Halisaurinae

The Halisaurinae are a subfamily of mosasaurs, a diverse group of Late Cretaceous marine squamates. Members of the subfamily are informally and collectively known as "halisaurines" and have been recovered from North and South America, Europe, Asia and Africa,[1] indicating a more or less global distribution in the Late Cretaceous. Though all the genera included in the subfamily have at one point or another been included within Halisaurus, the genera currently seen as valid halisaurines are Eonatator, Halisaurus, Phosphorosaurus and Pluridens.

Halisaurinae
Temporal range: Late Cretaceous 86.3–66 Ma
Skeleton of Halisaurus arambourgi.
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Superfamily: †Mosasauroidea
Family: †Mosasauridae
Subfamily: †Halisaurinae
Bardet et al., 2005
Genera

Halisaurines were small to medium-sized mosasaurs averaging 4.5-6+ meters in length. Of all known hydropedal[note 1] mosasaurids, they were the least adapted to a marine lifestyle. The earliest known remains of halisaurines occur in rocks of Santonian age and the subfamily persists until the latest Maastrichtian. The etymology of this group derives from the genus Halisaurus (Greek halis = "sea" + Greek sauros = "lizard").

Description
Restoration of Halisaurus.

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 and Tethysaurus, 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.[2]

Despite their shortcomings, the Halisaurinae had diversified and reached a global distribution by the end of the Maastrichtian whilst other subfamilies, particularly the Plioplatecarpinae and Tylosaurinae, appear to have been in decline.[3] There is also evidence supporting that they were accomplished intercontinental migrators, with a known large-scale migration event involving Eonatator during the Campanian from the Western Interior Seaway of North America to what today is southern Sweden.[4]

Bardet et al. (2005, p. 464[5]) diagnosed the Halsaurinae as all mosasaurs more closely related to Halisaurus than Moanasaurus. Unambiguous character states were listed as follows: "premaxilla-maxilla 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 and fibula long and slender with slightly expanded extremities. Ambiguous characters include "dorsal median ridge borne on the anterior two-thirds of the frontal; frontal with ventral 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 extending ventrally to the ventral surface of the centrum; fused haemal spines."

Phylogeny
Speculative restoration of Pluridens based on Halisaurus.

Designation of this subfamily followed many decades of confusion surrounding the type genus, Halisaurus, which had long been seen as (and remains) an important taxon in studies of mosasaur phylogeny. The confusion was especially centered on H. sternbergii, a species from the Mooreville Chalk Formation of Alabama and the Niobrara Chalk of Kansas. Originally, the species had been referred to the mosasaurine Clidastes,[6] then to Halisaurus (Russell, 1967; p. 369 [7]), which was also considered a member of the Mosasaurinae at that time. Later workers [8][9] questioned the phylogenetic position and monophyly of Halisaurus, in part because of striking morphological differences between H. sternbergii and the other known species of the taxon.

Several discoveries throughout the 1980s and 1990s helped shed light on Halisaurus, 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).[10] In 2005, the species Halisaurus sternbergii was reassigned to its own genus, Eonatator, 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.[5] Since the description of Eonatator, two more species of Halisaurus, 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 Mosasaurinae instead of the whole Mosasauridae. This would mean that the halisaurines are more closely related to the mosasaurines than the russellosaurines (genera such as Tylosaurus and Plesioplatecarpus) are.[11]

Below is a cladogram following an analysis by Takuya Konishi and colleagues (2015) done during the description of Phosphorosaurus ponpetelegans, which showcases the internal relationships within the Halisaurinae.[2] The analysis excluded the dubious Halisaurus onchognathus and the genus Pluridens.

Halisaurinae
 

Halisaurus arambourgi

 
 
 

Halisaurus platyspondylus

 
 

Eonatator coellensis

 
 
 

Eonatator sternbergii

 
 
 

Phosphorosaurus ortliebi

 
 

Phosphorosaurus ponpetelegans

 
 
 
 
 

Species and taxonomy

Halisaurinae

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References
  1. "Fossilworks: Halisaurinae". fossilworks.org. Retrieved 2017-08-20.
  2. 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
  3. Polcyn, Michael J.; Jacobs, Louis L.; Araújo, Ricardo; Schulp, Anne S.; Mateus, Octávio (2014-04-15). "Physical drivers of mosasaur evolution". Palaeogeography, Palaeoclimatology, Palaeoecology. Physical drivers in the evolution of marine tetrapods. 400: 17–27. doi:10.1016/j.palaeo.2013.05.018.
  4. Lindgren, Johan; Siverson, Mikael (2005). "Halisaurus sternbergi, a Small Mosasaur with an Intercontinental Distribution". Journal of Paleontology. 79 (4): 763–773. doi:10.1666/0022-3360(2005)079[0763:HSASMW]2.0.CO;2. JSTOR 4095048.
  5. Bardet, 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.
  6. 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).
  7. Russell DA. 1967. Systematics and morphology of American mosasaurs. Peabody Museum of Natural History, Yale University, Bulletin 23.
  8. 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.
  9. 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.
  10. Lingham-Soliar, T. 1996. The first description of Halisaurus (Reptilia Mosasauridae) from Europe, from the Upper Cretaceous of Belgium. Bulletin de l'Institut royal des Sciences naturelles de Belgique, Sciences de la Terre, 66, 129–136.
  11. Simões, Tiago R.; Vernygora, Oksana; Paparella, Ilaria; Jimenez-Huidobro, Paulina; Caldwell, Michael W. (2017-05-03). "Mosasauroid phylogeny under multiple phylogenetic methods provides new insights on the evolution of aquatic adaptations in the group". PLOS ONE. 12 (5): e0176773. doi:10.1371/journal.pone.0176773. ISSN 1932-6203. PMC 5415187. PMID 28467456.

Notes
  1. In mosasaurs, the terms "hydropedal" and "plesiopedal" refers to varying limb conditions and varying degrees of adaptations for marine life. Plesiopedal mosasaurs, such as Dallasaurus or Tethysaurus were primitive and largely coastal, while later hydropedal mosasaurs were streamlined and well-adapted to marine life.

Further reading

Bardet N, Suberbiola XP. 2001. The basal mosasaurid Halisaurus sternbergii from the Late Cretaceous of Kansas (North America): a review of the Uppsala type specimen. Comptes Rendus de l'Académie des Sciences, Série IIA 332: 395-402.

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