Pholidosaurus

{{Short description|Extinct genus of reptiles}} {{Automatic taxobox | fossil_range = Early Cretaceous, {{fossilrange|Berriasian|Albian}} | image = Pholidosaurus.jpg | image_caption = ''Pholidosaurus meyeri'' skull fossil at the Museum für Naturkunde, Berlin | image_upright = 1.15 | taxon = Pholidosaurus | authority = Meyer, 1841 | subdivision_ranks = Species | subdivision = * {{extinct}}''P. meyeri'' (Dunker, 1844) * {{extinct}}''P. purbeckensis''? Salisbury, 2002 * {{extinct}}''P. schaumburgensis'' Meyer, 1841 (type) | synonyms = *''Macrorhynchus'' Dunker, 1844 *''Petrosuchus laevidens'' Owen, 1878 (''vide'' Andrews, 1913) *''Steneosaurus purbeckensis''? Mansel-Pleydell, 1888 }}

'''''Pholidosaurus''''' is an extinct genus of neosuchian crocodylomorph. It is the type genus of the family Pholidosauridae. Fossils have been found in northwestern Germany. The genus is known to have existed during the Berriasian-Albian stages of the Early Cretaceous. Fossil material found from the Annero and Jydegård Formations in Skåne, Sweden and on the island of Bornholm, Denmark, have been referred to as a mesoeucrocodylian, and possibly represent the genus ''Pholidosaurus''.<ref name=S-WRL09>{{cite journal |doi=10.1016/j.cretres.2009.07.011 |last=Schwarz-Wings |first=D. |author2=Rees, J. |author3= Lindgren, J. |year=2009 |title=Lower Cretaceous Mesoeucrocodylians from Scandinavia (Denmark and Sweden) |journal=Cretaceous Research |volume=30 |issue=5 |pages=1345–1355|bibcode=2009CrRes..30.1345S }}</ref>

==Description== thumb|250px|left|Skull of the possible species ''Pholidosaurus purbeckensis'' (originally classed under the genus ''Petrosuchus'') An early description of the genus by Lydekker (1888) mentioned that the orbit is slightly smaller than the supratemporal fossa, the nasals reach the premaxillae, and the vomer appears on the palate.<ref name=LR88>{{cite book |last=Lydekker |first=R. |year=1888 |chapter=Crocodilia |title=Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History): The orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynochocephalia, and Proterosauria |publisher=Trustees of the British Museum |location=London |pages=42–130}}</ref> It is similar in appearance to and about as large as the modern gharial.

==Species== The type species of ''Pholidosaurus'' is ''P. schaumburgensis'', named in 1841 from the Wealden of Bückeburg, Germany.<ref name=LR88/><ref name=LR87>{{cite journal |doi=10.1017/S001675680019377X |last=Lydekker |first=R. |year=1887 |title=Note on the Hordwell and other crocodilians |journal=Geological Magazine |volume=4 |pages=307–312 |issue=7|bibcode=1887GeoM....4..307L |s2cid=140634029 |url=https://zenodo.org/record/2237814 }}</ref> ''P. schaumburgensis'' was named on the basis of a natural mold of part of a thorax discovered in around 1830 from the Berriasian Obernkirchen Sandstein.<ref name=MH41>{{cite journal |last=Meyer |first=H. |year=1841 |title=''Pholidosaurus schaumburgensis'' ein Saurus aus dem Sandstein der Wald-Formation Nord-Deutschlands |journal=Neues Jahrbuch für Mineralogie, Geologie und Paläontologie |volume=4 |pages=443–445}}</ref> This mould is known as IMGPGö 741-1. The individual that the mould belonged to is thought to have been around {{convert|25|cm|in}} in length.<ref name=SSW02>{{cite book |last=Salisbury |first=S. W. |year=2002 |chapter=Crocodilians from the Lower Cretaceous (Beriassian) Purbeck Limestone Group of Dorset, Southern England |title=Life and environments in Purbeck times |editor1=A. R. Milner |editor2=D. J. Batten |series=Special Papers in Palaeontology |volume=68 |publisher=Wiley-Blackwell |pages=121–144}}</ref>

