halisaurus platyspondylus'' (Squamata: Mosasauridae) from the Navesink Formation (Maastrichtian) of New Jersey. ''Journal of Vertebrate Paleontology'' 8 (Supplement 3): 29A-30A.Bell GL. Jr. 1997. A phylogenetic revision of North American and Adriatic Mosasauroidea. pp. 293-332 In: Callaway JM. and Nicholls EL, (eds.), ''Ancient Marine Reptiles'', Academic Press: 501 pp. questioned the

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

position and

monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...

of ''Halisaurus'', in part because of striking morphological differences between ''H. sternbergii'' and the other known species of the

taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

. Several discoveries throughout the 1980s and 1990s helped shed light on ''

'', with more complete specimens of the type species ''H. platyspondylus'' being discovered and '' Phosphorosaurus ortliebi'' being momentarily reassigned to the genus by Lingham-Soliar (1996). In 2005, the species ''Halisaurus sternbergii'' was reassigned to its own genus, ''

'', along with the description of the new species ''Halisaurus arambourgi'' by Nathalie Bardet and colleagues. With the description of ''Eonatator'' as a closely related genus to ''Halisaurus'', the two genera were grouped into the new subfamily Halisaurinae. Their study supported the model that the group was a sister-group to more advanced mosasaurs. Since the description of ''

'', two more species of ''

'', ''H. ortliebi'' and ''H. walkeri'' were recognised as separate genera; '' Phosphorosaurus'' and '' Pluridens'' respectively. The recentmost major phylogenetic analysis of mosasaurs, conducted by Tiago R. Simões and colleagues in May 2017, recovered ''Halisaurus'' and the rest of the Halisaurinae as a sister group to the

instead of the whole

Mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on th ...

idae. This would mean that the halisaurines are more closely related to the mosasaurines than the russellosaurines (genera such as ''

Tylosaurus ''Tylosaurus'' (from the ancient Greek (') 'protuberance, knob' + Greek (') 'lizard') is a genus of mosasaur, a large, predatory marine reptile closely related to modern monitor lizards and to snakes, from the Late Cretaceous. Description A ...

'' and '' Plesioplatecarpus'') are. Below is a

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

following an analysis by Takuya Konishi and colleagues (

2015 File:2015 Events Collage new.png, From top left, clockwise: Civil service in remembrance of November 2015 Paris attacks; Germanwings Flight 9525 was purposely crashed into the French Alps; the rubble of residences in Kathmandu following the Apri ...

) done during the description of '' Phosphorosaurus ponpetelegans'', which showcases the internal relationships within the Halisaurinae. The analysis excluded the dubious '' Halisaurus onchognathus'' and the genus '' Pluridens''.

Species and taxonomy

Halisaurinae *''

'' **''H. platyspondylus'' **''H. arambourgi'' **''H. onchognathus'' ('' nomen dubium''; holotype destroyed during

World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposin ...

) *'' Phosphorosaurus'' **''P. ortliebi'' **''P. ponpetelegans'' *''

'' **''E. sternbergii'' **''E. coellensis'' *'' Pluridens'' **''P. walkeri'' **''P. calabaria'' **''P. serpentis''

References

Notes

External links