''Macrorhynchus'' is a junior synonym of ''Pholidosaurus''.<ref name=MCC34>{{cite journal |doi=10.1086/624165 |last=Mook |first=C. C. |year=1934 |title=The evolution and classification of the Crocodilia |journal=The Journal of Geology |volume=42 |issue=3 |pages=295–304 |bibcode=1934JG.....42..295M|s2cid=128568052 }}</ref> It was named in 1843 from the same stratigraphic unit and region as ''P. schaumbergensis'', with the type species being ''M. meyeri''.<ref name=DW44>{{cite journal |last=Dunker |first=W. |year=1844 |title=Ueber den norddeutschen sogenannten Walderton und dessen Versteinerungen |journal=Programm der Casseler Gewerbeschule}}</ref> Because ''M. meyeri'' bears a strong resemblance to ''Pholidosaurus schaumburgensis'', it is now regarded as a species of ''Pholidosaurus''. It was reassigned to the genus ''Pholidosaurus'' in 1887 by Richard Lydekker because of this synonymy, and also because the name ''Macrorhynchus'' was preoccupied by a genus of fish named in 1880.<ref name=LR87/> ''P. meyeri'' differs from ''P. schaumburgensis'' in that the bar separating the supratemporal fenestrae is rounded, while in the type species it is angular.<ref name=LR88/>

==Misassigned species== ''Pholidosaurus decipiens'' was erected for a partial cranium, NHMUK 28432, that was originally assigned to the new genus and species ''Petrosuchus laevidens'' by Richard Owen in 1878.<ref name=OR78>{{cite journal |last=Owen |first=R. |year=1878 |title=Monograph on The Fossil Reptilia of the Wealden and Purbeck Formations, Supplement no. VII. Crocodilia (''Goniopholis'', ''Pterosuchus'', and ''Suchosaurus'') |journal= Palaeontographical Society Monograph |pages=1–15}}</ref> ''Petrosuchus laevidens'' was based on this cranium and a mandibular ramus called BMNH 41099, both of which were collected from Swanage, England. A later study in 1911 concluded that the material belonged to two different species; NHMUK 28432 was reassigned to ''Pholidosaurus'' and NHMUK 41099 was designated the lectotype of ''Petrosuchus laevidens''. The species name ''decipiens'' was coined in reference to Owen's oversight, and ''Petrosuchus'' is now considered a junior synonym of ''Goniopholis simus''.<ref name=SSW02/>

Another species from England, ''P. purbeckensis'', was originally described as a species of ''Steneosaurus'' in 1888.<ref name=M-P88>{{cite journal |last=Mansel-Pleydell |first=J. C. |author-link=John Clavell Mansel-Pleydell |year=1888 |title=Fossil reptiles of Dorset |journal=Proceedings of the Dorset Natural History and Antiquarian Field Club |volume=9 |pages=1–40}}</ref> The holotype is an almost complete cranium, referred to as DORCM G97, missing the anterior portion of the rostrum. The skull was found from either Swanage or the Isle of Purbeck (hence the species name), although the exact locality from which the skull originated is not specified by the author of the original description.<ref name=WS80>{{cite book |last=Woodward |first=A. S. |author2=Sherbron, C. D. |year=1980 |title=A catalogue of British Fossil Vertebrata |volume=8 volumes |location=London |pages=396pp}}</ref> This material was also once referred to ''Macrorhynchus''. The author of the 1888 description considered ''S. purbeckensis'' an intermediate form between ''Steneosaurus'' and ''Teleosaurus''.<ref name=M-P88/> However, in 2002, a new study showed that ''S. purbeckensis'' was conspecific with ''P. decipiens'', creating the new combination ''Pholidosaurus purbeckensis''.<ref name=SSW02/>

Another species of ''Pholidosaurus'', ''P. laevis'', was named in 1913 from Swanage, based on the partial cranium NHMUK R3414.<ref name=ACW13>{{cite journal |last=Andrews |first=C. W. |year=1913 |title=On the skull and part of a skeleton of a crocodile from the Middle Purbeck of Swanage, with the description of a species (''Pholidosaurus laevis''), and a note on the skull of ''Hylaeochampsa'' |journal=Annals and Magazine of Natural History |volume=8 |pages=485–494 |doi=10.1080/00222931308693345|url=https://zenodo.org/record/1430095 }}</ref> This has been considered a junior synonym of ''P. purbeckensis'' by both Salisbury ''et al.'' (1999) and Salisbury (2002).<ref name=SSW02/><ref name=SWPS99>{{cite journal |last=Salisbury |first=S. W. |author2=Willis, P. M. A. |author3=Peitz, S. |author4= Sander, P. M. |year=1999 |title=The crocodilian ''Goniopholis simus'' from the Lower Cretaceous of northwestern Germany |journal=Special Papers in Palaeontology |volume=60 |pages=121–148}}</ref>

In an SVPCA abstract, Smith ''et al.'' (2016) noted that ''Pholidosaurus purbeckensis'' is not congeneric with the type species, and instead is closely related to ''Fortignathus'' and members of Dyrosauridae.<ref>Thomas J. Smith, Lorna Steel, and Mark T. Young, 2016. Re-evaluation of ''Pholidosaurus purbeckensis'' (Crocodyliformes: Tethysuchia) from the Early Cretaceous of England, with implications for the evolution of Pholidosauridae and Dyrosauridae. ''Symposium of Vertebrate Palaeontology and Comparative Anatomy 2016 Abstract Book''. p. 29. {{cite web |url=http://svpca.org/years/2016_liverpool/Abstract%20Bookv3.pdf |title=Archived copy |access-date=2016-08-24 |url-status=dead |archive-url=https://web.archive.org/web/20160915041248/http://svpca.org/years/2016_liverpool/Abstract%20Bookv3.pdf |archive-date=2016-09-15 }}</ref>

==Classification== {|style="float:right; text-align:center;" |'''Phylogenetic position of ''Pholidosaurus''''' |- |{{Cladogram|title='''Sereno ''et al.'' 2001'''<ref name=SLSG01>{{cite journal |doi=10.1126/science.1066521 |last=Sereno |first=P. C. |author2=Larsson, H. C. E. |author3-link=Sidor, C. A |author3=Sidor, C. A. |author4= Gado, B. |year=2001 |title=The giant crocodyliform ''Sarcosuchus'' from the Cretaceous of Africa |journal=Science |volume=294 |issue=5546 |pages=1516–1519 |pmid=11679634|bibcode = 2001Sci...294.1516S |s2cid=22956704 |url=http://doc.rero.ch/record/16412/files/PAL_E3596.pdf }}</ref> |clades={{clade| style=font-size:80%;line-height:80% |label1=Neosuchia |1={{clade |1={{clade |1={{clade |1=''Sunosuchus'' |2=''Goniopholis''}} |2={{clade |1='''''Pholidosaurus''''' |2={{clade |1=''Dyrosaurus'' |2={{clade |1=''Terminonaris'' |2=''Sarcosuchus''}} }} }} }} |2={{clade |1=''Bernissartia'' |2=Crocodylia }} }} }} }} |- |{{Cladogram|title='''Pol 2003'''<ref name=PD03>{{cite journal |doi=10.1671/A1015-7 |last=Pol |first=D. |year=2003 |title=New remains of ''Sphagesaurus huenei'' (Crocodylomorpha: Mesoeucrocodylia) from the Late Cretaceous of Brazil |journal=Journal of Vertebrate Paleontology |volume=23 |issue=4 |pages=817–831|bibcode=2003JVPal..23..817P |s2cid=86592576 }}</ref> |clades={{clade| style=font-size:80%;line-height:80% |label1=Neosuchia |1={{clade |1={{clade |1=''Theriosuchus'' |2=''Alligatorium''}} |2={{clade |1={{clade |1=''Sokotosuchus'' |2=''Dyrosaurus''}} |2={{clade |1='''''Pholidosaurus''''' |2={{clade |1=''Pelagosaurus'' |2={{clade |1=Teleosauridae |2=Metriorhynchidae}} }} }} }} |3=''Eutretauranosuchus'' |4={{clade |1=''Goniopholis'' |2={{clade |1=''Bernissartia'' |2={{clade |1=''Hylaeochampsa'' |2=''Borealosuchus'' |3=''Gavialis'' |4=''Crocodylus'' |5=''Alligator'' }} }} }} }} }} }} |}

Richard Lydekker assigned ''Pholidosaurus'' to the family Goniopholididae in 1887 along with ''Hylaeochampsa'', ''Theriosuchus'', ''Goniopholis'', and ''Petrosuchus'' because the vertebrae are amphicoelus and the orbit communicates with the lateral temporal fossa.<ref name=LR87/> [[Image:Pholidosaurus tooth.jpg|thumb|left|Possible ''Pholidosaurus'' tooth (DK164) from the Jydegaard Formation at the Geological Museum in Copenhagen]] ''Pholidosaurus'' has often been grouped with other longirostrine, or long-snouted, crocodylomorphs, including dyrosaurids and thalattosuchians. Buckley and Brochu (1999) concluded that ''Pholidosaurus'', ''Sokotosuchus'', Dyrosauridae, and Thalattosuchia formed a longirostrine clade that was the sister taxon to Crocodylia.<ref name=BB99>{{cite journal |last=Buckley |first=G. A. |author2=Brochu, C. A. |year=1999 |title=An enigmatic new crocodile from the Upper Cretaceous of Madagascar |journal=Special Papers in Palaeontology |volume=60 |pages=149–175}}</ref> However, Thalattosuchia was traditionally considered a more basal clade of crocodylomorphs, being a more basal lineage of Mesoeucrocodylia than dyrosaurids or ''Pholidosaurus'', both of which were considered neosuchians.<ref name=Hetal08>{{cite journal |last=Hill |first=R. V. |author2=McCartney, J. A. |author3=Roberts, E. |author4=Bouaré, M |author5=Sissoko, F. |author6= O'Leary, M. A. |year=2008 |title=Dyrosaurid (Crocodyliformes: Mesoeucrocodylia) fossils from the Upper Cretaceous and Paleogene of Mali: implications for phylogeny and survivorship across the K/T Boundary |journal=American Museum Novitates |issue=3631 |pages=19pp |url=http://digitallibrary.amnh.org/dspace/bitstream/2246/5940/1/N3631.pdf}}</ref> The results of the phylogenetic analysis by Buckley and Brochu (1999) were attributed to the similarity in characters associated with snout elongation seen in these crocodylomorphs, even though these characters may have been independently derived in each group. More recent studies have revealed Thalattosuchia as a more basal clade when dyrosaurids are removed from the data set.<ref name=BBKP00>{{cite journal |doi=10.1038/35016061 |last=Buckley |first=G. A. |author2=Brochu, C. A. |author3=Krause, D. W. |author4= Pol, D. |year=2000 |title=A pug-nosed crocodyliform from the Late Cretaceous of Madagascar |journal=Nature |volume=405 |issue=6789 |pages=941–944 |pmid=10879533 |bibcode=2000Natur.405..941B|s2cid=4426288 }}</ref>

More recent studies show that ''Pholidosaurus'' is closely related to the Thalattosuchia, with both taxa closely related to a clade containing ''Terminonaris'' and the Dyrosauridae.<ref name=WRC01>{{cite journal |doi=10.1671/0272-4634(2001)021[0492:TACNMF]2.0.CO;2 |last=Wu |first=X.-C. |author2=Russell, A. P. |author3= Cumbaa, S. L. |year=2001 |title=''Terminonaris'' (Archosauria: Crocodyliformes): new material from Saskatchewan, Canada, and comments on its phylogenetic relationships |journal=Journal of Vertebrate Paleontology |volume=21 |pages=492–514 |issue=3|s2cid=86206630 }}</ref> In a phylogenetic analysis conducted by Sereno ''et al.'' (2001), ''Pholidosaurus'' was placed as a distant sister taxon to the other longirostrine crocodylomorphs, with ''Terminonaris'' and the newly named ''Sarcosuchus'' being closely related to one another and ''Dyrosaurus'' being the next closest taxon to the group.<ref name=SLSG01/> The later phylogenetic analysis of Brochu ''et al.'' (2002) again showed that ''Pholidosaurus'' was closely related to Thalattosuchia. In the study, both taxa formed a clade that was the sister taxon to a clade containing ''Sokotosuchus'' and Dyrosauridae.<ref name=Betal08>{{cite journal |doi=10.1666/0022-3360(2002)076<1060:ADCBFM>2.0.CO;2 |last=Brochu |first=C. A. |author2=Bouare, M. L. |author3=Sissoko, F. |author4=Roberts, E. M. |author5= O'Leary, M. A. |year=2002 |title=A dyrosaurid crocodyliform braincase from Mali |journal=Journal of Paleontology |volume=76 |pages=1060–1071 |issue=6|url=http://doc.rero.ch/record/14734/files/PAL_E1860.pdf }}</ref> Jouve ''et al.'' (2006) concluded that ''Pholidosaurus'' was closely related thalattosuchians were also included within the family, which would be considered paraphyletic without them. Jouve ''et al.'' (2006), like Buckley and Brochu (1999), attributed this result to phylogenetic problems that exist among longirostrine crocodylomorphs due to similarities in their morphology.<ref name=JIBA06>{{cite journal |doi=10.1111/j.1096-3642.2006.00241.x |last=Jouve |first=S. |author2=Iarochène, M. |author3=Bouya, B. |author4= Amaghzaz, M. |year=2006 |title=A new species of ''Dyrosaurus'' (Crocodylomorpha, Dyrosauridae) from the early Eocene of Morocco: phylogenetic implications |journal=Zoological Journal of the Linnean Society |volume=148 |issue=4 |pages= 603–656|doi-access=free }}</ref>

==References== {{Reflist}}

==External links== * [https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=36602 ''Pholidosaurus''] in the Paleobiology Database

Category:Early Cretaceous reptiles of Europe Category:Early Cretaceous crocodylomorphs of Europe Category:Tethysuchia Category:Prehistoric pseudosuchian genera